Biology MGMT of Rice Insects PDF

BIOLOGY AND
MANAGEMENT
OF RICE INSECTS
BlOLOGY AND
OF RICE
Edited by
E.A. HEINRICHS
PUBLISHING FOR ONE WORLD

WILEY EASTERN LIMITED
NEW AGE INTERNATIONAL LIMITED
New Delhi • Bangalore • Bombay • Calcutta • Guwahati
Hyderabad • Lucknow • Madras • Pune • London
MANAGEMENT
INSECTS
With contributions by
R.M. Aguda • A.T. Barrion • M. Bharathi • S. Chelliah • D. Dale
K.O. Gallagher • E.A. Heinrichs • K. Kiritani • J.A. Litsinger
M.E. Loevinsohn • K. Naba • P.A.C. Ooi • O. Parada
D.W. Roberts • M.C. Rombach • B.M. Shepard • C.M. Smith • G. Weber
IRRI
INTERNATIONAL RICE RESEARCH INSTITUTE
Copyright © 1994 Wiley Eastern Limited • New Age International Limited
WILEY EASTERN LIMITED • NEW AGE INTERNATIONAL LIMITED

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FOREWORD
INSECT PESTS ARE severe constraints to rice production throughout the world. Although rice insect
outbreaks are recorded in antiquity, evidence indicates that the severity and frequency of attack has
increased in the last several decades. The increase appears to have accompanied crop intensifica-
tion. Thus, for high yielding, green revolution varieties to achieve near their yield potential, various
constraints, including those imposed by insects, must be mitigated.
Integrated pest management has been accepted as the rationale approach to the regulation of
rice insect populations. Integrated pest management is an economically attractive and ecologically
and sociologically acceptable means of managing rice insect populations. To most effectively
implement integrated pest management programs requires first of all a proper identification of the
pest complex and an understanding of the biological and ecological factors that regulate pest
populations. With these building blocks as a firm foundation for properly characterizing and
monitoring insect populations, various pest control tactics have been developed and integrated into
rice insect management programs.
Success in the development and implementation of effective rice insect management programs
has been limited in scope and has yet to be even partially achieved in many of the rice growing
regions of the world. Success cannot be achieved without a development of the required skills. The
authors have written this book to provide the basic information needed to develop those skills.
There has been an acute need for a book that draws together the fundamentals of rice insect
taxonomy, biology, and ecology, and the various control tactics, and explains how these are in-
tegrated into a successful rice insect management program. The authors have drawn upon their
extensive experience in rice entomology and have succeeded remarkably well in filling that need
through the completion of this book. Its publication is evidence of our desire to promote and
accelerate the development of integrated pest management systems for rice throughout the rice
growing world.
KLAUS LAMPE
Director General
International Rice Research Institute
PREFACE
INSECTS ARE A MAJOR constraint in the production of rice throughout the world. They are especially
severe in tropical Asia and are increasing in importance in Africa and South America. With the
continued increases in the human population and losses of arable land, there is an ever increasing
need to increase rice production per unit of land through the development of rice production
strategies that are sustainable and are economically, environmentally, and socially acceptable. Rice
entomologists are playing a major role in the development of sound rice production strategies and
will continue to do so in the challenging years ahead.
Development of rice insect management strategies is promoted by the availability of literature
upon which to draw. During the last three decades there has been a virtual explosion of rice
entomology research papers scattered in diverse publications throughout the world, but there has
not been a comprehensive treatment of rice entomology under one cover. The few books on rice
entomology are out of date, very limited in scope, and are not available to most rice scientists.
During my tenure at the International Rice Research Institute (IRRI) (1975-1985), I often felt the
need for a reference book on rice insects. This book was produced to provide students, rice
entomologists, and scientists in other rice disciplines with a comprehensive reference on rice
entomology that contains the information needed to understand the principles of rice entomology
and to develop research and insect management programs.
The fundamentals of rice entomology, the biology and ecology of rice insects, the various tactics
for rice control, and the implementation of rice insect management systems are covered in this
book. Chapter 1 discusses the global importance of rice as a major food crop and the role of insects
in causing plant damage. Chapter 2 consists of a taxonomic key for the identification of rice insect
pest species and their parasites and predators. This is the first taxonomic key published covering
essentially all of the world’s important rice insect pests and their natural enemies. Chapter 3
explains the biology, distribution, life cycle, ecology and damage caused by the major rice insect
pests throughout the rice growing regions of the world. Chapter 4 is a treatment of how ecological
factors determine the presence and abundance of rice insect pests in different rice cropping
environments. Chapters 5-9 discuss the status of the various rice insect control tactics: host plant
resistance; cultural, mechanical and physical controls; predators and parasitoids; pathogens; and
insecticides, respectively. Chapter 10 covers the integration of the various insect control tactics.
Chapter 11 is a case study of the development of a management strategy for the brown planthopper,
Nilaparvata lugens (Stål), and Chapters 12 and 13 explain the development and implementation of
rice insect management systems designed for Japan and Latin America, respectively.
viii PREFACE
Effective rice insect management programs are interdisciplinary in nature and involve not only
entomologists with diverse expertise, but also rice breeders, plant physiologists, plant biochemists,
soil scientists, economists, weed scientists, plant pathologists, and sociologists. All of these dis-
ciplines will find this book to be a valuable resource for their research program. This book will also
serve as a supplement to texts in entomology and other agricultural courses in colleges and
universities.
A great deal of effort has gone into the production of this book. I thank the authors for the
excellence of the chapters that they contributed and extend my sincere appreciation to Jan Ray for
her assistance in the preparation of the manuscript, and to Kristin Heinrichs and Wendy Heinrichs
for preparing the subject and species indices. To Danny Amalin, Entomology Division, IRRI, I
express my appreciation for the superb quality of his approximately 2,000 drawings of insects
included in Chapters 2 and 3. Last, but not least, I owe a debt of gratitude to Dr. N.C. Brady,
Director-General, IRRI (1973-1983), and Dr. M.D. Pathak, Director, Research Coordination and
Training, IRRI (1962-1989) for their inspiration and encouragement and for providing me the
opportunity to conduct research on rice, the most important food crop of the developing world.
Baton Rouge, Louisiana E.A. HEINRICHS

CONTENTS
Foreword .......................................................................... v
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
I. FUNDAMENTALS
1. Rice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
– E.A. Heinrichs
2. Taxonomy of Rice Insect Pests and Their Arthropod Parasites and Predators . . . . . . . . . 13
– Alberto T. Barrion and James A. Litsinger
II. BIOLOGY AND ECOLOGY

3. Insect Pests of the Rice Plant — Their Biology and Ecology . . . . . . . . . . . . . . . . . . . . . . . . 363
– D. Dale
4. Rice Pests and Agricultural Environments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
– Michael E. Loevinsohn
III. CONTROL TACTICS AND STRATEGIES

5. Host Plant Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 517
– E.A. Heinrichs
6. Cultural, Mechanical, and Physical Control of Rice Insects . . . . . . . . . . . . . . . . . . . . . . . . 549
– James A. Litsinger
7. Predators and Parasitoids of Rice Insect Pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 585
– P.A.C. Ooi and B.M. Shepard
8. Pathogens of Rice Insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 613
– Michiel C. Rombach, D. W. Roberts, and Remedios M. Aguda
9. Insecticide Management in Rice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
– S. Chelliah and M. Bharathi
X CONTENTS
10. Integration of Rice Insect Control Strategies and Tactics . . . . . . . . . . . . . . . . . . . . . . . . . . . 681

– C. Michael Smith
11. The Brown Planthopper: Promises, Problems, and Prospects . . . . . . . . . . . . . . . . . . . . . . . 693
– Michiel C. Rombach and Kevin D. Gallagher
IV. IMPLEMENTATION OF RICE IPM SYSTEMS

12. Development and Implementation of Rice IPM in Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . 713
– Keizi Kiritani and Kunihiko Naba
13. Development of an Integrated Pest Management System for Rice in
Latin America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 733
– G. Weber and O. Parada
Subject Index ..................................................................... 749
Species Index .................................................................... 763
CONTRIBUTORS
Remedios M. Aguda Michael E. Loevinsohn

Entomology Division International Development Research Centre
International Rice Research Institute Ottawa, Canada
Manila, Philippines Kunihiko Naba
Albert T. Barrion Division of Entomology
Entomology Division Hiroshima Prefectural Agricultural Experiment
International Rice Research Institute Station
Manila, Philippines Hagashihiroshima, Hiroshima, Japan
M. Bharathi P.A.C. Ooi
Tamil Nadu Rice Research Institute CAB International Institute of Biological
Aduthurai, Tamil Nadu, India Control
S. Chelliah (IBC) Malaysian Station, Batu Tiga
Tamil Nadu Rice Research Institute Selangor, Malaysia
Aduthurai, Tamil Nadu, India
O. Parada
D. Dale Federacion Nacional de Arroceros
Department of Entomology Bogota, Colombia
Kerala Agricultural University
Kerala, India D.W. Roberts
Insect Pathology Research Center
Kevin O. Gallagher Boyce Thompson Institute for Plant Research
Department of Entomology at Cornell University
University of California at Berkeley
Ithaca, New York
Albany, California
Michiel C. Rombach
EA. Heinrichs
Insect Pathology Resource Center
Department of Entomology
Boyce Thompson Institute for Plant Research
Louisiana State University Agricultural Center
at Cornell University
Baton Rouge, Louisiana
Keizi Kiritani C. Michael Smith
Division of Entomology Department of Plant, Soil and Entomological
National Institute of Agro-Environmental Science
Sciences, Yatabe, Tsukuba, Japan University of Idaho
Moscow, Idaho
James A. Litsinger
Entomology Division G. Weber
International Rice Research Institute Centro Internacional de Agricultura Tropical
Manila, Philippines Cali, Colombia
I. FUNDAMENTALS
1
Rice
E.A. HEINRICHS
Baton Rouge, Louisiana, USA
"So dependent upon rice are the Asian countries that throughout history a failure of that crop
has caused widespread famine and death."
—Robert F. Chandler, Jr., 1979
1. IMPORTANCE OF RICE
RICE IS THE PRIMARY staple for more than two billion people in Asia and for hundreds of millions
of people in Africa and Latin American (IRRI 1985). Among the major food crops it is the only one
that is almost exclusively a human food. It constitutes half of the diet of the more than 1.6 billion
people and 400 million rely on rice for a fourth to a half of their diet (Swaminathan 1984). Rice
provides about 80% of the calories for two billion Asians and one-third of the caloric intake of one
billion persons in Africa and Latin America (Chang 1984). Consumption per capita varies from 186
kg/year in Burma to a low of 4 kg/year in USA (IRRI 1988).
2. ORIGIN OF RICE
Rice an annual grass (Gramineae) is cultivated from 53° N to 40° S latitude. Rice belongs to the
genus Oryza which includes twenty wild species and two cultigens, O. sativa and O. glaberrima
(Table 1) (Chang 1985). The origin of rice is lost in antiquity. Chang (1976) (Fig. 1) has postulated
that the original habitat was the supercontinent of Gondwanaland. When Gondwanaland broke up
Oryza species drifted to what became Africa, Antarctica, Australia, Malagasy, South America, and
Southeast Asia.
4 RICE
TABLE 1
Species of Oryza, chromosome numbers, and geographical distributions
(from Chang 1985)
Species name x =12 Distribution

(synonym) 2n =
O. alta Swallen 48 Central and South America
O. australiensis Domin 24 Australia
O. barthii A. Chev. 24 West Africa
(O. breviligulata)
O. brachyantha A. Chev. & 24 West and central Africa
Roehr.
O. eichingeri A. Peter 24,48 East and central Africa
O. glaberrima Steud. 24 West Africa
O. gandiglumis (Doell) Prod. 48 South America
O. granulata Nees & Arn. 24 South and Southeast Asia
ex Hook f.
O. glumaepatula Steud. 24 South America and West Indies
(O.perennis subsp. cubensis)
O. latifolia Desv. 48 Central and South America
O. longiglumis Jansen 48 New Guinea
O. longistaminata A. Chev. & 24 Africa
Roehr. (O. barthii)
O. meridionalis N.Q. Ng 24 Australia
O. meyeriano (Zoll. & Morrill) 24 Southeast Asia, southern China
ex Steud.) Baill.
O. minuta J.S. Presl ex C.B. Presl 48 Southeast Asia
O. nivara Sharma & Shastry 24 South and Southeast Asia and Southern
(O. fatua, O. sativa f. China
spontanea)
O. officinalis Wall. ex Watt 24 South and Southeast Asia, southern China,
New Guinea
O. punctata Kotschy ex. Steud. 48,24 Africa
O. rideyi Hook f. 48 Southeast Asia
O. rufipogon W. Griffith 24 South and Southeast Asia and southern
(O. perennis, O. fatua, O. China
perennis subsp. balunga)
O. sativa L. 24 Asia
O. schlechteri Pilger – New Guinea
Domestication of O. sativa occurred some 10,000 years ago in the river valleys of South and
Southeast Asia and China (De Datta 1981). Rice specimens have been found in China that date
from the third millenium B.C. (Grist 1975). Earliest historical references are found in Chinese
writings of about 3,000 B.C. when it was stated that of the five principal food plants in the country
rice was the most important.
Rice was introduced into Kyushu, Japan from China in about 100 B.C. and from there it spread
reaching Hokkaido in the extreme north only in the eighteenth century. The Portuguese introduced
HEINRICHS 5
Fig. 1. Evolutionary pathway of two cultigens. Taxa boxed by solid lines are wild perennials. Taxa
boxed by broken lines are annuals. Arrow with solid line indicates direct descent. Arrow with broken
line indicates indirect descent. Double arrows indicate introgessive hybridization (from Chang 1985).
rice into Brazil and the Spaniards introduced rice into Central America and parts of South America.
The crop was brought into Italy in the fifteenth century.
Rice cultivation in the USA dates from about 1646 when it was introduced into the James River
region of Virginia and in 1685 when it was first grown in the colony of South Carolina. In 1718 it was
introduced into Louisiana but did not assume importance until 1887.
3. RICE CLASSIFICATION
In Asia O. sativa differentiated into three subspecies based on geographic conditions; indica,
javanica, and japonica (sinica). Indica refers to the tropical and subtropical varieties grown
throughout South and Southeast Asia and southern China whereas japonica refers to the short and
roundish grained varieties of the temperate zones of Japan, China, and Korea. Javanica designates
the Indonesian bulu (awned) and gundil (awnless) rices with long panicles and bold grains growing
alongside of indicas in Indonesia. A further classification for rice that refers to its habitat in terms
of soil and water, is upland, rainfed shallow, deep water, floating, and irrigated. Rice area of the
five types in India and China is indicated in Fig. 2. Most of the rice area in China is irrigated
whereas in India less than half is irrigated (IRRI 1985). In Latin America and Africa most of the
rice is upland whereas in the U.S.A. all rice is grown under irrigated conditions.
6 RICE
Fig. 2. Percentage area of the various types of rice culture in India and China
(from IRRI 1985).
Rice is adaptable to a wide range of environmental conditions because it has an efficient system
of air passage from shoot to root. This system allows rice to grow in waterlogged soils. Because of
the air passage system roots respire aerobically and utilize carbohydrates to produce energy even
when they are growing in an anaerobic environment.
In much of the world the rice crop is grown on soils with 5 to 10 cm of standing water. Rice also
grows well on dry upland soils where it is cultivated like wheat. Rice can thus be grown in mixed
and multiple cropping patterns with other crops including wheat, maize, and soybeans
(Swaminathan 1984). Floating rice grows in water that is one to five m deep. It can elongate at the
rate of 25 cm per day in rapidly rising water.
The rice plant passes through 10 stages during its growth cycle: (1) germination and emergence,
(2) seedling, (3) tillering, (4) stem elongation, (5) panicle initiation, (6) panicle development,
(7) flowering, (8) milk grain, (9) dough grain, and (10) mature grain stage (Reissig et al. 1986).
Traditional varieties require about 150 days of growth to reach the mature grain stage whereas the
HEINRICHS 7
Fig. 3. Portions of the rice plant attacked by insects (from Reissig et al. 1986).
modern, high yielding very early maturing varieties can be harvested in as few as 90 days after
sowing.
4. DAMAGE CAUSED BY RICE INSECT PESTS
4.1 YIELD LOSSES

Insect pests attack all portions of the rice plant and all stages of plant growth. Grain insects remove
milk from developing grains, defoliators feed on the leaves, sucking insects feed on the leaves and
stem, stem borers feed within the stem, and some species feed on the roots (Fig. 3).
8 RICE
Insects are important constraints to rice production and occur in all rice growing environments
(see Chapter 4). Insects reduce yields substantially, especially in tropical Asia. Cramer (1967)
estimated the rice yield losses caused by insects by reviewing the literature up to 1966. Because of
the inadequacy of much of the data he adopted a conservative approach. Yet he estimated losses
ranging from 31.5% in Asia (excluding The People’s Republic of China) to 2% in Europe and
Oceania (Table 2). The severity of insect damage in Asia is further indicated by the results of 117
experiments conducted on the IRRI (International Rice Research Institute) farm in Los Baños,
Philippines. The average yield of insecticide-protected plots was 4.9 t/ha whereas yields of plots
receiving no insecticide was 3.0 t/ha, a yield loss of about 40%. However, losses on experiment
station plots are usually greater than in farmer’s fields. Thus a further series of experiments
coordinated by IRRI was conducted in farmers’ fields in six Asian countries. The increase in yields
of fields receiving fertilizer, and insect and weed control was 0.9 t/ha in the wet season and 1.7 t/ha
in the dry season (Herdt 1979); 45% and 35% of those increases, respectively, were due to insect
control (Fig. 4).
TABLE 2
Rice yield losses caused by insect pests on a world basis (from Cramer 1967)
Region yield loss

(%)
Asia (excluding People’s Republic of China) 31.5
People’s Republic of China 15.0
Africa 14.4
South America 3.5
North and Central America 3.4
Europe 2.0
Oceania 2.0
4.2 FACTORS AFFECTING EXTENT OF INSECT-CAUSED YIELD LOSSES

Rice insects are as old as rice itself. Historical records from Korea and Japan report on the history
of rice insect pests during the last two millenia. In 701 insect outbreaks were reported in western
Japan. In 875 a migratory locust outbreak occurred in the Ise District of Japan and in 1071 an
Fig. 4. Factors affecting differences in yields of farmers’ fields following farmers’ practices and
improved production practices in six Asian countries 1974-1977 (from Herdt 1979).
HEINRICHS 9
armyworm outbreak occurred near Kyoto. A brown planthopper outbreak in 1733 was reported as
one of the most damaging insect outbreaks in the history of rice production in Japan. About 2.6
million persons were affected and 12,000 died from Hunger (Okutani 1980).
Although rice insect outbreaks have been recorded over the last 1,300 years, they have become
much more frequent and the insect pest complex has changed in the last three decades. Certain
insects such as the leaf- and planthoppers have increased in severity, whereas others such as certain
stem borer species have declined in importance.
Crop intensification has been suggested as a reason for the changes in the insect pest fauna in
rice (Loevinsohn 1985). Loevinsohn defines intensification as “an increase in resources devoted to
rice cultivation.” Intensification involves (1) an increase in the number of crops grown per year,
(2) an increase in the use of agricultural chemicals; fertilizer and pesticides, (3) planting of varieties
responsive to fertilizers and pesticides, and (4) increased plant densities, Loevinsohn provides
evidence that ecologically specialized species have been favored by crop intensification.
Changes in rice cultural practices have accompanied the widespread adoption of modern
varieties. The photoperiod insensitivity and reduced growth duration of the modern varieties have
made it possible to grow two and even three crops per year, where water, and temperatures are
adequate. The spread of the dry season crop has caused shifts in the composition of pest fauna
(Loevinsohn et al. 1988). Although most of the insects in the tropics have increased in severity,
some species such as the white stem borer Scirpophaga innotata (Walker), have decreased in
importance because its larvae aestivate in stubble and are destroyed by the dry season land
preparation. Species dependent on standing water, such as the whorl maggot Hydrellia philippina
Ferino, and the rice caseworm Nymphula depunctalis (Guenee), have become more abundant.
Fig. 5. Annual light trap collections indicate the replacement of the dark-headed stem borer Chilo
polychrysus with the yellow stem borer Scirpophaga incertulas with the expansion of double rice
cropping in Titi Serong, Malaysia, 1964-1975 (from Lim and Heong 1977).
10 RICE
The increase in the area planted to dry season rice as a result of expanded irrigation has
reduced the area planted to alternate crops and the cover of wild grasses in fields formerly left
fallow during the dry season. This change has favored specialist insect pests with monophagous
feeding habits such as the yellow stem borer Scirpophaga incertulas (Walker) while the population
of the polyphagous dark-headed borer Chilo polychrysus (Meyrick) has decreased (Lim and Heong
1977) (Fig. 5).
The most dramatic effects of modern cultural practices on abundance of rice insects specialists
is seen in the plant sucking Homoptera; the nearly monophagous brown planthopper Nilaparvata
Iugens (Stal), and the most monophagous species of the tropical leafhopper complex, Nephotettix
virescens (Distant). The brown planthopper rose from the status of a secondary pest to a major yield
constraint beginning in the 1960s (Heinrichs and Mochida 1984). In Indonesia losses in 1976-1977
were estimated at 365,000 metric tons of milled rice; equivalent to the annual consumption of 3
million people (Dyck and Thomas 1979). Of the various changes in production practices the
increased use of resurgence-inducing insecticides has been considered as the major cause of
intensified brown planthopper problems in Asia (see Chapters 9 and 11).
5. DEVELOPMENT OF INTEGRATED PEST

MANAGEMENT STRATEGIES
As rice scientists and farmers have gained experience in the cultivation of the modern varieties and
the agronomic practices that have accompanied the ‘green revolution’ they have become acutely
aware of the need to develop more effective and economical pest management strategies. This has
resulted in a shift from a primarily unilateral approach of insect control with a strong reliance on
insecticides to a multilateral approach involving a mix of control approaches.
Alternative pest control tactics have been developed and integrated into control programs.
This approach known as integrated pest management (IPM) in the simplest terms is referred to as
“a broad ecological attack combining several tactics for the economic control and management of
pest populations” (Smith and Calvert 1978). Integrated control practices rely on combinations of
biological, chemical, and cultural control methods and insect resistant rice varieties. Integrated
management systems manipulate multiple plant protection tactics in an effective and coordinated
way.
IPM programs are having a significant impact on minimizing the adverse effects of insecticides,
and in increasing the profitability of rice production. It has been estimated that the cost savings
from research leading to increased insect pest management efficiency on rice in South and
Southeast Asia will be $973 million in insecticide saved by the year 2,000 (Flinn 1984).
Chapters in this book discuss the basic information required to develop sound rice insect pest
management systems; an understanding of the taxonomy, biology and ecology of rice insect pests.
These chapters are followed by a section dealing with the various control tactics including host plant
resistance, cultural, mechanical, and physical controls, biological control agents, and insecticides.
Chapters on the implementation of rice insect management systems discuss the integration of the
HEINRICHS 11
~ ~~~ ~
various control tactics and the deployment of insect pest management systems in Japan and Latin
America.
During a visit to IRRI in the Philippines in the early 1960s Norman Cousins, then editor of the
Saturday Review, wrote that “any genuine improvement in the human condition on this planet must
be concerned with rice.” Entomologists throughout the rice growing regions of the world are
striving diligently to do their part in improving the quality of life by developing and implementing
strategies to more effectively, safely, and economically manage rice insect pests.
REFERENCES
Chang, T.T. 1976. The origin, evolution. cultivation dissemination and diversification of Asian and African rices.
Euphytica 25 : 425-441.
Chang, T.T. 1984. Conservation of rice genetic resources: luxury or necessity? Science 224: 251-256.
Chang, T.T. 1985. Crop history and genetic conservation: rice – A case study. Iowa State J. Res. 59: 425-455.
Cramer, H.H. 1967. Plant Protection and World Crop Protection. Pflanzenschutz Nachr. 20 1-524.
De Datta, S.K. 1981. Principles and Practices of Rice Production, John Wiley, New York.
Dyck, V.A., and B. Thomas. 1979. The brown planthopper problem: In Brown Planthopper: Threat to Rice Production in
Asia. International Rice Research Institute, Los Baños. Philippines, pp. 3-17.
Flinn, J.C. 1984. Report to the CGIAR on the future input of IRRI (mimeo). International Rice Research Institute, Los
Baños, Philippines.
Grist, D.H. 1975. Rice. Fifth ed., Longman, London.
Heinrichs, E.A., and O. Mochida. 1984. From secondary lo major pest status: The case of insecticide-induced rice brown
planthopper, Nilaparvata lugens, resurgence Pror Ecol. 7: 201-218.
Herdt, R.W. 1979. An overview of the constraints project results In Farm Level Constraints to High Rice Yields in
1974-1977. International Rice Research Institute, Los Baños, Philippines. pp. 395-411.
International Rice Research Institute (IRRI). 1985. International Rice Research 25 Years of Partnership. Los Baños,
Philippines.
International Rice Research Institute (IRRI). 1988. IRRI Rice Facts 1988. Los Baños, Philippines.
Lim, D.S., and K.L. Heong. 1977. Habitat modification for regulating pest population of rice in Malaysia. Malaysia
Agricultural Research and Development Institute Report No. 50.
Loevinsohn, M.A., J.A. Litsinger, and E.A. Heinrichs. 1988. Rice insect pests and agricultural change. In Harris, M.K.,
and C.E. Rogers (eds.).The Entomology of Indigenous and Naturalized Systems in Agriculture.Westview Press,
Boulder, CO., USA.
Loevinsohn, M.E. 1985. Agricultural intensification and rice pest ecology: Lessons and implications. A paper presented
at the International Rice Research Conference 1-5 June, 1985. International Rice Research Institute. Los Baños,
Philippines.
Okutani, T. 1980. Principles of rice insect pests control. In Rice Protection in Japan. Part 2 Entomologv. Japan
International Cooperation Agency, Hyogo, Japan.
Reissig, W.H., E.A. Heinrichs, J.A. Litsinger, K. Moody, L. Fiedler, T.W. Mew, and A.T. Barrion. 1986. Illustrated Guide
to Integrated Pest Management in Rice in Tropical Asia. International Rice Research Institute. Los Baños. Philippines.
Smith, R.F., and D.J. Calvert. 1978. Insect pest losses and the dimensions of the world food problem. In Pimentel, D.
(ed.). World Food, Pest Loses. and the Environment. Westview Press, Boulder, Co, USA.
Swaminathan, MS. 1984. Rice. Sci. Am. 250(1): 81-93.
2
Taxonomy of Rice Insect Pests and their

Arthropod Parasites and Predators
ALBERTO T. BARRION AND JAMES A. LITSINGER

Division of Entomology
P.O. Box 933, Manila, Philippines 1099
1. INTRODUCTION
RICE IS GROWN WORLDWIDE in over 124 million hectares under diverse cultural conditions and over
a wide geographical range. The need for higher levels of production has caused expanded areas to
be planted and more crops per year to be harvested. The greater apparency of rice in agricultural
ecosystems will tend to increase species richness of both pests and natural enemies. Grist and Lever
(1969) were the last to catalogue the insect pests of rice worldwide.
This chapter provides a practical and comprehensive classification of insects and spiders
associated with rice worldwide. It includes both rice pests and natural enemies (predators and
parasites) of rice pests. The keys are based on diagnostic characters for each species and have
necessarily had to involve morphological features relevant to each taxonomic group. The richly
illustrated keys can be used by individuals with minimal experience in insect morphology and
taxonomy. We have maintained technical accuracy by using precise morphological and taxonomic
terminology but also give non-technical synonyms. With the exception of leafhoppers and plant-
hoppers the keys are based primarily on adults which have a greater array of morphological
features.
This chapter is more complete for Asia than Africa or Latin America and South America
because the keys were based on specimens in our reference collection and literature available to us.
However, it is hoped that this will contribute to a better understanding of the rice ecosystem and
will serve as a taxonomic guide for further studies in rice entomology.
14 TAXONOMY OF RICE INSECT PESTS ETC.
2. KEY TO INSECTS AND SPIDERS

1 With 3 distinct body regions – head, thorax, and segmented abdomen; 6-legged;
winged or wingless (insects) (Fig.1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1' With only 2 body regions – cephalothorax (fused head and thorax) and unsegmented
abdomen; 8-legged; wingless (spiders) (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 729
2(1) Wings present (with few veins or well developed) ............................ 3

2' Wingless ............................................................. 713
3(2) With only 1 pair of wings ................................................. 4

3' With 2 pairs of wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
4(3) Forewings reduced to a pseudohaltere, leather-like and without venation; functional

hindwings markedly expanded; head with large berry-like eyes; antennae 4- or 7-seg-
mented; pro- and mesothorax much reduced, metathorax very large; trochanters absent;
twisted winged parasites, males (STREPSIPTERA) .......................... 5
4' Forewings developed, membranous and with venation; hindwings reduced to club-like
halteres; pro- and metathorax greatly reduced; trochanters present; fly pests, parasites,
and predators (DIPTERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5(4) Tarsi 2-segmented; antenna 4-segmented, 3rd segment laterally flabellate and originates
at the base of the segment; parasites of delphacids and fulgorids [Eienchidae] . . . . 6
5' Tarsi 3-segmented; antenna 7-segmented, the 3rd to 5th prolong laterally, and 7th
segment elongate; parasites of cicadellids [Halictophagidae] . . . . . . . . . . . . . . . . . . . 7
6(5) Postlumbium (Fig. 3) prominently large and semicircular; postscutellum large, about 0.7
of entire metathorax; vein A straight to slightly convex; male cephalotheca conspicuous-
ly convex and oval (Fig. 4) . . . . . . . . . . . . . . Elenchus yasumatsui Kifune & Hirashima
6' Postlumbium (Fig. 5) small; postscutellum subtriangular with a narrow and rounded
apex, occupies about 0.4 of the metanotum; vein A straight with a bent apex; male
cephalotheca globular (Fig. 6) . . . . . . . . . . . . . Elenchus japonicus Esaki & Hashimoto
7(5) Hindwing without Mf; Rf short and broad; Cu clearly shorter than 1A; 2A very short and
situated near wing margin (Fig. 7); eyes with 12 facets; antennae less compact; flabella
of antennal segments III to VI and VII equally slender but gradually decreasing in
length, 7th segment narrowed medially (Fig. 8); aedeagus with almost straight, sharp,
dorsal process longer than ventral protrusion; male cephalotheca conspicuously convex,
transversely elongate - elliptical with ventral margin trapezoidally protruded (Fig. 9)
host – Thaia oryzivora Ghauri . . . . . . . . . . . . . . . . . . . . . Halictophagus thaiae Kifune
7' Hindwings with long Mf; Rf long and broad; Cu and 1A almost equally long; 2A long
and distinctly separated from wing margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
BARRION AND LITSlNGER 15
BARRION AND LITSINGER 17
8(7') Basal portion of 3rd antennal segment about as long as that of 4th; flabella more or less
thickened (Fig. 10); aedeagus enlarged in the basal half, apex with a short dorsal process
and a long ventral protrusion (Fig. 11); male cephalotheca elliptical (Fig. 12); host –
Cofana spectra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halictophagus spectrus Yang
8' Basal portion of 3rd antennal segment about twice as long as that of 4th, flabella of
antennal segment V-VII almost equal in size and shape but smaller than III and IV with
a long, flat and stout flabella (Fig. 13); aedeagus enlarged in basal third, stem almost
straight, hook rectangularly dorsally pointed with sharp tip (Fig. 14); male cephalotheca
conspicuously convex and elongate-elliptical (Fig. 15); hosts – Nephotettix nigropictus
and N. virescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halictophagus bipunctatus Yang
9(4') Third antennal segment with 4 or more freely articulated flagellomeres; maxillary palp
3 to 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9' Third antennal segment usually consolidated into a compound segment, typically with a
terminal to dorsal bristle-like arista; maxillary palp 1 or 2-segmented . . . . . . . . . . . .18
10(9) Mesonotum with a V-shaped suture; ocelli absent; antennae 12-segmented with a
circular pedicel, bases of funicles 2 to 10 swollen and ringed with hairs; head grey
dorsally with 3 grooves above antennal sockets, front elongate-cylindrical projected
forward with an apicomedian process; postscutellum globular with small bulbous
median point; wings (Fig. 16) transparent except brownish bands along subcosta, and
cells between R1 and R2, wing length a little longer than body length; haltere as long as
abdominal segment II and III; basal tarsal segment of leg I longer than abdomen
[Tipulidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tipula aino Alexander
10' Mesonotum without a V-shaped suture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
11(10') Costa continuing around wing margin with wing venation reduced, longitudinal vein Rs
unbranched (Fig. 17) and without discal cell; antennae more than 10 segments with
bead-like joints; maxillary palpi 3 or 4-segmented [Cecidomyiidae] . . . . . . . . . . . . . .12
11' Costa disappearing beyond wing tip; veins equally developed and 2nd basal cell longer
than 1st basal cell (or basal cell imperfectly separated to apically open in 2nd basal cell);
R branched; vein M branched or unbranched; antennae with 15 segments . . . . . . . .13
12(11) Asian rice gall midge; apical end of hypoproct rounded (Fig. 18); gonostyle thinner in
apical one-third (Fig. 19); anterior margins of abdominal tergites VII and VIII without
distinct pigmented areas; maxillary palpi 4- segmented . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Orseolia oryzae (Wood-Mason)
12' African rice gall midge; apical end of hypoproct slightly notched (Fig. 20); gonostyle very
slightly tapered in apical one-third (Fig. 21); anterior margins of abdominal tergites VII and
VIII with irregular narrow pigmented bands; maxillary palpi 3-segmented (Fig. 22). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orseolia oryzivora Harris & Gagne
Figs. 16-24
13(11') Front legs not lengthened; vein M 2-branched (Fig. 23); antennae (Fig. 24) 15-seg-
mented, segments III-X much shorter than XI-XV; body brown with fore- and midlegs
yellow; forefemora without subapical pale band; anterior part of femora II and III, tibiae
II, and dorsal side of tibiae III with a series of long hard spines; ventral side of basitarsi
II and III and second tarsal segment of leg III with rows of small but dense spines; apical
tarsal segments of all legs with several long spines [Ceratopogonidae] . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nilobezzia acanthopus (de Meijere)
13' Front legs lengthened; vein M unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
14( 13') Forewings (Fig. 25) with 4-5 cloudy patches on the subapex of wing membrane in
between longitudinal veins R4+5 to A (or cell R5, M, Cu, and A); scutum with dark
brown markings; legs yellow with dark brown bands on knees, distal part of tibiae and
tarsi I-IV, tarsi V entirely black; abdominal tergites uniformly dark brown with narrow
apical pale band on tergites I-VII; dorsal appendage (Fig. 26) hook or sickle shaped
apex and enlarged basal portion with a rounded tip; style (= gonostylus) slender
towards apex with long apical bristles (Fig. 27); small species, wing length 2.8-2.9 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chironomus kiiensis Tokunaga
14' Forewings without cloudy patches on cells Rs, M, Cu, and A; wing length usually more
than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15( 14') Dorsal appendage strongly C to sickle-shaped (Fig. 28) with a well developed large
tubercle in its inner portion; ventral appendage large and distinctly long, slightly nar-
rowed before base and gradually narrowly rounded apically; style almost uniformly stout
except reduction towards a rounded apex; anal point 2-ridged (Fig. 29); scutellum with
13 bristles and 7 small setae; all tarsal segments apically dark and last 2 segments fuscous
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chironomus crassiforceps Keiffer
15’ Dorsal appendage large, spine-like, slightly curved at apex or subcylindrical with a
moderately enlarged apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16(15') Dorsal appendage large, spine-like, straight and pointed (Fig. 30); ventral appendage
greatly swollen, sounded in lateral aspect, densely hairy in the inner surface; styles short
and oval; anal point short or deep in lateral aspect and laterally flanged (Fig. 31); thorax
dull grey with pits of the dorsocentral bristles very distinct . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chironomus tepperi Skuse
16’ Dorsal appendage not acutely pointed, tips slightly curved or swollen; anal point long to
moderately long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
17( 16') Ventral appendage (Fig. 32) with about 17 setae, slightly pointed in the inner-apical
area; dorsal appendage (Fig. 33) slightly knob-like reaching midhalf of the ventral
appendage (sometimes inwardly curved at tip); anal point prominently developed, long,
rounded at apex and narrowed before a little broad base; style gradually widened at
midhalf in the outer margin with 7 setae along margins and 2 setae in the middle behind
the former set (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chironomus dorsalis Meigen
Figs. 25-35
17' Ventral appendage (Fig. 34) with 10-14 setae and bluntly shaped apex; dorsal ap-
pendage (Fig. 35) straight with an inwardly curved tip; anal point small, widened basally
and rounded at apex; style with a swollen basal half and narrowed apical half from the
outside margins, inner apical margins with 6 setae (Fig. 34) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Chironomus javanus Kieffer
18(9') Ptilinal suture and lunule absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
18' Ptilinal suture and lunule present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
19(18) Head as large as thorax and subspherical, bounded mainly by the enormous eyes;
antennae with dorsal arista; R4 + 5 unbranched; big-headed fly parasites [Pipunculidae]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19' Entirely not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
20(19) Stigma absent on wings between Sc+ R1; cross vein r-m almost midway discal cell . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20' Stigma present between Sc + R1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
21(20) Humeri yellowish white; wings hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
21' Humeri grey or black (Fig. 36); brown tinge present on wings; eyes not joining on front
with narrow convergence point on anterior of ocelli (Fig. 37); hosts - nymphs and
adults of Nephotettix spp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tomosvaryella oryzaetora Koizumi
22(21) Hind trochanter trapezoidal (Fig. 38); piercer of ovipositor relatively straight (Fig. 39);
genitalia of male symmetrical in dorsal outline (Fig. 40), 9th segment glossy and broad
ventrally; host — Nephottetix spp. . . . . . . . . . . . . . Tomosvaryella subvirescens (Loew)
22' Hind trochanter with a tubercle-like process; male genitalia moderately tapered at tip,
dorsally asymmetrical (Fig. 41); ovipositor of piercer downcurved (Fig. 42); hosts — N.
cincticeps, Psammotettrix striatus (L.) . . . . . . . . . . . . . Tomosvaryella sylvatica (Meigen)
23(20') Latero-basal half of lst abdominal tergum without prominent setae; in males,
membranous area in the rounded genitalia absent, genitalia as long as but not as wide as
5th abdominal segment (Fig. 43) . . . . . . . . . . . . . . . . . . . . . Pipunculus roralis (Kertesz)
23' Latero-basal half of 1st abdominal tergum with a row of prominent setae; membranous
area present in male genitalia, dorsally absent in P. javanensis . . . . . . . . . . . . . . . . . 24
24(23') Membranous area of male genitalia caudo-ventrally present, genitalia notched dorso-
apically (Fig. 44); in females, tip of ovipositor prominently upcurved extending to
posterior margin of 5th abdominal sternite starting from apex of abdomen (Fig. 45); 3rd
antennal segment dark brown; legs black with yellow patches along tibiae and tarsi ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pipunculus javanensis de Meijere
24' Membranous area dorsally present bisecting the segment; ovipositor upcurved or
straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25(24') Broad longitudinal cleft present middorsally in male genitalia (Fig. 46); piercer of
female’s ovipositor long and nearly straight (Fig. 47); host — Nephotettix cincticeps . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pipunculus orientalis (Koizumi)
25' Narrow oblique cleft present in the right side of male genitalia (Fig. 48); piercer of
female’s ovipositor strongly upcurved (Fig. 49); 3rd antennal segment of female light
yellowish brown; legs yellow except femora and distal end of tarsi . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pipunculus mutillatus Loew
26(19') Wing venation reduced, discal cell and crossvein m absent (Fig. 50); anterior veins very
much thickened and crowded basally; oblique veins M1+ 2, M3 and Cul weak; head
subglobular with oblong eyes and 1-segmented antenna (Fig. 51); thorax yellow and
humpback; hindlegs long with laterally flattened femora; females with 4 scutellars and 2
pairs of equal supra-antennals; humpbacked fly parasites [Phoridae] . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megaselia scalaris (Loew)
26' Wing venation developed; discal cell, crossvein m and r-m present; oblique veins
prominent; antennae 3-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27(26') Arista dorsal of the cone-shaped 3rd antennal segment (Fig. 52); third costal section
equal to or longer than the fourth; crossvein M shorter than apical portion of M3 + 4;
acrostichals present; femora swollen medially and thinner apically . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsicnemus sp.
27’ Arista in front of 3rd antennal segment; rest of characters not as above . . . . . . . . . 28
28(27') Black body with whitish yellow legs and hyaline wings; head and antenna black (Fig. 53);
hind tibiae without extensor bristles; subcostal vein a little inside the basal one-third of
costal length; base of vein R + Cu at midlength of subcostal vein; postical vein erased in
apical half (Fig. 54) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drapetis (Drapetis) sp.
28' Brownish red body with yellowish brown legs; head and antenna reddish brown (Fig.
55); hind tibiae with two extensor bristles; subcostal vein reaches midlength of costal
length; base of vein R + Cu at basal one-fourth of subcosta; postical vein prominent, not
erased (Fig. 56); midabdominal tergites blackish brown . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drapetis (Elaphropeza) sp.
29(18') Second antennal segment nearly always with longitudinal seam; lower calypter usually
large and thorax nearly always with a complete transverse suture before wings . . . . . . . 30
29' Second antennal segment without longitudinal seam; lower calypter vestigial and thorax
without complete transverse suture before wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
30(29) Hypopleural setae absent; and vein not reaching wing margin, 2nd anal vein not curving
forward around apex of first; terminal 3 abdominal segments with a pair each of grayish
black spots (Fig. 57), sometimes absent in segment V (Fig. 58); females with a pair of
small anterior plates on tergite VIII of ovipositor (Fig. 59); male trifoliate process with
the stalk enlarged around middle; shoulder weak, median piece membranous and
lateral plates developed; hypopygial prominence with 2 lateral arms widely separated
and diverging outwards from tergal surface (Fig. 60), seeding maggot . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atherigona oryzae Malloch
30' Hypopleural and pteropleural bristles very prominent; postscutellum well developed;
R5 cell narrowed or closed distally; abdominal terga with strong bristles with edges
overlapping sterna; arista usually bare; parasitic flies [Tachinidae] . . . . . . . . . . . . . 31
31(30') Face elevated into a very strong, broad roof-like facial keel running from epistome to
lunula; eyes bare; palpi highly reduced, much shorter than 3rd antennal segment;
sternopleuron with 2 setae; pteropleural setae absent; proboscis very long and slender,
length much greater than height of head; R4 + 5 without setae, apical and fused with
straight and oblique M1 and m-cu arises at about apical one-third of M (Fig. 61);
parasitic on larvae of scarabaeid genera Adoretus, Anomala, and Leucopholis . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prosena siberita (Fabricius)
31' Face not as above, without such facial keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32(31') Bend of vein M abruptly angulate (Fig. 62); M1 concave basally and convex apically;
petiole (R4 + 5 + M1) slightly curved upwards producing a cleft at their junction; basal
node of R4 + 5 with fine hairs ventrally, hairs shorter than r-m in length; base of costa
without long hairs; palpi absent; lower calypters with only very short marginal fringe;
abdomen long and slender, subcylindrical; T1+ 2 with excavation in the basal one-fifth
to one-sixth; parasitic on pentatomid bug Eysarcoris . . . . . . . . . . . . Cylindromyia spp.
32' Not entirely as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33(32') Pteropleural setae absent; eyes and prosternum bare; epistome strongly projecting,
highly visible in profile; vibrissae high above level of epistomal margin . . . . . . . . . . 34
33' Pteropleural setae prominent; eyes hairy or bare; prosternum hairy or setulose and with
vestiture; epistome not projecting and not visible in profile or weakly projecting; vibris-
sae a little over level of epistomal margin; 3 post intra-alar setae present, anterior one as
close to the transverse suture as to the posterior one (Goniinae) . . . . . . . . . . . . . . .35
34(33) Hindcoxae with posterodorsal fine soft hairs; veins R1 and R4 + 5 bare except for fine
hairs on the basal node of R4 + 5; antennae inserted a little above midlength of eye, 2nd
segment moderately long, 3rd segment short and broadly rounded; parafrontal, parafa-
cial, and wide gena hirsute; peristomal setae 8 and strongly developed; eye small and
ovoid (Fig. 63) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cuphocera varia F.
34' Hindcoxae bare posterodorsally; vein R1 setulose and vein R4 + 5 setulose up to or

beyond r-m (Fig. 64); 2nd costal sector haired underneath; antennae very small, in line
to the basal one-third of eye; eyes largely oblong; gena small; peristomal hairs less than
8 (Fig. 65); orange-yellow eyes, silvery white face; frons wider than facial region as eyes
rather converged ventrally (Fig. 66); abdomen slender, concolorous with eyes except for
silvery white apices of tergites 1 + 2 and 3 and blackish tip . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halydaia luteicornis Walker
35(33') Pre-alar setae strong and long, longer than 1st post intra-alar (ia) setae and Ist post
dorsocentrals (dc); sternopleuron with 4 setae; 2nd costal sector without hair ventrally;
apical setae not enclosed by the nonconverging subapicals . . . . . . . . . . . . . . . . . . . . 36
35' Pre-alar setae weak and small, shorter than Ist post ia and Ist post dc; sternopleuron
with less than 4 setae; 2nd costal sector with or without hairs ventrally; apicals some-
times enclosed by the converging subapical setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
36(35) Barette completely haired; humeral callus with 5 strong setae, the main 3 arranged in a
triangle; tibia II with a single submedian anterodorsal seta; sternopleuron with 2 setae;
eyes hairy; antennae high above epistome, arista long, segment II of antennae long
almost 2/3 of segment III; vibrissa long (Fig. 67); tergite V of abdomen broadest basally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemorilla maculosa Meigen
36' Barette haired only anteriorly or bare and setae on humeral callus not more than 4 . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37(36') Head shape rather abnormal (Fig. 68), strongly triangular in profile; frons almost
horizontal; profrons much wider than gena; arista thickened in its entire length, with 2nd
segment elongate and 6X as long as broad; antennae inserted up high in level to the
lower end of the apical one-third of height of eye; 3rd antennal segment beyond
epistome, long and subcylindrical; eyes, parafacials and facial ridge all bare; cell R5
triangular with short M2 and petiole towards wing apices . . . . . . . . Diatraeophaga sp.
37' Head normal, other characters not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38(37') Genal depth reduced to a narrow area smaller than the length of profrons and
obliterated by the very large eyes; parafacials bare; propleural setae present; humeral
callus with no more than 4 setae, 3 main ones arranged in a straight line (Carceliini) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
38' Gena as wide as or wider than profrons; head with one or more pairs of reclinate orbital
setae; vibrissae inserted above the level of the epistomal margin . . . . . . . . . . . . . . . . 42
39(38) Sternopleuron with 4 setae; frontal setae normal or mostly inclinate; apical scutellar
setae horizontal; tibiae III with 2 or more anterodorsal setae; antennae long with 3rd
segment almost reaching epistome; eyes large forming narrow gena (Fig. 69); parasitic
on larvae of rice skippers Pelopidas and Pamara . . . Thecocarcelia oculata (Baranov)
39' Sternopleuron with 2 or 3 setae; eyes bare or hairy . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40(39') Eyes haired (Fig. 70); post dorsocentral with 4 setae; proclinate orbital setae absent;
frontal setae 5; profrons twice the height of gena; parafrontal hairy unlike parafacial;
antennae inserted just above midlength line of eyes; 3rd segment not reaching epistome;
vibrissa1 setae not crossing arista . . . . . . . . . . . . . . . . . . . . . . . . . . Carcelia excisa Fallen
40' Eyes bare; post dorsocentral with 4 setae; propleural setae present though small some-
times; sternopleuron with 2 or 3 setae (Argyrophylax) . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41(40') Third antennal segment reaching epistome (Fig. 71); palpi yellow; head generally dark
to blackish red except silvery white tinge along parafacial and facial ridge; marginal
setae of abdominal tergite IV extended beyond tip of abdomen; surstylus very small and
well separated from cerci (Fig. 72); cercus tapers towards apex, in lateral view biconcave
dorsally; epandrium wider than long and hairy dorsally (Fig. 73) . . . . . . . . . . . . . . . . . . .
............................................. Argyrophylax nigrotibialis Baranov
41' Third antennal segment relatively far from epistome (Fig. 74); palpi yellow; surstylus
(Fig. 75) very well developed, broad and rounded at apex; cercus outwardly curved in
lateral view and closely attached to surstylus; epandrium (Fig. 76) slightly hairy dorsally
............................................... Argyrophylax phoeda Townsend
41' ' Similar to the above two species except genitalia; palpi black with yellowish tip (Fig. 77);
marginal setae of abdominal tergite IV relatively close to tip of abdomen. . . . . . . . . . . .
................................................ Argyrophylax fransseni Baranov
42(38') Eyes densely hirsute ..................................................... 43
42' Eyes bare or with minute hairs ............................................. 44
43(42) Parafacials entirely clothed with strong hairs (Fig. 78); antennae far from epistome,
yellow-brown except segment III; head silvery white except dark brown interfrontal area
with shallow, fine longitudinal grooves; abdomen same color as head, cerci triangular in
apical view (Fig. 79); surstylus elongate and rounded at tip (Fig. 80); legs blackish brown
with whitish yellow broad pulvilli; parasitic on rice stem borers – Chilo spp. and
Scirpophaga nivella ............................. Sturmiopsis inferens Townsend
43' Parafacials bare; facial ridge rather flattened with strong setulae beyond half its length;
4 strong setae lined peristomal area; interfrontal area as long as parafrontals; antennae
inserted in line to lower apical 1/3 of eye, 3rd segment long, nearly touches epistome and
vibrissa (Fig. 81); basal node of R4 + 5 with about 2-4 small setae; male abdomen with
light and uneven silvery grey tinges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.................................. Pseudoperichaeta insidiosa Robineau-Desvoidy
44(42') Ocellar setae very prominent, equal in size to reclinate orbital setae and inserted out of
the triangle in front of anterior ocellus; yellow epistome straight; parafacials bare; eyes
with few minute hairs (Fig. 82); antennae and aristae black; silvery white along parafron-
tals; abdomen silvery white except black T1+2 and mid- and posterior margins of
tergites; legs black with silvery white femora . . . . . . . . . . . . . Zygobothria ciliata Wulp
44' Ocellar setae small, weak, and wiry; parafacials with fine hairs apically; well defined
hair-patch unusually small or usually large and thick with very long hairs converging into
a fascicle present on each side of venter of abdominal segment IV in males; tibiae II with
one submedian anterodorsal seta; aedeagus bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . 45
45(44') Palpi moderately short, yellowish brown; sternopleuron with 2 setae; peristomal area
with 4 setae; hair-patch large, covers most of venter of abdominal segment IV (Fig. 83);
upper occiput without black setulae behind postocular row in females; frons narrow;
legs brownish black; abdomen black with few silvery dust; paralobes with spinules
apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palexorista lucagus (Walker)
45' Palpi long beyond epistome, yellow; sternopleuron with 4 setae; peristomal area with 9
setae; hair-patch very small (Fig. 84), sometimes obliterated; female’s upper occiput
with a distinct irregular row of black setulae at the back of postocular; antennae long
with 3rd segment nearly 3 times as long as 2nd segment; antennal axis high above ocular
axis; parafacials with fine hairs on upper half . . . . . . . . . Palexorista solensis (Walker)
46(35') Parasites of acridid grasshoppers; scutellum with 3 pairs of strong marginal setae,
apicals crossed in basal one-third, subapicals widely diverging, basals cross subapicals
slightly converging and closed to distal one-third of apicals, lateral setae absent; basal
node of R4 + 5 with one strong setula; eyes bare; bend of M oblique to meet R4 + 5
before apex (Fig. 85) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceracia spp.
46' Not parasitic on acridid grasshoppers; arrangement of marginal setae of scutellum
different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
47(46') Subapical scutellar setae strongly convergent and crossed at their apices; 2nd costal
sector haired below up to 3rd sector; costal margin conspicuously broken and incised at
the apex of subcosta; vein R4+5 extensively setulose, R1 partly or entirely setulose;
propleuron not hairy; hind tibiae with 3 dorsal preapical setae; eyes bare; proboscis
normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
47' Subapical scutellar setae parallel or diverging; 2nd costal sector bare below, costal
margin inconspicuously broken and not incised at apex at subcosta; T1+ 2 fully ex-
cavated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
48(47) Two equally long and strongly diverging (one directed upward, the other downward)
prostigmatic setae present on thorax; head silvery white except yellow interfrontal area;
eyes strongly parallel-sided (Fig. 86); parafrontals with a few short hairs; antennae dark
brown with brown segment II and base of segment III; peristomal area with 3 setae;
palpi yellow; Sc with a bend and 2-3 hairs above bend; R1 with 13 hairs; basal node of
R4 + 5 with 15-16 hairs extended to top level of bend of M . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peirbaea orbata (Wiedemann)
48’ One strong prostigmatic seta directed upwards present on thorax; sternopleuron com-
pletely hairless laterally in front of midcoxae; eyes moderately small, widen toward
genae (Fig. 87) with 2 peristomal setae; epistome curved forward; wings with strong
setulae along costal margins, R1, R4 + 5 and Cul; M1 near R4 + 5 but clearly separated,
bend of M not angled (Fig. 88) . . . . . . . . . . . . . . . . . . . . . . . . . Ceromya silacea Meigen
49(47') Dorsocentral with 3+3 setae, post dorsocentral with 3 setae arranged in a triangle; head
grey except black occiput, reddish bottom part of parafacials; reddish brown frontal
vitta; palpi pale orange-yellow; antennae dark grey with reddish base of segment III;
cerci slender, rounded at apices (Fig. 89) and distinctly divergent in rear view; abdomen
orange with grey pollinose patches; legs reddish brown but pale along femora and tibiae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lixophaga diatraeae Townsend
49' Dorsocentral with 3-4 setae; post dorsocentral with 4 setae; slender palp brownish
yellow; ocellar setae very strong; parafacial setae absent (Fig. 90); antenna black, far
from epistome, inserted almost in line to top of eye; cerci pointed at tip and surstylus
with a long process (Fig. 91); legs black except white pulvilli and greyish outer femora;
abdomen black with silvery white bands along apical half of tergites III to V . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Exorista xanthaspis (Wiedemann)
50(29') Costa without breaks; subcosta perfect, nearly always separate from R1 and ending
independently in costa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
50’ Costal break just before end of subcosta and or near humeral crossvein; or costa
constricted, partially fractured or broken before end of R1; subcosta imperfect . . . 53
51(50) Eyes normal, close to each other and not on a stalk; wings blackened by markings leaving
3 transverse to oblique white bands towards apex and patches of irregularly arranged
white spots in lower basal (Fig. 92) area; anal cell without an acute lobe; scutellum
broadly rounded at apex with long setae along margins; base much wider than apex;
head in dorsal view pointed forward below base of antennae; body with bluish reflec-
tions; face, genae, and antennae ochreous yellow; legs brownish with tarsi ochreous
yellow except its apex [Platystomatidae] . . . . . . . Poecilotraphera taeniata (Macquart)
51' Eyes situated at the ends of long stalks; in dorsal view at least 3 times as wide as long
with antennae well separated; scutellum bituberculate; front femora much swollen
[Diopsidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52(51') Wings with a large, round, black apical spot (Fig. 93); frons with a dark brown stripe
connecting bases of stalk (Fig. 94); yellow to pale yellow-brown with black tip; scutellum
pollinose, dark to blackish brown; dorsum of thorax shining; body length (head to tip of
abdomen) nearly as long as width of 2 compound eyes; species barely 6 mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diopsis apicalis Dalman
52' Wings without such black apical spot (Fig. 95); frons with a light brown stripe (Fig. 96);
facial teeth rather long and more widely separated than in D. apicalis; scutellar spines
long, yellow-brown except one-fourth to one-third apical area black; scutellum yellow;
distance from eye to eye much longer than body length; species 7-9 mm in length . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diopsis macrophthalma Dalman
53(50') Anal cell absent; ocellar triangle conspicuously large and reddish brown; crossvein r-m
beyond midhalf of discal cell; cell CuP and vein CuA + 1A absent, cell M confluent with
discal cell; mesonotum with 3 broad longitudinal and 2 short subposteromarginal
blackish brown bands (Fig. 97); body dark yellow, legs yellow [Chloropidae] . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlorops oryzae Matsumura
53' Anal cell present; ocellar triangle less conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

54(53') Costa broken only near end of R1; postocellars divergent; oral vibrissae present
[Agromyzidae] .......................................................... 55
54' Costa broken near end of subcosta and also near humeral crossvein; distinct oral
vibrissae absent; postventricals divergent [Ephydridae] ....................... 56
55(54) Subcosta developed throughout its length coalescing with R1 before reaching costa;
costa extending strongIy to vein M1+2; crossvein r-m slightly beyond midpoint of discal
cell (Fig. 98); 3 pairs (4 + 2) of dorsocentrals and aerostichals in 6 rows; frons narrowly
projecting above eyes (Fig. 99); orbital setulae sparse and reclinate; third antennal
segment axe-shaped with slightly pubescent arista (Fig. 100); tibia II with 2 strong lateral
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agromyza oryzae (Munakata)
55' Subcosta becoming a fold distally and ending in costa separately and basad of R1; costa
ending at vein R4+5; mesonotum with 3 dorsocentrals and 4 aerostichals in irregular
rows; frons not projecting above eyes; orbital setulae upright; third antennal segment
with blunt angle at upper corner (Fig. 101) . . . . . . Pseudonapomyza asiatica Spencer
56(54') Leg I mantis-like, enlarged femur with spines and tibiae with apical spur (Fig. 102);
basal tarsal segment of leg III shorter than combined length of segments II-V; basal
tarsal segment of leg I much shorter than femora I; basal ventral spine of femora I not
reaching apex of basal tarsal segment of leg I; legs greyish black, tarsi of leg II slightly
yellow with blackish shades and black terminal segment; sides of tergite II with a mass
of hairs; abdomen brownish with silvery tinge laterally . . . . Ochthera sauteri Cresson
56' Leg I as above; basal tarsal segment of leg III as long as rest of segments (II-V); basal
tarsal segment of leg I slightly longer than femora 1; basal ventral spine of femora I
reaching apical end of basal tarsal segment of leg I; legs blackish brown with dense
yellow-white dust, tarsi and tibiae silvery in front; tarsi of leg II all yellow; labial palp
dark brown; abdomen dark olive green with white dusted triangular spots at sides of
segments; sides of tergite II without mass of hairs (Fig. 103) . . . . . . . . . . . . . . . . . . . . . . .
............................................... Ochthera brevitibialis de Meijere
56'' Foreleg and tibia normal, apical spur absent ................................. 57
57(56'') Mesonotum without well developed setae; dorsocentrals absent; ocellars behind
anterior ocellus or between posterior ocelli; 2nd antennal segment with dorsal spine-like
strong bristles (Fig. 104) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psilopa spp.
57' Mesonotum with well developed setae; head and thoracic bristles long and strong; arista
with several long dorsal hairs; 1 or 2 orbitals present and 2-3 verticals; 2-3 dorsocentrals
(dc) present [Notiphilinae] ............................................... 58
58(57') Midtibiae without strong dorsal bristles; dull gray to brownish black; eyes with pubes-
cence; 3rd antennal segment with 1 or 2 weak dorsal bristles; several facials feeble and
hair-like; 1st costal section without spine-like bristles apically; 2-4 dorsocentrals (dc)
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
58' Midtibiae with 3-4 erect dorsal bristles; 1 reclinate fronto-orbital strong; 3-5 weak to
moderate facial bristles; costa extending to or slightly beyond tip of 3rd vein; arista with
many long hairs dorsally [Notiphila] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
59(58) Four dorsocentral setae present; face bordered with 6 facials; frons and cheeks silvery
white; mesonotum greyish with silvery white and brown tinges, scutellum silvery white to
grey; antennae dark grey with light silvery tinge in the inner portion of 2nd segment, 7-10
aristal hairs (Fig. 105); abdomen silvery white to grey with blackish brown medians in
the basal three segments; legs yellow except femora I to III; inner portion of femur I with
about 10-12 spines (Fig. 106). . . . . . . . . . . . . . . . . . . . . . . . . Hydrellia philippina Ferino
59' Four dorsocentral setae present; face golden yellow covered with silvery tinge, bordered
with 4 weak to moderate facials, frons black with greyish tinge; cheeks greyish white;
mesonotum and scutellum brown; antennae black with 5-6 aristal hairs (Fig. 107); head
index 5:1; abdomen greyish to dark brown, legs black with greyish tinge and yellow
metatarsi, inner portion of femur I armed with more than 25 small teeth (Fig. 108)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrellia griseola (Fallen)
59'' Two dorsocentral setae present; face and lunula silvery white with 5-6 moderate facials
(Fig. 109); frons and antennae black; mesonotum and scutellum black with dense brown
tinge; third antennal segment with whitish pubescence, 8 aristal hairs present; abdomen
half shining black but yellow apically; tibiae and tarsi all yellow; ventral process of
hypopygium thin and very slender . . . . . . . . . . . . . . . . . . . . . Hydrellia tomiokai Miyagi
59''' Two dorsocentral setae present; face golden yellow to yellowish brown with 6 hair-like
facials (Fig. 110); frons blackish brown with velvety black laterals; antennae black, 3rd
segment with brownish pubescence; 7 aristal hairs; cheeks and occiput dark grey;
abdomen brown, coxae yellow, femora black, tibiae yellow except midtibiae brownish
yellow, basal half of hind tibiae blackish brown . . . . Hydrellia sasakii Yuasa et Isitani
60(58') Root maggot; midtibiae with 4 extensor bristles (Fig. 111); several facial bristles weak
and hair-like; 5th tergite in male forms a cylindrical tubercle with 2 stout upcurved
bristles apically; Japanese species . . . . . . . . . . . . . . . . . . . . . . Notiphila sekiyai Koizumi
60' Leaf miner; midtibiae with 3 extensor bristles (Fig. 112) . . . . . . . . . . . . . . . . . . . . . . 61
61(60') Antennae with 12-14 aristae tip and outer edge of 3rd antennal segment dark brown;
ocellar bristles set relatively closer to the front of lateral ocelli; 2 strong parafacial
bristles (Fig. 113); eyes moderately oblong; legs dark grey except yellow apices of
femora, and brownish yellow tibiae and tarsi; abdomen silvery white with broad blackish
brown spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notiphila similis de Meijere
69(68') Head uniformly black (Fig. 120); anterior margin of pronotum with a pair of dark brown
outwardly acute calli (Fig. 121); posterior portion straight; baso-clypeus nearly as wide
as distance between eyes taken across lateral ocelli; antennal socket in a deep groove;
fronto-clypeal inflection medially recurved below median ocellus; basal segments of
antennae blackish brown; mirror wider than long (Fig. 122) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plebeiogryllus plebejus (Saussure)
69’ Head with a faint to distinct yellow band above antennal bases running towards dorsal
eye margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
70(69') Head with clear dark yellow band in the internal margin of eye (Fig. 123); in males,
mirror (Fig. 124) longer than wide, width less than 3.1 mm (sometimes with a closed
cell); stridulatory file much closer to each other, with teeth number ranging from
210-292; females with 11-13 veins in the lateral field of forewings, veins IV and V mostly
widely separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Teleogryllus occipitalis (Serville)
70’ Head with faint yellow band (Fig. 125); in males, mirror (Fig. 126) as long as wide
(normally without closed cell), width greater than 3.2 mm; stridulatory file more widely
separated, bears 134 teeth with a maximum of 215 in females; with 13-15 equally spaced
veins in the lateral field of forewing and terminal V or VI branched from Sc . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Teleogryllus mitratus (Burmeister)
71(67') Ovipositor without differentiated apical valves; sinuated and S-shaped (Fig. 127); elytra
extended almost to apex of abdomen; anterior tibia with external and internal tym-
panum of equal size (Fig. 128); posterior tibia serrulated between spines, long spines
more than 6 pairs; pale brownish yellow gryllids; phytophagous
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euscyrtus concinnus (Haan)
71' Ovipositor not S-shaped; posterior tibia not serrulated; predatory gryllids . . . . . . 72
72(71') Elytral venation different in both sexes, male with neat anal field and mirror (Fig. 130);
female elytra feebly convex, without false veins, and with 5 straight veins in the dorsal
field; head without V-shaped mark dorsally; scape and pedicel pale yellow; terminal
segment of maxillary palpi triangular with a broad truncated tip; 2nd tarsal segment
yellow (Fig. 129); head, thorax, wings, and legs yellow . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anaxipha longipennis (Serville)
72’ Elytral venation similar in both sexes; male without mirror; head with a light yellow to
white mark dorsally, scape and pedicel black; terminal segment of maxillary palpi black
with brown tinge, and with a moderately broad and truncated tip; 2nd tarsal segment
black (Fig. 131); generally black with pale brownish red tinges in wings and abdominal
dorsum, legs whitish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . Metioche vittaticolis (stål)
73(64') Fastigium projected obliquely above eyes by 2 eye lengths, deeply notched frontally in
line with base of antennae (Fig. 132); face diagonal, lateral lobes of pronotun almost 2
times longer than deep, hind lateral part of lobe rounded above coxa I; ovipositor
sword-like, as long as hind tibia; venter of tibia III with 20-21 spines; forewings as long
as hindwings; hind femora slightly swollen . . . . . . . . .Euconocephalus varius (Walker)
73' Fastigium not projected as above; lateral lobes of pronotum broadly rounded above
coxae I, or narrowly rounded along its lower posterior margin behind coxae I and
slightly indented above it; forewing shorter than the hindwing, green or brownish yellow
with brownish tinge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
74(73') Hindwings usually 1/3 longer than the forewings, wing tips more rounded, base of hind
femora noticeably swollen, venter spineless; ovipositor very short but broad . . . . . 75
74' Hindwings a little longer than forewings; wing tips pointed; prosternum bispinose;
venter of femora with or without spines; ovipositor very long . . . . . . . . . . . . . . . . . . 76
75(74) Hind femora prominently attenuate; lateral lobes of pronotum longer than deep, slightly
concave in the antero-dorsal 1/5 with pointed postero-dorsal tip projected relatively
upward (Fig. 133); cerci (Fig. 134) very long, wide subapically prior to a tapering end;
subgenital plates elongate and bifid (Fig. 135); supra-anal plate knob-like; costal margin
of forewings with a brown tinge . . . . . . . . . . . . . . . . . . Phaneroptera gracilis Burmeister
75' Hind femora not attenuate; lateral lobes of pronotum as long as deep, partially concave
along antero-dorsal 1/3 with pointed postero-dorsal tip slightly curved downwards;
subgenital plate robust, elongate, and sulcate medially above and below; cerci with a
long thin process at tip (Fig. 136); supra-anal plate deeply bifid (Fig. 137); costal margin
of forewing uniformly green except for a pair of brownish black spots along basal 1/4;
abdominal tergite X enlarged (Fig. 138) . . . . . . . . . . . . . . . . . Phaneroptera furcifera Stål
76(74') Fastigium of vertex narrow (Fig. 139); forewings unicolorous green; hind femora with
3-7 externo-ventral spines (Fig. 140); cerci with 1 internal spine, globular at apex (Fig.
141); subgenital plate with excised apex; ovipositor (Fig. 142) relatively straight, 13-16
mm in length; thorax and abdomen yellow . . . . . Conocephalus longipennis (de Haan)
76' Fastigium moderately wide; forewings (Fig. 143) pigmented with large dark spots,
without markings along costal and precostal areas; hind femora (Fig. 144) without
ventral spines; cerci (Fig. 145) of males with 1 internal spine, pointed at tip; ovipositor
very short 6.6-8.2 mm (Fig. 146) . . . . . . . . . . . . . Conocephalus maculatus (Le Guillou)
77(63') Fastigium of vertex with a deep thin median furrow; lower basal lobe of hind femora
longer than upper portion [Pyrgomorphidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
77' Mediolongitudinal sulcus in the anterior of fastigium absent; lower basal lobe of hind
femur as long as or shorter than upper portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
78(77) Pronotum rough and strongly tuberculated (Fig. 147); dorsally black with anterior
tubercles and lateral lobes yellow; wings large, tegmina long with parallel sides olive
green with numerous subglobular yellow spots (Fig. 148); abdomen black with reddish
orange bands; body length 37-55 mm in males and 47-69 mm in females . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aularches miliaris (Linnaeus)
78' Pronotum rather smooth, non-tuberculated; wings of different colors . . . . . . . . . . 79

79(78') Tegmina narrow but well developed with rounded tip; femora shorter than tegmina
(Fig. 149); head conical, shorter than pronotum; face oblique (Fig. 150); fastigial furrow
short reaching less than 1/2 distance to anterior margin of eyes; pronotum dorsally
rounded with feeble median carina lightly bissected by 2 transverse sulci; hind tibia with
terminal spines sharply pointed; outside spines shorter than inside . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tagasta marginella (Thunberg)
79' Tegmina long and narrow with tips pointed; head acutely conical . . . . . . . . . . . . . . 80
80(79') Fastigium (Fig. 151) of vertex longer than the length of compound eyes; head lanceolate;
ratio of fastigium length to least width between eyes 2.5-3 in females and 3-3.3 in males
(Fig. 152); exceptionally slender, pronotum (Fig. 153) of females with 2 wide apart sulci
without cross “veins” . . . . . . . . . . . . . . . . . . . . . . . Atractomorpha psittacina (de Haan)
80' Fastigium (Fig. 154) of vertex well developed, shorter than the length of compound eyes;
head conical; ratio of fastigium length/least width between eyes – 2.25 in males and
1.7-2.1 in females (Fig. 155); hindwings light purplish red; membranous area of lateral
lobe prominent; sulci of pronotum with a crossvein (Fig. 156); phallic parts small; valves
also small, conically short but slightly curved; small species; less than 20 mm in males
and greater than 20 mm in females . . . . . . . . . . . Atractomorpha crenulata (Fabricius)
81(77') Prosternal peg absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
81' Prosternal peg present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
82(81) Sword-like antennae with basal segments flat and wide . . . . . . . . . . . . . . . . . . . . . . 83
82' Antennae filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
83(82) Anterior tip of hind femora slender with a pair of spur-like processes (Fig. 157), base
partially enlarged; compound eyes much closer to the anterior of head; head strongly
elongated, much longer than pronoturn with a median transverse furrow; face strongly
oblique (Fig. 158); tip of tegmina pointed (Fig. 159) . . . . . . . . Acrida willemsei Dirsh
83' Anterior tip of hind femora moderately swollen, spur absent (Fig. 160); compound eyes
midway head length; head shorter than pronotum; pronotum laterally compressed,
median and lateral keels distinct; three transverse furrows present with the 2 anterior
not visible dorsally; eyes, sides of vertex and pronotum and occiput with reddish brown
suffusion (Fig. 161) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonista bicolor (de Haan)
84(82') Pronoturn constricted anteriorly with tubercles on median carina in front of posterior
transverse furrow (Fig. 162); rod-like antennae slightly swollen in apical 1/3 and little
longer than combined length of head and pronotum; fastigium of vertex angular with
straight frons slightly oblique; metazonal posterior margin obtuse apically; hind femora
robust with 2 black spots dorsally; general coloration chocolate brown . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trilophidia annulata (Thunberg)
84' Pronotum without tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
85(84') Hindwings with blackish brown bands across middle enclosing yellow areas . . . . . . 86
85' Hindwings without blackish brown markings along midhalf of its length . . . . . . . . . . 87
86(85) Grasshoppers, barely 25 mm long; pronotum short with sides parallel anteriorly but
expanded posteriorly (Fig. 163); dorsally bears a distinct light x-mark; apical half of
tegmen semi-transparent, basal 1/2 of leading edge with 3 pale areas (Fig. 164); hind-
wings with a broad dark band enclosing basal greenish yellow area; coloration greenish
brown with hind tibiae red distally . . . . . . . . . . . . . . . . Oedaleus abruptus (Thunberg)
86' Body length ranges 26-33 mm in males and 36-52 mm in females; antero-dorsal tip of
pronotum strong arched forward, posterior tip acutely angled and pointed (Fig. 165);
tegmina with a white transverse band, 1/3 of distance to apex, 2nd band indistinct (Fig.
166); hind femora ventrally and internally yellow, outer margins with series of small
black spots dorsally and ventrally . . . . . . . Gastrimargus marmoratus grandis Saussure
87(85') Mid-tegmina lined with dense row of thickened parallel veins (Fig. 167); antennae
filiform, apical 1/2 of segments black, length longer than the combined length of head
and pronotum (Fig. 168); inner apical spine of hind tibiae much longer than the outer
one; general coloration: brown to dark blackish brown in head dorsum and pronotum
and reddish in hind tibiae . . . . . . . . . . . Heteroptemis respondens respondens (Walker)
87' Mid-tegmina without thickened parallel veins ................................ 88
88(87') Pronotum lower than head top (Fig. 169); median carina not clearly elevated; midseg-
ments of antennae barely 2 times longer than broad; eyes closer to each other; hind
tibiae reddish brown along mid-half and both ends with greenish bands in between, hind
femora black to reddish brown but greenish dorsally and ventrally; inner 1/2 of fore-
wings costa green; body 30 mm in length . . . Ailopus thalassinus tamulus (Fabricius)
88' Pronotum a little higher or head in line with the median carina; head rounded anteriorly
(Fig. 170); eyes widely separated; mid-antennal segments slender, nearly 5 times longer
than broad; tegmina with brownish tinges on veinlets (Fig. 171); hind tibiae yellowish
with brown to blackish tinges distally and proximally; body more than 34 mm long
......................................... Locusta migratoria manilensis (Meyen)
89(81') Row of thickened parallel veinlets at right angles to the main longitudinal veins in the
radial area of tegmina absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
89' Row of thickened parallel veinlets present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
90(89) Venter external lobe of femur III with an extended spine at apex . . . . . . . . . . . . . . . 91
90' Venter external lobe of femur III without an extended spine apically . . . . . . . . . . . . 99
91(90) Head elongated, conical fastigium well developed, its length nearly as long as the longest
eye diameter; slender species: males 20-27 mm, females 25-32 mm long [ Quilta ] . . . . . .
......................................................................... 92
91' Head subconical; fastigium wider than long; more robust grasshoppers . . . . . . . . . .93
92(91) Fastigium long and narrow, its length nearly subequal to eye length measured from
above (Fig. 172); ratio of length of fastigium to the minimum distance between eyes
taken dorsally more than 2; body length 23-27 mm in males, 27-31 mm in females . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Quilta oryzae Uvarov
92' Fastigium broader but short, its length relatively shorter than eye length (Fig. 173); ratio
of length of fastigium to the minimum distance between eyes taken dorsally less than 2;
smaller species: body length 20-23 mm in males, 25-32 mm in females . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Quilta mitrata Stal
93(91') Apical half of hind tibiae expanded; tegmina and hindwings not abbreviated; general
coloration: green with brownish to bluish band latero-apically [Oxya] . . . . . . . . . . . .94
93' Apical 1/2 of red hind tibiae with black spine tips expanded; head conical; pentagonal
fastigium wider than long without median keel-like ridge; length of head and pronotum
as long as antennae; median pronotal carina weak, weakly intersected by 3 transverse
sulci; tegmina and hindwings cut short before tip of abdomen (Fig. 174) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caryanda diminuta (Walker)
94(93) Supra-anal plate feebly trilobate (Fig. 175 as in O. hyla intricata) with a tubercle on each
side of a median apical structure; cerci conical (Fig. 176), narrows apically to a small tip;
inner tooth-like pair of lophi (= lobes of epiphallus wanting; female subgenital plate
(Fig. 177) without longitudinal ridge ventrally . . . . . . . . . . . . . Oxya hyla intricata (Stal)
94' Lateral tubercles absent in the supra-anal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .95
95(94') Valve-like (valvular) plate of cingulum prominently long, upwardly curved and swollen
at apex (Fig. 178); subgenital plate (Fig. 179) of female with a median pair of widely
separated spines; cerci (Fig. 180) acute; supra-anal plate of male with a rather broad
triangular median lobe (Fig. 181); tegmen with a poorly developed bulge in the costa .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxya velox (Fabricius)
95' Valve-like plate short and fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .96
96(95') Supra-anal plate bears a lobe-like apical part that extends posteriorly when flat, basal
folds absent (Fig. 182); tegmina short not reaching 4th tergite of abdomen, cerci (Fig.
183) simple, conical with slightly acute tip; subgenital plate (Fig. 184) of females with
lateral longitudinal ridges in the posterior 1/2 of the ventral surface; small grasshoppers
barely 26 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxya minuta Carl
96' Supra-anal relatively flat with or without developed basilateral folds . . . . . . . . . . . .97
97(96') Supra-anal plate (Fig. 185) with well developed basilateral folds; cerci (Fig. 186) trun-
cate or apically subacute; lateral longitudinal ridges of the females’ subgenital plate
(Fig. 187) without spines except at tip . . . . . . . . . Oxya japonica japonica (Thunberg)
97' Supra-anal plate without basilateral folds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
98(97') Cerci conical (Fig. 188), taper towards subacute tip; supra-anal plate less triangular
without basilateral folds (Fig. 189); valve-like plate of cingulum broad; apical valves of
penis stout; in the females tegmina short not extended beyond apex of hind femora . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxya yezoensis Shiraki
98' Cerci with a blunt to rounded apex (Fig. 190); supra-anal plate (Fig. 191) rounded
triangular with basilateral folds; valvular plate of cingulum and apical valves of phallus
slender; tegmina of females beyond apices of hind femora . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxya chinensis (Thunberg)
99(90') Mesosternal lobes either acute or rectangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
99' Mesosternal lobes usually rounded; pronotum without any carinae . . . . . . . . . . . . 104
100(99) Pronotum, moderately coarse, strongly recurved (median keel raised) and distinctly cut
by 3 transverse furrows (Fig. 192); frontal ridge strongly grooved below ocelli with
parallel margins; tegmina and hindwings well developed, extended beyond hind knees,
hind tibiae purple red with 8-9 outer and 10-11 inner yellow spines with blackened tips;
coarsely punctured body, length 50-60 mm in males and 65-86 mm in females . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chondracris rosea (de Geer)
100' Pronotum rather flat to moderately raised . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
101(100') Hind femur long to moderately slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
101' Hind femur short and broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
102(101) Pronotum slightly tectiform to saddle-shaped (Fig. 193); posterior portion rounded
(Fig. 194); face not sloping (Fig. 195); fastigium trapezoidal; median keel low, inter-
sected by 3 prominent transverse sulci, 2nd and 3rd lateral longitudinal sulci with 2
transverse lines; gena without dark brown band; mesosternum rounded; tegmina with
numerous moderately broad longitudinal bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schistocerca americana gregaria (Forskal)
102' Pronotum roof-shaped, not tectiform, laterals with 2 transverse dark brown bands
bordered by pale yellow bands (Fig. 196); posterior part moderately acute; face sloping;
fastigium more or less rounded; gena with a dark brown band reaching bottom of eye;
mesosternum acute; tegmina (Fig. 197) without numerous brown. bands but with pale
yellowish median band on its flattened dorsal surface . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patanga succincta (Linnaeus)
103(101') Tegmina with numerous slightly diagonal bands (Fig. 198); dorsolateral sides of
pronotum with velvety dark brown bands (Fig. 199); prosternal process bent backwards
almost reaching mesosternum; face reclinate, frontal ridge narrow, parallel sided, and
partially depressed at ocellus; pronotum slightly constricted medially, median keel low
and distinct, intersected by 3 transverse sulci; posterior margin angular; tegmina and
hindwings well developed reaching beyond hind femora . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyrtacanthacris tatarica (Linnaeus)
103' Tegmina fairly broad without dark brown fasciae (Fig. 200); prosternal process only
slightly bent backwards; frontal ridge rather reclinate, surfaces finely punctated;
pronotum more or less tectiform, constricted along metazona; median keel thick,
convex but not well elevated, posterior margin distinctly pointed (Fig. 201); general
coloration yellowish brown with bluishblack markings . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valanga nigricornis (Burmeister)
104(99') Anterior part of pronotum slightly narrowed, posterior more or less rounded at tip;
frons slightly convex to moderately reclinate; combined length of pronotum and head
longer than antennae; prosternal lobes short and subconical; hind femora broad, 3-4
times as long as broad, dorsal portion with dark patches (Fig. 202); general coloration
dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catantops pinguis (Stal)
104' Not as above, pronotum constricted at midhalf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
l05(104') Prosternal tubercle conical, acute apically and moderately inclined towards the back;
antennae filiform, total length either longer or shorter than combined length of head
and pronotum; frons straight in profile; hind femora slender with 4 broad black spots
dorsally (Fig. 203); cerci of males short, gradually narrowed in the apical 1/3 and
incurved apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenocatantops humilis (Serville)
105’ Prosternal tubercle strongly broadened, flat laterally, and curved backwards; antennae
slightly longer than head and pronotum; frons sloping behind; pronotum constricted
medially; hind femora 4-5 times as long as broad; tegmina well beyond hind knee by
more than a pronotal length; hind femora (Fig. 204) with 3 black spots on the dorsal
margin connected to a midlongitudinal black band with a serrated appearance in the
lower area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenocatantops splendens (Thunberg)
106(89') Spine-like apex in the lower external lobe of hind femora absent . . . . . . . . . . . . . . 107
106' Spine-like apex in the lower external lobe of hind femora present (Fig. 205); pronotum
long, cylindrical and posteriorly expanded, with 3 transverse sulci, third very much
behind middle; short conical prosternal tubercle obtuse apically; a broad dark band
runs from lateral margins of fastigium to lateral lobes of pronotum to anterior margin of
tegmina; valves of ovipositor strongly toothed, apex with strong recurved tooth . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gesonula mundata (Walker)
107(106) Supra-anal plate broader than long, tips acutely angular; male cerci very large, trilobed
with the median piece greatly expanded (Fig. 206); subgenital plate of female also
trilobed, lateral lobes shorter and less broad than the slightly acute to rounded midlobe
(Fig. 207); pronotum with 2 narrow longitudinal black bands (Fig, 208); tegmina with
longitudinal black stripe running from pronotum to post-cubital vein; terminal ab-
dominal tergite black at the center edges (Fig. 209); anterior end of hind femora dark
black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parahieroglyphus bilineatus (I. Bolivar)
107' Supra-anal plate longer than broad, sides curved outwards toward apex; male cerci
medium-sized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
108(107') Apices of male cerci notched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

108' Apices of male cerci not notched, simple ................................... 110
109(108) Ventral side of males’ cerci straight in lateral view, tip acutely pointed with a blunt
inwardly curved tubercle in the apical 1/4 (Fig. 210); subgenital plate of females longer
than broad, semi-ovate with the median lobe pointed (Fig. 211); 2nd and 3rd sulci
slightly wavy; hind femora moderately slender, reaching end of abdomen . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hieroglyphus banian (Fabricius)
109' Ventral side of males’ cerci (Fig. 212) strongly recurved in lateral view, tip blunt and
curved downwards with a large tubercle above it; subgenital plate of female trilobed,
median lobe longer than lateral lobes (Fig. 213) fastigium 2-3 times as broad as long, all
sulci wavy (Fig. 214); apodeme (Fig. 215) widely U-shaped viewed from above; hind
femora not reaching end of abdomen . . . . . . . . . . . Hieroglyphus tonkinensis I. Bolivar
1l0(108') Pronotum rounded along the posterior margin .............................. 111
110' Pronotum shaped otherwise, usually obtuse-angular ......................... 112
111(110) Subgenital plate of males (Fig. 216) sulcated and truncated apically; cerci (Fig. 217)
simple, slightly C-shaped facing each other dorsally, tip blunt and upcurved laterally;
females’ subgenital plate 3-lobed with small round outer lobes much shorter than the
pointed midlobe (Fig. 218); posterior of pronotum rounded (Fig. 219); integument
smooth and shallowly pitted; tegmina and wings in line with tip of abdomen . . . . . . . . .
.................................................. Hieroglyphus oryzivorus Carl
111' Subgenital plate of males notched apically (Fig. 220); cerci (Fig. 221 nearly truncated at
tip with an upturned tubercle before apex (seen laterally); females’ subgenital plate
3-lobed with shorter, angular outer lobes, median lobe longer and subacute (Fig. 222);
tegmina and hindwings a little beyond tip of abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hieroglyphus daganensis Krauss
112(110') Cerci (Fig. 223) of male subacute along apices, prominently longer than supra-anal plate
with a subacute tip (Fig. 224); females’ subgenital plate wide before the subacute
median lobe, with two parallel slightly wavy ridges (Fig. 225); pronotum intersected by
3 black sulci; posterior sulcus deeper and wider (Fig. 226) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hieroglyphus annulicornis Shiraki
112' Cerci (Fig. 227) of males oblique, longer than supra-anal plate; subgenital plate (Fig.
228) of females with strongly rounded outer lobes and highly acute median lobe; 3 sulci
of pronotum interconnected near margins (Fig. 229); three distal abdominal sternites
hairy; tegmina and hindwings beyond tip of abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hieroglyphus nigrorepletus I. Bolivar
113(62') Small, usually slender insects with asymmetrical rasping and piercing mouthparts; wings
short and narrow, with reduced venation and wide marginal fringe; tarsi with reversible
apical bladders (last segment swollen); cerci absent; apex of abdomen rounded, point of
ovipositor dissected ventrally; narrow wings pointed apically; antennae 6- to 9-seg-

mented (THYSANOPTERA); thrips [Thripidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
113' Not with the above combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
114(113) Forewings (Fig. 230) with longitudinal veins, sometimes cross vein and setae on vein;
segment X of abdomen (Fig. 231) usually pointed (conical type), rarely tubular, always
divided longitudinally on the ventral surface; saw-like ovipositor present in females;
wing membrane with numerous microtrichia (Terebrantia) . . . . . . . . . . . . . . . . . . 115
114' Forewing without longitudinal veins if present; microtrichia in the wing membrane
absent; segment X of abdomen tubular (Fig. 232) in both sexes without a longitudinal
division dorsally and ventrally; females with a ventrally everted membranous ovipositor
between segment IX and X; head long and parallel-sided; prothorax with 8 long pointed
anteroangular and posteroangular setae (Fig. 233); antennae 8-segmented, brownish
except lighted yellow band 3rd segment (Fig. 234); segment III with 0-2 sense cones and
4 cones on IV; postocular seta well developed, long, about 50µ; (Tubulifera) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplothrips aculeatus (F.)
114" Allied to H. aculeatus; prothorax with 10 long cylindrical setae (Fig. 235), cup-like at tip;
pelta inverted T-shaped; antennae brownish in basal 2 segments, VII-VIII and apical 1/2
of VI, III-IV-V and basal 1/2 of VI yellow; segment X of abdomen 1/2 the width of
segment VII in females . . . . . . . . . . . . . . . . . . . . . . . . Haplothrips ganglbaueri Schmutz
115(114) Forewing broad (Fig. 236), apically rounded with 4 cross veins distinct, 2 dark cross
bands distinctly separated from pale areas; short setae on head and pronotum (Fig.
237); antennae 9-segmented, segment III apically shaded, sensoria on segments III and
IV linear (Fig. 238); prothorax without setae post-marginally; metanota indented
laterally; anteocellar setae absent; maxillary palpi 3-segmented . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aeolothrips fasciatus (Linnaeus)
115' Forewing slender and acute apically, visible cross vein only 1; antennae with 6-9 seg-
ments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
116(115') Head much smaller than pronotum, distinctly produced in front of eyes (Fig. 239);
antenna segment II produced exteriorly at apex (Fig. 240); asymmetrical, III and IV with
sense cones simple; trapezoidal pronotum bears 2 pairs of major posteroangular setae;
sternites III-VIII of abdomen each with an ovoid glandular area in males (Fig. 241) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chirothrips manicatus (Haliday)
116' Head not produced in front of eyes; segment II of antennae normal . . . . . . . . . . . 117
117(116') Antennae (Fig. 242) 7-segmented with forked sense cone on segments III and IV; head
with 1 pair of anteocellar setae (Fig. 243); postocular seta III longer than I; forewings
uniformly (Fig. 244) dark greyish brown; tibiae brown . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenchaetothrips biformis (Bagnall)
117' Antennae 8- or 9-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Figs. 230-234
118(117') Pronotum (Fig. 245) normal with short setae at posterior margin; metanota with a long
lateral arm and 4 setae; vein series of setae in the forewing interrupted (Fig. 246);
antennae (Fig. 247) 9-segmented with simple sense cone in segment 1 and a forked sense
cone in III; diameter of spiracle on tergite VIII about 0.2 times the length of tergite;
body yellowish green with some light brown spots... Anaphothrips obscurus (Muller)
118' Pronotum (Fig. 248) with long setae at the posterior margin; prothorax with 2 pairs each
of long setae on anterior and posterior margins, anteromarginal setae shorter than
antero-angular setae; interocellar setae in ocellar triangle; forked sense cones in anten-
nal segments III and IV (Fig. 249); fringe cilia wavy; body dark yellowish brown; wings
light yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frankliniella tenuicornis (Uzel)
119(113') Both pairs of membranous wings largely covered with overlapping scales; mouthparts in
a coiled proboscis; antennae many segmented; moths, butterflies (LEPIDOPTERA)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . 120
119' Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
120(119) Antennae gradually or abruptly clubbed, tip sometimes hooked .............. 121
120' Antennae not clubbed or hooked . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . 133
121(120) Antennae approximated at base, clubbed distally without hooked tip; some peripheral
veins stalked; front wings more or less triangular, some veins especially Sc swollen at
base, radius 5-branched; moths usually with brownish or greyish circular spots
[Satyridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
121' Antennae widely separated at base, scape with scale-tuft, flagellum dilated apically with
hooked tip; ocelli absent; chaetosemata present; haustellum naked; forewings without
retinaculum, veins arising separately from discal cell, CuP absent; 1A + 2A simple or
forming basal fork; hindwings without frenulum, Sc connected to Rs near base by R1;
moths usually with glass-windows on wings [Hesperiidae] . . . . . . . . . . . . . . . . . . . . 125
122(121) Eyes hairy; all veins basally swollen, termen of both wings entire, evenly rounded;
undersurfaces of wings brighter than the dull brown uppersides (each wing) with a pale
postdiscal band and a series of submarginal ocelli (eye spots) (Mycalesis) . . . . . . 123
122' Eyes smooth; tarsal claws on the mid and hind legs cleft; forewings falcate at vein 5;
hindwings caudate at vein 4; uppersides of hindwings with a white dot or a white
pupilled ocellus (Melanitis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
123(122) Forewings (Fig. 250) with a pair of eye spots (swollen one along veins 7-9 and the large
spot almost along veins 3-5) dorsally, though a much smaller spot may be present and
contiguous to the upper one, similar underneath except for the upper part bordered
with a spot each anteriorly and posteriorly; hindwing bare dorsally, underneath bears
6-7 eye spots with 5 and 1 broader than rest (Fig. 251) . . . . . Mycalesis gotanma Moore
123' Uppersides of forewings with only 1 large spot, ringed pale yellow-brown between veins
2 and 3 (Fig. 252); undersides with 2 minute white dots, one under the large dorsal spot
and the other between veins 6 and 7; hindwings bare dorsally, underneath with a small
dark brown spot each between veins 1a to vein 7 . . . Mycalesis nr. mineus (Linnaeus)
124(122') Uppersides of forewings dark brown with two 1-2 mm subglobular white spots between
veins 3-4 and 4-5 ringed brown to bluish violet with 4 highly wavy orange hues in the
centers, a broad brown spot with violet tinge dorsad of 2 white spots (Fig. 253);
undersides of forewings with 2 white spots of the upperside; hindwings with 2 brown
spots ringed light brown to yellow and centered white between veins 6 and 7, and 2 and
3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanitis leda (Linnaeus)
124' Upperside blackish brown and with unmarked forewings in males, light to pale brown
in females, apical area paler than basal in both sexes; forewings of females with a round
to ovoid white spot each between veins 3-4 and 4-5, lower 1/2 bordered with orange band
in the inside, veins 5-7 with an inner orange and outer dark brown band; hindwing with
a small white spot between veins 2-3 dorsally (Fig. 254) . . . . . Melanitis phedima Stoll
125(121') Vein 5 (M2) of forewings straight from its point of origin; vein 5 of hindwings distinct;
antennal club not constricted before the apiculus; cell of hindwings about 2/5 wing
length; uppersides of both wings with prominent orange-yellow markings (Fig. 255), 3
bands along postmedian and a contiguous one along costa and median, broadening at
median; hindwing with 3 nearly contiguous bands starting from vein 2 upwards . . . . . .
..................................................... Ampittia maro Fabricius
125' Vein 5 (M2) downcurved from its origin; vein 5 of hindwings wanting; antennal club
constricted before apiculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
126(125') Midtibiae smooth; wings with yellow markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

126' Midtibiae spined or smooth; wings brown with small hyaline spots . . . . . . . . . . . . . 128
127(126) Third segment of palpi short and stout; vein 2 of forewings nearer the end of the cell than
to their base; uppersides of male and female forewings orange-yellow with brown
markings (Fig. 256), male forewing with a narrow oblique band from midvein 1b to the
origin of vein 4; termen of forewing straight; post-discal band extends from vein 1b to
vein 8 and an orange spot in the cell in hindwings of male; ventral edge of male valva
without a projecting shoulder . . . . . . . . . . . . . . . . . . . . . . . . . Telicota augias (Linnaeus)
127' Third segment of palpi long and thin; antennal club not flattened with a very fine
apiculus and at an angle to the club; undersides of forewings with the lower basal portion
of vein 12 (Sc) fringed with hair scales; vein 2 (Cu2) closer to end of cell, vein 4 farther
from vein 3 than vein 5; post-discal band on uppersides of forewings broad running from
dorsum to the costa, hindwings with 6 orange spots (Fig. 257) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Potanthus confucius (Fruhstorfer)
128(126') Midtibiae heavily spined; undersides of hindwings with a spot in the cell (Pelopidas) . .
........................................................................ 129
128' Midtibiae smooth and not spined ........................................ 130
129(128) Band on uppersides of forewings meet vein 1b on the basal side of the origin of vein 2 in
males, with 4 small spherical post-discal white spots and 2 rather elongate ones in the
lower apical edge of the discal cell; females with broader white spots, 4 post-discals and
4 more below the apical end of the discal cell; 2 discal cells perpendicular to the nearly
triangular spot above vein 1b (Fig. 258) . . . . . . . . . . . . . Pelopidas mathias (Fabricius)
129' Band on the uppersides of forewings meets vein 1b right below origin of vein 2 or nearer
termen in males (Fig. 259); spots similar to P. mathias except for a more parallel discal
spot and less oblique band between veins 3-4; females (Fig. 260) as in above species
except for a more shoe-like band between veins 3-4, more rounded spot above vein 1b
not perpendicular to the 2 discal cells; moths larger than P. mtathias . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pelopidas agna (Moore)
130(128') Antennae about 1/2 length of each forewing, antennal club obtuse, uppersides of
hindwings with 4 elongate to ovoid white spots; forewings with 2 elongated spots in the
discal cell, 2 groups of post-discal (3 small spots lining the radial spaces and 3 broad,
more rectangular spots below it, and 1 spot above vein 1b (Fig. 261) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polytremis pellucida (Murray)
130' Antennae very short, approximately 2/5 length of forewing; apiculus very short . . 131
131(130') Discal cells with 2 thin parallel white spots; 3 small post-discals and 2 broader bands in
the lower apical end of cell, the 7 spots C-shaped; hindwing with 4 white spots (Fig. 262);
male valva uniform in width with a straight inner edge . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parnara guttata Bremer and Grey
131' Discal cells without prominent spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
132(131') Upperside of hindwings with 2 small, irregular white postdiscal spots faint and diffuse
(Fig. 263); inner margin of valva strongly convex . . . . . . Parnara naso bada (Moore)
132' Hindwings with 4 white spots (Fig. 264); forewings similar to P naso; uppersides of wings
with a more golden luster than P. naso; male valva recurved in its inner margin below
the projecting shoulder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parnara ganga Evans
133(120') Maxillary palpi 4-segmented (sometimes 2-3); haustellum reduced; chaetosemata

usually present; forewings without areole; Sc + R1 approximated to or shortly fused to
Rs beyond discal cell, M2 approximated to M3 at base in the hindwing [Pyralidae] . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
133' Maxillary palpi 1-segmented; haustellum usually strong or reduced; forewings with or
without areole; chaetosemata absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
134(133) Wings immaculate white with brown specks on the uppersides of both wings (Fig. 265);
larvae semiaquatic, with tracheal gills and concealed inside cylindrical leaf tube;
caseworm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nymphula depunctalis (Guenée)
134' Wings never immaculate, usually brown to brownish orange; grass webworms or leaf-
folders/leafrollers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
134'' Wing coloration not as above; stem borers ................................ 139
135(134') Wings fuscous (dark brown) with postmedial line undulated in both (fore and hind
wings); forewings with the antemedial line outwardly curved and oblique; medial very
short and dorsad of the inwardly lower half of postmedial (Fig. 266); proboscis strongly
coiled in between porrect labial palpi (Fig. 267); body uniformly dark brown; body
length about 12.5 mm, wing span 22-25 mm; grass webworm . . . . . . . . . . . . . . . . . . . . . . .
........................................... Herpetogramma licarsisalis (Walker)
135' Wings orange-brown with dark brown subterminal to terminal areas; antemedial,
medial, and postmedial lines more prominent; body length 6-9 mm; wing span 11-18
mm; larvae yellow, with or without brown spots in the pronotal shields and sclerotized
base of setae (leaffolders/leafrollers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
136(135') Forewings with veins R2 + R1 stalked; costal region without any strigulae; postmedial
line of both wings diagonal and straight and distinctly interconnected (Fig. 268); an-
temedial short and replaced with androconial hair tuft in the males (Fig. 269); tip of
abdomen with a longitudinal black band, broader in females and slender in males;
genitalia of both sexes illustrated in insets . . . . . . Cnaphalocrocis medinalis (Guenée)
136' Forewings with veins R2 and R1 free; costal region with or without strigulae; postmedial
line not as above; genitalia of both sexes illustrated in insets (Figs. 270-275) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marasmia . . . . . 137
137(136') Costa of forewings with silver-yellow and minute black strigulae (Figs. 270-271); sac-
culus with a spur-like structure; cornuti with about 60 teeth; valva elongated to ovate;
saccus knob-like; bursa copulatrix with minute teeth or scobinations in the proximal
duct (ductus bursae) and none in the distal sac (corpus bursae) as illustrated in inset
(Fig. 270) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marasmia ruralis (Walker)
137' Costa without strigulae ................................................. 138
138(137') Wing span 12-14 mm; forewings with a greyish brown C-shaped band covering the
subterminal and terminal areas; hindwings with 2 parallel lines traversing the wing (Figs.
272-273); aedeagus with 2 robust teeth; apical one twice as long as the basal; bursa
copulatrix broadly truncate basally and surrounded with composite-like granulations
along the signum illustrated in inset (Fig. 272) . . . . . . . . . . . Marasmia exigua (Butler)
138' Wing span more than 15 mm; postmedial line bent inwards to a point near the posterior
tip of a short medial line where it runs posteriorly; postmedial line of hindwings about
1/2 the length of antemedial posteriorly (Fig. 274); abdomen of male with a small
interconnected black band an penultimate segment while the female has short lon-
gitudinal black band on the tip (Figs. 274-275); genitalia of both sexes illustrated in
insets ............................................. Marasmia patnalis Bradley
139(134'') Proboscis absent (Fig. 276); hindwings with median nerve not pectinate on upper side
........................................................................ 140
139' Proboscis present (Fig. 277); hindwings with a pectinate median nerve on upper side.
........................................................................ 146
140(139) Palpi upturned (Fig. 278); forewings immaculate white, veins R2 to R4 stalked; males
have 24-45 mm wing span and white anal tuft; females 19-34 mm and with orange brown
tuft; American species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rupela albinella Cramer
140' Palpi porrect (Figs. 276-277); R2 usually free ............................... 141
141(140') Forewings yellow with a black spot at lower angle of cell in the females (Fig. 279) and
ochreous with minute markings in males (Fig. 280); frenulum double-bristled . . . . . . . .
............................................... Scirpophaga incertulas (Walker)
141' Forewings white to ochreous white; frenulum with 1-2 bristles; anal tuft ochreous yellow,
white or greyish ........................................................ 142
142(141') Frenulum double-bristled (Fig. 281). ...................................... 143
142' Frenulum single - bristled (Fig. 282) ....................................... 144
143(142) Both sexes immaculate white; anal tuft white (Fig. 283); length of labial palp 2 times
diameter of compound eyes in males; subteguminal process with a spine (Fig. 284) . . .
................................................ Scirpophaga innotata (Walker)
143' Females white; anal tuft ochreous yellow (Fig. 285); males ochreous; four fuscous spots
on the submedian fold and lower angle of cell and series of small fuscous dots on the
terminal area of forewings; subteguminal process large, without spine but with sinous
margin (Fig. 286); 3 cornuti (Fig. 287) . . . . . . . . . . . . . Scirpophaga nivella (Fabricius)
144(142') Length of labial palps as long as diameter of compound eyes (Fig. 288); forewings white
to ochreous white; subteguminal process rounded (Fig. 289); cornuti without spine (Fig.
290) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scirpophaga virginia Schultze
144' Length of labial palps longer than diameter of compound eyes (Fig. 291); subteguminal
process not as above .................................................... 145
145(144') Subteguminal process flattened and rounded with smooth margins (Fig. 292); moths
dark ochreous in males and white in females; length of labial palps 1.5 times diameter of
compound eye; anal tuft dark grey; cornuti bear 3 spines (Fig. 293) . . . . . . . . . . . . . . . . .
................................................. Scirpophaga gilviberbis Zeller
145' Subteguminal process bilobed (Fig. 294); moths of both sexes white; length of labial
palps 1.3 times diameter of compound eye; anal tuft pale ochreous white; cornuti
without spines (Fig. 295) . . . . . . . . . . . . . . . . . . . . . . Scirpophaga occidentella (Walker)
146(139') Forewings without vein R5 (or vein 7); (Phycitinae) . . . . . . . . . . . . . . . . . . . . . . . . . 147
146' Forewings with vein R5(7) (Crambinae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
147(146) Proboscis well developed; forewings with black suffusions along termen and tornus
curved downwards (Fig. 296); fore and hindwings with M3 (vein 4) present; American
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elasmopalpus lignosellus (Zeller)
147' Proboscis weak as in Fig. 277; forewings with a dark brown band parallel to the costal
margin; tornus rounded upwards (Fig. 297); fore and hindwings without M3; median
nervure with only 3 veins; African species . . . . . . . . . Maliarpha separatella (Ragonot)
148 Forewings rather narrow without metallic scales (Fig. 298); frons subrounded, slightly
forward but without a corneous point at apex; ocelli absent; tufts of hair of 2nd
abdominal segment and hind tibiae in males prominent; American species . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diatraea saccharalis (Fabricius)
148' Forewings somewhat broad with metallic scales (Fig. 299); frons conically produced
forward (Fig. 300) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
149(148') Hindwings with vein M1 (or vein 6) weakly developed and slightly above middle of
discocellulars; forewings with R5 (or vein 7) stalked with R4 (vein 8-9), subterminal area
with wavy light brown and silvery white bands, terminal area with series of black spots
and indented (Fig. 301) . . . . . . . . . . . . . . . . . . . . . . . . Acigona chrysographella (Kollar)
149' Hindwings not as above, vein M1 well developed and R5 free . . . . . . . . . . . . . . . . 150
150(149') R1 or vein 11 free (Fig. 302) ............................................. 151
150' R1+ Sc (or vein 12) fused (Fig. 303) ...................................... 155
151(150) Face broadly rounded without corneous point and ventral ridge (Fig. 304); forewings
with fuscous dots on the subterminal areas (Fig. 305); labial palps 3-4 times as long as
diameter of compound eyes; ocelli well developed; aedeagus curved without ventral
arm, but with bulbose basal projection (Fig. 306); costa of harpe with a tooth projected
upward, juxta1 arm long and tapers to points and without subbasal teeth (Fig. 307);
African and European species . . . . . . . . . . . . . . . . . . . . . Chilo agamemnon Bleszynski
151' Face conical with distinct point; ventral ridge may be present or absent . . . . . . . . 152
152(151') Ventral ridge absent (Fig. 308); forewings dull yellow dusted with brown scales, terminal
dots prominent, subterminal lines with series of lustrous metallic golden scales; labial
palps 4 times as long as diameter of eye; costa of harpe without teeth, arms of juxta long
and each bears a subapical tooth (Fig. 309); aedeagus distinctly long, hairy, apically
pointed, ventral arm absent (Fig. 310) . . . . . . . . . . . . . . . . . . Chilo plejadellus Zincken
152' Ventral ridge present; labial palps 3-3.5 times as long as diameter of eye . . . . . . . 153
153( 152') Face with vestigial ridge (Fig. 311); forewings yellow to brown dusted with fuscous
scales; subterminal line a light brown line (Fig. 312); vein M1 of hindwing arises from Rs
before angle of cell; costal margin of harpe with a strong median tooth; juxta bell-shaped
with 2 notches basally (Fig. 313) . . . . . . . . . . . . . . . . . . . . . . . Chilo partellus (Swinhoe)
153' Face not as above; vestigial ridge well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
154(153') Arms of juxta-plate symmetrical and distinctly swollen near apices; tips of harpe not
tapering (Fig. 314); subapical teeth absent (male genitalia); female genitalia with heavily
sclerotized ostial pouch and slightly demarcated from ductus bursae; frons with
prominent ridge and corneous point (Fig. 315); forewing length 11-14 mm (Fig. 316) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chilo suppressalis (Walker)
154' Arms of juxta-plate narrow and not dilated medially; subapical teeth present (Fig. 317);.
female genitalia with small ostial pouch and well demarcated from ductus bursae; larger
moths with forewing length 12-17 mm . . . . . . . . . . . . . . . . . . . . Chilo hyrax Bleszynski
155(150') Face produced forward, smooth, or with a small point; Oriental region . . . . . . . . 156
155' Face rounded without corneous point (Fig. 304) . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
156(155) Terminal dots ill-defined to absent (Fig. 318); forewing with subterminal metallic spots
roundly curved almost parallel to termen from vein M2 to inner margin; aedeagus
without pointed projection (Fig. 319); costa of harpe with tooth, harpe broad and more
or less triangular (Fig. 320) . . . . . . . . . . . . . . . . . . . . . . . . Chilo polychrysus (Meyrick)
156' Terminal dots distinct (Fig. 321); forewings with subterminal metallic spots regularly
curved and not parallel to termen; aedeagus with projection (Fig. 322); costa of harpe
without tooth, harpe not broad (Fig. 323) . . . . . . . . . . . . . . . Chilo auricilius Dudgeon
157( 155') Aedeagus with basal part curved and bears a short ventral arm (Fig. 324); arms of juxta
with subapical tooth; harpe broad and blunt apically (Fig. 325); forewing length 8-13 mm
and generally orange yellow to dirty yellow . . . . . . . Chilo diffusilineus (J. de Joannis)
157' Aedeagus without ventral arm (Fig. 326); juxta1 arm without tooth, harpe tapers apically
(Fig. 327); forewing length 10-14 mm and ochreous yellow . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chilo zacconius Bleszynski
158( 133') Haustellum reduced; forewing without areole; Sc + R1 swollen at base, fused with Rs to
near middle of discal cell then diverges, M2 nearer to M3 than to M1 in the hindwing;
dorsal black band of thorax tapers on both ends; abdomen pinkish red with black
transverse median band dorsally; lateral sides with 2 longitudinal rows of paired black
spots; forewings brownish white with a broad transverse black band at center, obliquely
interrupted postmedially; hindwings white with a pair of gray spots in terminal area (Fig.
328) (Arctiidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Creatonotus gangis (Linnaeus)
158’ Haustellum strong; forewings with areole; Sc + R1 shortly fused with Rs near base,
rarely fused to about 1/2 of discal cell, M2 weak arising closer to Ml than to M3 (trifid)
or strong and arising nearer to M3 than to M1 (quadrifid) in the hindwing [Noctuidael
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
159(158') Defoliators; moth body length 9-11 mm; wing span 16-22 mm; whitish brown to dark
yellow or orange brown with or without diagonal bands . . . . . . . . . . . . . . . . . . . . . 160
159' Defoliators; moth body length greater than 15 mm; body and wing coloration variable
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
159'' Stem borers; moth body length 10-17 mm; forewings usually with suffused red markings
radiating from wing base towards discal area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
160(159) Triangular-shapedmoths and head always directed downwards at rest (Fig. 329); wings
dull whitish yellow with a small brown short triangular-like postmedial band, subter-
minal to terminal area brownish with 8 tiny terminal spots; labial palps slightly porrect
and broadened by scales apically (Fig. 330); frons pointed; larvae green with a pair of
white longitudinal stripes dorsally, body with long hairs . . . Rivula atimeta (Swinhoe)
160' Moths not triangular at rest; wings orange to yellow-brown with a pair of parallel,
diagonal ferruginous bands (Fig. 331); labial palps curved, hind segment small (Fig.
332); larvae green like Rivula but without long hairs; prolegs only present in abdominal
segments V, VI, and anal region; larvae make looping movement . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Naranga aenescens Moore
161(159') Antemedial line of forewings thin, entire and prominent, basally curved costally and
continued posteriorly in the hindwing as a white diagonal band projected towards tip of
abdomen; postmedial broad and basally curved; top subterminal with a broad brown
spot contiguous downwards by a thinner band; hindwings dark bluish violet with 3-4
white spots in the termen (Fig. 333); labial palps upcurved, above eyes and in line with
antennal bases; moth body length 16-22 mm, wing span 50 mm . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Achaea janata (Linnaeus)
161' Antemedial line of forewings absent or less prominent; hindwings never bluish violet
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
162(161') Wings brown to grey-brown; darker in the forewings than the hindwings; forewings with
a slant L-shaped band formed by an elongated reddish brown band below discal cell,
above midlength of vein 1A + 2A and 1 more diagonal band radiating from apex to outer
1/3 of posterior margin (Fig. 334); subterminal with about 7 minute black dots; reniform
spot traceable; hindlegs covered with a thick hair tuft . . . . Mocis frugalis (Fabricius)
162' Medium-sized moths without a slant L-shaped band; abdominal prolegs of larvae
complete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
163(162') Pronotal hair tuft with a pair of recurved black eyelash-like bands (Fig. 335); antennae
of males bipectinate, branches of moderate length (Fig. 336); forewings with a distinct,
short, transverse black bar extending distad from reniform spot; pale orbicular transver-
sely ovate, narrowed distally (Fig. 337); hindwings nearly white except apex and part of
termen; larvae are cutworms. . . . . . . . . . . . . . . . . . . . . . . . . . Agrotis ipsilon (Hufnagel)
163' Pronotal hair tuft without recurved black eyelash bands, if present, antennae of males
not bipectinate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.64
164(163') Valves of male genitalia 5-7 times longer than broad; vesica long, coiled, and barbed
with spines; appendix bursae of females’ genitalia with a long curving “tail” at the apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
164' Valves broad, not as slender as in Heliothis; vesica different . . . . . . . . . . . . . . . . . . . 166
165(164) Hindwings black terminal fascia interrupted by a pale patch between veins M3 + Cu2;
postmedial and subterminal lines discernible (Fig. 338); apex of vesica with a normal
coil valves 5 times as long as wide in males (Fig. 339); females’ lumen surface of
appendix bursae clothed with spicules; larval coloration varies from green to brown with
whitish short stripes; cremaster spines basally separated by twice basal diameter of
spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heliothis (Helicoverpa) armigera (Hubner)
165' Hindwings black terminal fascia not interrupted; postmedial and subterminal lines
wanting (Fig. 340); apex of vesica curved tail-like; valves 6-7 times as long as broad in
males (Fig. 341); spicules on lumen surface short and compactly set; small signum of
bursae dorsad to 3 elongate signa; larval color as above; cremaster spines closer to each
other, separated by a basal diameter of spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
................................... Heliothis (Helicoverpa) punctigera Wallengren
166(164') Eyes not hairy; indentation between valvula and cucullus small or wanting (Spodoptera)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
166' Eyes hairy; indentation between valvula and cucullus prominent to very wide (Mythim-
na) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
167(166) Reinform spot oblique and arrow head-like; orbicular oblique with black edges and
dark brown center; subbasal line angled and oblique; antemedial slightly waved and
curved inwards toward the base in the lower half, subterminal with 7 transverse bIack
bars, 2nd bar from top the smallest and basal one bifurcated or split into 2 black bars
(Fig. 342); moths with dark purplish brown forewing. . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spodoptera litura (Fabricius)
167' Wing patterns or marks not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
168(167') Western Hemisphere species; females’ forewings dark brown without any contrasting
markings, superficially rounded at apex with an ovate, oblique and black centered
orbicular spot; males’ usually brown with subterminal line of forewing fading out
midway to inner margin; harpe short and straight; valves broad about 1/4 longer than
wide (Fig. 343); ductus bursae long and cylindrical (Fig. 344) . . . . . . . . . . . . . . . . . .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . Spodoptera frugiperda (J.E. Smith)
168' Not restricted to the Western Hemisphere; wing patterns different . . . . . . . . . . . . 169
169(168') Indentation between cucullus and valvula absent in male’s genitalia . . . . . . . . . . . 170
169' Indentation present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
170(169) Central margin of harpe broad with strong curved processes on distal and proximal
ends, distal process slightly beyond margin of valvula; valva half as broad at the valvular
region; sacculus and transtilla broad (Fig. 345); females’ corpus bursa a globular sac;
heavily sclerotized ductus bursa ribbed with minute sclerotized tubercles; signum
scobinate and transverse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spodoptera pecten Guenée
170' Central margin of harpe slender, curved distal process not reaching margin of valvula;
transtilla not clear (Fig. 346); males’ forewings without pale diagonal band running from
near costa between orbicular and reniform spots toward inner apical angle of wing;
apical tibial spurs of middle legs very slender with reduced specialized setose (Fig. 347);
females with circular or shortly ovate orbicular spot, never oblique and smaller than the
reniform; larvae with pale spiracles rimmed black; dorsum pale with a narrow medial
dark stripe enclosing a broken pale middorsal line . . . . . Spodoptera exigua (Hubner)
171(169') Indentation broad and shallow between the broadly rounded cucullus and valvula;
transtilla elongate and thumb-like pedunculi very prominent with thick fan-like hair tuft;
valvula clothed with a dense patch of hairs mid-dorsally; harpe with a strongly
sclerotized apical spine, distal process curved and 2 proximal processes on either side
of it present (Fig. 348); aedeagus with a long sclerotized spine distally; ductus bursa not
sclerotized but ribbed longitudinally . . . . . . . . . . . . . . . . Spodoptera abyssinia Guenée
171' Indentation deep to moderately deep; other characters not as above; reniform spot
prominently black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
172(171') Ventral margin of males’ genitalial valve incised near apex (Fig. 349); cucullus twice as
broad as long; distal and proximal processes of harpe sharply pointed and hardly
curved, tip of distal process slightly beyond incision line (Fig. 350); aedeagus with a
single cornutus, smooth and tapered apically, and about 1/2 as long as aedeagus;
sclerotized ductus bursa 2 times longer than ostium bursa; reniform spot prominently
black as in succeeding species; in both sexes, the band between postmedial and subter-
minal fasciae narrow, reddish brown shading between veins M2 and Cul reduced (Fig.
351) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spodoptera triturata (Walker)
172' Ventral margin of males’ genitalial valve incised well below apex; cucullus as long as
broad or a little longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
173(172') Foretibiae with a thick mass of long hair and scales on outer side partly reaching apex
of tarsus; outer spur of tibiae II slightly shorter than inner spur in males; basal 1/2 of
inner spur with specialized, non-squamose densely setose area (Fig. 352); male
genitalial valves elongate and narrow except in mid-valvula, the broadest (Fig. 353); anal
angle in the indentation present between cucullus and valvula; transtilla slender; small
finger-like labis present; harpe bears a finger-like apical spine reaching anal angle . . .
....................................................................... 174
173 ' Foretibial hairs and scales lightly produced and never up to 1st tarsal segment; both
spurs on tibiae II very broad and lanceolate, specialized setose areas broad and well
developed (Fig. 354); females’ forewings with elongate-ovate, strongly oblique orbicular
spot, reniform spot wanting (Fig. 356); males’ forewings with a pale diagonal band
between orbicular and reniform spots running from near costa to inner apical wing
margin, genitalia1 valves narrowed towards saccus, cucullus blunt to squarish on top,
harpe curved from the central region (L-shaped), distal end acutely pointed and
downwardly curved at apex (Fig. 355). . . . . . . . . . . . . . . Spodoptera exempta (Walker)
174(173) Males: Medial band reduced to small spot between the reniform and the orbicular,
terminal area a small patch near apex; females: white subterminal fascia very slender
and terminal area marked with reddish brown irrorations (Fig. 357); Oriental, Indo-
Australian and Pacific species . . . . . . . . . . Spodoptera mauritia acronyctoides Guenée
174' Males: medial with a conspicuous white band, space between subcostal vein and Cu2
white except the brown margins and central reddish area of the orbicular spot; and a
large fuscous black reniform spot joined to brown costa by a vague median band; veins
M1 to M3 with a small but conspicuous fuscous spot; terminal area immaculate white at
apex; terminal row with interneural fuscous spots (Fig. 358); African species . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spodoptera mauritia mauritia Boisduval
175(166') Moths pale brown with slight traces of orbicular and reniform spots (Fig. 359);
postmedial curved represented by a series of black spots on veins, excurved from radial
branches to vein 5 and oblique from vein 4 to outer margin; and marginal series of black
specks fairly prominent; light brown oblique band radiates from apical area to bottom
of forewings, hindwings pale suffused with reddish brown tinge; cucullus and valvula
with a wide anal angle, the former with a long spur apically and moderately slender base
opposite anal angle (Fig. 360); female genitalia with a long less rigid arm encircling the
bursa copulatrix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mythimna separata (Walker)
175’ Moths brown ochreous or reddish brown without traces of paired orbicular and
reniform spots; apex of discal with a whitish spot on entire median of cell bisected by a
whitish stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
176(175') Discal cell bisected by a whitish stripe; forewings slight reddish brown with lustre,
postmedial line with a series of reddish black spots on veins 2 and 3 running inward-up-
ward to costal margin; antemedial line a reddish black spot below fork of vein 2 (Fig.
361); meson of thorax with whitish stripe on a reddish brown background; hindwings
pure white; cucullus sausage-like, more than 4 times as long as wide; sacculus not
prolonged to midpoint of cucullus (Fig. 362) . . . . . . . . . . . Mythimna venalba (Moore)
176' Discal cell with only a whitish spot on tip; cucullus kidney-shaped to large and normal
........................................................................ 177
177(176') Moths brownish-ochreous mottled with fuscous scales; males with prominent ventral
lateral tuft on the base of abdomen; females as in males with hair fringes underneath
discal cell; wing veins whitish speckled with grayish brown scales; reniform spot, a small
white spot at lower angle of cell; antemedial line only a single blackish brown spot below
median vein; postmedial with a series of concolorous spots on veins evenly excurved
from costal margin to inner margin (Fig. 363); sacculus with a horn-like projection,
outcurved, somewhat slender and pointed; cucullus large and normal (Fig. 364) . . . . . .
................................................. Mythimna loreyi (Duponchel)
177' Moths ochreous ground; apex of forewings rounded, veins white and their interspaces
with fine red-brown streaks; median vein more prominent at lower cell angle beyond a
slight fuscous fascia; undersides of wings both without series of specks; inner margins of
saccullus toothed, highly sclerotized; cucullus kidney-like shaped (Fig. 365) . . . . . . . . .
................................................. Mythimna roseilinia (Walker)
178(159') Asian species; forewing with a pale red horizontal band radiating from wing base to mid
termen (Fig. 366); male clasper with a strong furcated projection . . . . . . . . . . . . . . . . . .
.................................................... Sesamia inferens (Walker)
178' African species; forewings without the radiating pale red band (Fig. 367) and male
claspers not as above; melanism notable in adult moths . . . . . . . . . . . . . . . . . . . . . . 179
179(178') Juxta broad trapezoid (Fig. 368), costal spine short and bifid apically (Fig. 369); fore-
wing more ochreous with minimal fuscous suffusion, fringe ochraceous white and lightly
infuscate at middle and tip; postmedial fascia fuscous black and slightly dentated;
pectus pinkish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sesamia calamistis Hampson
179' Juxta flask-shaped with a long neck (Fig. 370), costal spine long, curved with a strong
sub-apical tooth (Fig. 371); forewing with more fuscous suffusion; postmedial fascia
curvely dentated extending from vein R2 (10) to Culb (2); pectus white . . . . . . . . . . . . .
......................................... Sesamia botanephaga Tams & Bowden
180(119') Beak arising from front or hind part of head; front wings thickened at base and
membranous at tips (hemelytra), tips overlapping at rest (HEMIPTERA) . . . . . . . 181
180' Mouthparts not beak-like, usually mandibulate ............................. 344
181(180) Forewing in the form of tegmina of more or less uniform texture, without sharp differen-
tiation into corium and membrane and usually held roofwise over abdomen; labium
inserted close to prosternum without an intervening sclerotized gula . . . . . . . . . . . 182
181' Forewing in the form of hemelytra with thickened corium and apical membrane, usually
folding flat on the abdomen with widely overlapping apices; labium inserted far from
prosternum with an intervening sclerotized gula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
182(181) Tarsi 1- or 2-segmented; abdomen with or without siphunculi . . . . . . . . . . . . . . . . . 183
182' Tarsi 3-segmented; abdomen never with siphunculi . . . . . . . . . . . . . . . . . . . . . . . . . 194

183(182) Tarsi, when present, usually 1-segmented and bearing a single claw; with a combination
of dorsal ostioles, ventral circuli, triocular pores and usually with 1-18 pairs of cerarii;
anal ring usually with an outer and inner row of pores; tubular ducts not invaginated at
inner end; siphunculi absent; all forms wingless except males; mealybugs [Pseudococ-
cidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
183' Tarsi usually present, and 2-segmented (sometimes 1-segmented, but rarely atrophied);
claws paired; antennae 1-to 6-segmented (commonly 5-to 6-segmented), last segments
bear an elongate processus terminalis; both sexes either winged or apterous; when
winged, hindwing with 2 oblique veins; abdomen without or with a pair of siphunculi;
foliar and root aphids [Aphididae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
184(183) Antennae 6- or 7-segmented; quinquelocular pores numerous and evenly distributed
dorsally except at middle with almost a single row of triocular pores; 4 pairs of cerarii
confined in the 6th and posterior segments of abdomen; anal lobe cerarii each with 2 or
3 lanceolate setae, triocular pores absent (Fig. 372); anal ring not joined at anterior end
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brevennia rehi (Lindinger)
184' Antennae 8-segmented; quinquelocular pores absent; 16 pairs of cerarii present but
without ocular and preocular pores; anal lobe cerarii with a pair of conical setae
surrounded by a dense cluster of triocular pores and 6 auxiliary setae; anal ring with a
double row of pores (Fig. 373) . . . . . . . . . . . . . . Pseudococcus saccharicola Takahashi
185(183') Apterae with or without siphunculi, or with first two tarsal segments fused (showing
single segment appearance); antennae shorter than 1/4 of body length and 5-segmented.
Alate either without siphunculi, or with annular rhinaria borne on 6-segmented anten-
nae; median veins of wing unbranched (Fig. 374) . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
185' Apterae always with siphunculi; first 2 tarsal segments distinctly separated. Alate forms
always with siphunculi but without annular rhinaria; median veins of fore wings 1 to 2
branches. Antennae of both forms 6-segmented, processus terminalis of ultimate anten-
nal segment longer than base of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
186(285) Siphunculi distinct, conical in shape and with a rim; anal plate not displaced dorsally
(normal); abdominal segment VIII bears 2 dorsal hairs. First and second tarsal seg-
ments in apterae appear 1-segmented; antennae 5-segmented (Fig. 375); prominent
dorsal groups of facetted wax glands present on abdomen. Alatae with many annular
rhinaria on antennal segments III-V (Fig. 376); segment III and V of antennae each
prominently longer than segment IV . . . . . . . . . . Tetraneura nigriabdominalis (Sasaki)
186' Siphunculi absent; anal plate of both apterae and alatae displaced dorsally; abdominal
segment VIII with 6-8 dorsal hairs; second segment of hind tarsus shorter than the
ultimate rostral segment. Apterae hairy along antennae and dorsal body, Alatae with a
few rounded or irregularly-shaped rhinaria on antennal segments III-V (Fig. 377); and
such characters absent in apterae (Fig. 378); venter of abdomen without multifaceted
wax plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica lucifuga (Zehntner)
187(185') Siphunculi very short, conical, and lightly colored compared to the cuticle; segments
I-VI of abdomen each with 1-2 spinal hairs, 2-3 marginal pairs and a lateral pair; body
dorsum including head (Fig. 379) with long, stout, and pointed hairs in apterae (alatae
with abdominal hairs borne on brown sclerites); antennae 5-segmented with processus
terminalis about 1.75-2.25 times base of segment V and segment III with 2-4 spinal hairs;
cauda rounded with long hairs and a neck-like median (Fig. 380) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sipha flava (Forbes)
187' Siphunculi longer than wide at base; antennae usually 6-segmented . . . . . . . . . . . 188
188(187') Lateral abdominal tubercles usually absent from abdominal segments I-VII and variably
occurring on segments II-VI; spiracles of abdominal segments I and II near each other
with their pigmented areas nearly contiguous; frontal tubercles well developed; anten-
nal hairs capitate; ultimate rostral segment (Fig. 381) constricted near base and 0.7-0.8
times larger than the 2nd segment of the posterior tarsus; genital plate of females
subspherical with a pair of moderately long setae (Fig. 382); cauda with 8-9 hairs (Fig.
383) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Sitobion) avenae (F.)
188' Lateral abdominal tubercles usually present on abdominal segments I and VII, and
larger than those on II to VI; spiracles of abdominal segments I and II far apart; frontal
tubercles never well developed; alatae usually without a dorsal patch; antennal hairs
never capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
189(188') Cauda pale and not pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
189' Cauda pigmented brownish; concolorous at least to apices of siphunculi . . . . . . . 192
190(189) Cauda with 9-17 hairs (Fig. 384); antennae 2/3-3/4 times body; brownish black siphun-
culi shorter than greyish-yellow cauda (Fig. 385); alatae with dark crossbars on ab-
domen (Fig. 385) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis sacchari (Zehntner)
190' Cauda with fewer hairs about 4-6 only (Fig. 386) . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
191(190') Cauda pale with 4 hairs only; siphunculi uniformly dark, rough-imbricate, and much
longer than whitish cauda (Fig. 387); femora and first 2 antennal segments darkly
pigmented; segment IV of antennae pale but with a small brown apical ring; alatae with
median vein of forewing twice-branched; reddish brown aphids . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hysteroneura setariae (Thomas)
191' Cauda with 4-6 hairs; siphunculi more slender, smooth, and dark apically (Fig. 388);
femora and first 2 antennal segments pale; antennal segment IV uniformly dusky brown;
alatae with median vein of forewing single-branched; green aphids . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizaphis graminum (Rondani)
192(189') Aphids colonizing roots and stem bases; cauda with 4 hairs (Fig. 389); siphunculus
sculptured, cylindrical with apices colored brown similar to cauda (Fig. 390); segment
VIII of abdomen with 4-8 hairs dorsally; hairs on body and legs long and fine similar to
antennal hairs; antennae 5-segmented (rarely with 6 segments), antennal hairs much
longer than maximum diameters of segments (Fig. 391); greyish black aphids with
reddish tinge in the abdomen . . . . . . . . . . . . Rhopalosiphum rufiabdominalis (Sasaki)
192' Aphids common on the aerial parts; segment VIII of abdomen with only 2 dorsal hairs;
antennae of alatae and apterae 6-segmented with antennal hairs shorter than maximum
diameters of segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
193(192') Processus terminalis prominently long and usually 3-4 times base of antennal segment
VI; dorsum of abdomen with circular bead-like cuticular arrangement (Fig. 392);
siphunculi over twice length of cauda, very smooth, and distinctly clavate in the dusky
apical 1/2 and pale in the basal half (Fig. 393); brown aphids . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum nymphaeae (L.)
193' Processus terminalis short, only 1.7-2.5 times base of antennal segment VI; antennae
short, less than half body length, median of vertex elevated (Fig. 394); apterae with
antennal hairs longer than maximum diameters of segments; marginal tubercles located
above midlength of spiracles 1 and 2 and after 7th spiracle; siphunculi uniformly dark
and spinulose, up to 1.6 times longer than black cauda (Fig. 395); green to dark olive
green aphids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum maidis (Fitch)
194(182') Tegula lacking; midcoxa short and not widely separated; antennal pedicel rarely thicker
than scape, without wart-like sensillae; hind tibia long with rows of articulated small
spines, without tibial spur (Fig. 396); leafhoppers [Cicadellidae] . . . . . . . . . . . . . . 195
194' Tegula present on mesothorax; midcoxa elongate and widely separated; pedicel en-
larged, often bulbous with numerous wart-like sensillae; hind tibia with a large movable
spur (Fig. 397); planthoppers [Delphacidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
195(194) Adultleafhoppers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
195' Fourth and fifth instar nymphs of leafhoppers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
196(195) Opaque green leafhoppers with black markings on head, face, pronotum, tegmen, and
other body parts; vertex sharply ridged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
196' Color highly variable but never green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
197(196) Vertex without black submarginal band or with band partially developed or with traces
only behind ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
197' Vertex with a black submarginal band fully developed (entirely contiguous) . . . . . 199
198(197) Vertex slightly longer medially than next to eye (11:10), its cephalic margin rather
rounded or slightly bulging; comma-shaped transverse black band behind ocelli in males
(Fig. 398); absent in females (Fig. 399) . . . Nephotettix malayanus Ishihara & Kawase
198' Vertex much longer medially than next to eye (11:8), its anterior margin markedly
pointed; marginal and submarginal bands absent in both sexes (Fig. 400) . . . . . . . . . . . .
................................................. Nephotettix virescens (Distant)
199(197') Anterior margin of pronotum without black markings ........................ 200
199' Anterior margin of pronotum with black markings .......................... 201

200(199) Anterior margin of vertex rounded at center (Fig. 401) . . . . . . . . . . . . . . . . . . . . . . . . . . .
................................................. Nephotettix cincticeps (Uhler)
200' Anterior margin of vertex angularly projecting or distinctly pointed in center (Fig. 402)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephotettix parvus Ishihara & Kawase
201(199') Marginal black band of vertex rather broad and contiguously developed between ocelli
(Fig. 403); inner margin of clavus with a black band . . . . . Nephotettix nigropictus (Stål)
201' Marginal black band of vertex much reduced and present only in midcephalic margin
(Fig. 404); African species . . . . . . . . . . . . . . . . . . . . . . Nephotettix modulatus Melichar
202(196') Forewing maculate (spotted) or with zig-zag band ........................... 203
202' Forewing not as above .................................................. 204
203(202) Forewing with a black to dark brown longitudinal zig-zag band (Fig. 405); vertex and
pronotum whitish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Recilia dorsalis (Motschulsky)
203' Forewing with dusky markings (Fig. 406); vertex with a broad transverse band between
the compound eyes and a thin band running along the margin below the ocelli (Fig. 407);
pronotum and mesonotum with grey and white patterning . . . . . . . . . . . . . . . . . . . . . . . .
............................................... Scapitoideus morosus Melichar
204(202') Ocelli on disc of vertex closer to hind margin than to anterior margin; frontoclypeus
swollen; head with a median apical black spot (Fig. 408); large pale whitish leafhopper
...................................................... Cofana spectra (Distant)
204' Ocelli either on or closer to anterior margin of vertex or on face; frontoclypeus not
swollen ................................................................ 205
205(204') Vertex without spots; pronotum with paired light triangular markings (Fig. 409) . . . . . .
........................................................................ 206
205' Vertex with spots; Fronotum without such markings ......................... 207
206(205) Processes of aedeagus broader; median ridge of 7th sternite exserted in females
(Fig. 410) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thaia oryzivora Ghauri
206' Processes of aedeagus narrow; median ridge of 7th sternite not exserted in females
(Fig. 411) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thaia subrufa (Motschulsky)
207(205') Vertex with a black median spot touching the anterior margin (Fig. 412) . . . . . . . . . . . .
...................................... Empoascanara maculifrons (Motschulsky)
207' Vertex with 2 black spots of the anterior margin (Fig. 413); head and thorax golden
yellow; forewing unspotted or without a dark fuscous or black longitudinal stripe along
discal cell (Fig. 414) . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicadulina bipunctata (Melichar)
208(195') Ocelli prominent on crown of vertex between eyes; anterior margin of vertex bulging and
rounded; body with bluish tinge (Fig. 415) . . . . . . . . . . . . . . . .Cofana spectra (Distant)
208' Ocelli along the anterior margin of pointed or rounded vertex (Fig. 416); dorsal body
surface with spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
209(208') Spines present only on the dorsal posterior margin of the last 2-3 abdominal segments;
entire dorsum of abdomen may be black or orange except penultimate segment with 2
black spots; pronotum with a pair of L-shaped bands (Fig. 417)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicadulina bipunctata (Melichar)
209' Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
210(209') Body with spines including head, thorax, abdomen and wing pads . . . . . . . . . . . . . 211
210' Body with abdominal spines only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
211(210) Wing pads without spines; prominent black spot present in mid-anterior portion of
vertex (Fig. 418) . . . . . . . . . . . . . . . . . . . . . . Empoascanara maculifrons (Motschulsky)
211' Body clothed with spines including wing pads (Fig. 419) . . . . . . . . . . . . . . . . Thaia spp.
212(210') Head an thoracic markings more suffused and less prominent . . . . . . . . . . . . . . . . 213
212' Black or dark markings present on the body dorsum . . . . . . . . . . . . . . . . . . . . . . . 214
213(212) Frontal view of head with 2 curved but parallel orange bands running between 2
compound eyes (Fig. 420) . . . . . . . . . . . . . . . . . . . . . . Scaphoideus morosus (Melichar)
213' Frontal view of head without such bands; subdorsal band prominent in the abdomen
(Fig. 421) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Recilia dorsalis (Motschulsky)
214(212') Pale green to yellow leafhopper; body with minimal markings; last abdominal segment
with a dark spot on apical 2/3 (Fig. 422) . . . . . . . . . . . . Nephotettix virescens (Distant)
214' Greenish leafhoppers with dark body markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
215(214') Markings on last abdominal segment absent; anterior margins of vertex with 2 black
spots (Fig. 423) . . . . . . . . . . . . . . . . . . . . . . . Nephotettix malayanus Ishihara & Kawase
215' Markings on last abdominal segment distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
216(215) Bead-like band present on the last abdominal segment (Fig. 424); body markings
variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephotettix cincticeps (Uhler)
216' Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
217(216') Abdominal dorsum with light markings; median stripe on the last abdominal segment
poorly developed; sub-marginal band below ocelli pale; cheek unshaded, clypeus with
black median band (Fig. 425) . . . . . . . . . . . . . . . Nephotettix parvus Ishihara & Kawase
217' Abdominal dorsum with dark markings; median stripe on the last abdominal segment
distinct; clypeus with partial or entire black band . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
218(217') Vertex with 3 pairs of bands (Fig. 426); face entirely black . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephotettix nigropictus (Stål)
218' Vertex with fewer bands; upper median of clypeus and upper portion of cheek (below
compound eyes) with black spots (Fig. 427) . . . . . . . Nephotettix modulutus Melichar
219(194') Adult planthoppers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
219' Fourth and 5th instar planthopper nymphs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
220(219) Prominent fixed spines on basal segment of post-tarsi; frons not excavated; median
carina not intercepted (Fig. 428); mesonotum with 4 pale brown longitudinal markings
(Fig. 429); posterior margin of male pygofer with no central process ventrally;
parameters not bifurcated apically (Fig. 430) . . . . . . . . . . . . . Nilaparvata lugens (Stål)
220' No prominent spines on basal segment of post-tarsi . . . . . . . . . . . . . . . . . . . . . . . . . 221
221(220') Vertex quadrate, as long as broad, slightly narrower than eye; lateral pronotal carinae
concave and incomplete (Fig. 431); genae black, with yellowish brown lateral carinae;
mesonotum black with white-tipped scutellum; parameres simple and club-shaped
(Fig. 432) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laodelphax striatellus (Fallen)
221' Vertex longer than broad at base with prominent whitish yellow areas on vertex,
pronotum and mesonotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
222(221') Pronotal carinae rarely straight and almost reaching posterior margin; mesonotal carina
more or less strongly divergent; mesonotum whitish medially with outer sides of lateral
carinae dark brown (Fig. 433); forewings with black pterostigma; parameres broad
basally and bifurcated apically with no large spine; ovipositor moderately long and
never deeply curved dorsally in its basal half (Fig. 434) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sogatella furcifera (Horvath)
222' Pronotal carinae usually straight, never reach posterior margin; mesonotal carinae
parallel in the anterior half and weakly divergent; ovipositor long and slender, deeply
curved dorsad in its basal 1/2 and straight or weakly recurved distally; Latin American
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
223(222') Clavus without spot (Fig. 435); apex of style broad with the inner margins pointed and
with a marked carina; infuscation at apex of tegmen confined to the apical cells
posterior to M3 + 4; ovipositor large and more prominently serrated (Fig. 436); body 3-4
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tagosodes orizicolus (Muir)
223' Clavus with a spot (stigmata) between the common claval vein and the commissural
margin, in folded wing spots form a saddle-shaped stigmata (Fig. 437); apex of style
small, slender and curved inward; ovipositor (Fig. 438) relatively narrower and not
prominently serrated; body 2 mm long . . . . . . . . . . . . . Tagosodes cubanus (Crawford)
224(219') Vertex plate about as long as wide; head narrow; dorsal portion of body (thorax and
abdomen) with tinge of grey and white markings . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
224' Vertex plate wider than long, head broad; dorsal portion of body with brown pigmenta-
tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
225(224) Thoracic and abdominal dorsum uniformly marked with various amounts of grey and
white markings on a creamy white background; dorsal white band indistinct (Fig. 439)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sogatella furcifera (Horvath)
225' Thorax and abdomen not as above; dorsal white band distinct . . . . . . . . . . . . . . . . 226
226(225') Tip of vertex truncated; lateral margins of pronotum broad; subdorsal portion of
abdomen with 6 pairs of bands (Fig. 440) . . . . . . . . . . . Tagosodes cubanus (Crawford)
226' Tip of vertex partly emarginate; lateral margins of pronotum acute and pointed
downward (Fig. 441); sub-dorsal portion of abdomen with 7 pairs of bands . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tagosodes orizicolus (Muir)
227(224') Post-clypeus pale yellow; head width large (0.56-0.70 mm) (Fig. 442); apices of ab-
dominal segments with whitish dome-shaped indentations (Fig. 443) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nilaparvata lugens (Stål)
227' Post-clypeus dark brown; head width small (0.52 mm) (Fig. 444); median area of
abdomen whitish with darker margins (Fig. 445) . . . . Laodelphax striatellus (Fallen)
228(181') Antenna entirely concealed beneath (except in Ochterus), and shorter than head;
arolium absent . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
228' Antennae exposed or visible above, as long as or longer than head; arolium present or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
229(228) Ocelli present; flat and almost entirely black; all legs slender, not raptorial and legs II
and III without fringes of swimming hairs; beak slender reaching hind-coxae, enlarged
basal segment black and thin, slender apical segments brownish red; depressed antero-
lateral margin of pronotum brownish yellow along apical margins and yellowish basally;
posterior margin bordered yellow-orange, median area convex (recurved) (Fig. 446);
hemelytra with 2 yellow spots (base of membrane and at mind-length of embolium-ex-
ocorium) and a moderately long yellow band at base of embolium-exocorium (Fig. 447)
[Ochteridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ochterus marginatus (Latreille)
229' Ocelli absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
230(229') Fore-tarsi 1-segmented, scoop-like (palae) with a row of hairs; beak short, hidden and
unsegmented; head slightly overlaps with pronotum, yellow and moderately rounded at
tip (Fig. 448); pronotal disk brownish grey with procurved base; scutellum with a yellow
transverse band at mid-half; forewing brownish grey except transparent membrane and
base of clavus; membrane inwardly curved, tip truncated and apical margin brown; base
of clavus lightly punctated (Fig. 449); nymphs and adults root feeders; aquatic
[Corkidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micronecta quadristrigata Breddin
230' Fore-tarsi not scoop-like; beak segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
231(230') Body very strongly convex above; hind tarsi without claws; forelegs not raptorial . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
231' Body flattened or slightly convex; hind tarsus with claws; front leg raptorial, femur
usually thickened; membrane of elytra with veins . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
232(231) Hind leg short, with 2 claws; hind tibia cylindrical, without long hairs; abdomen without
fine laminate ventral keel on segments 2-6; small insects usually less than 4 mm long
[Pleidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
232 Hind leg long, oar-shaped, with 1 claw; flattened hind tibiae with fringe of long hairs;
abdomen with a broad median ventral keel; hemelytral commissure with an elliptical pit
at anterior end; usually more than 4 mm long [Notonectidae] . . . . . . . . . . . . . . . . 234
233(232) Body 2.2 mm long and 1.4 mm wide; head rugose-punctate with a brown longitudinal
median ridge between eyes, not reaching apex of pronotum; pronotum reticulate
punctate except smooth postero-median area (Fig. 450); scutellum with traces of reticu-
late punctations; hemelytra reticulate punctate with brown pits at each punctate;
propleuron carinate yellow at base of coxa I and with a small protrusion at its hind part;
mesopleural lobe strongly rounded cephalad and slightly indented posterad below
mid-half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraplea sobrina Stål
233' Body 1.6 mm long and 0.8 mm wide; head nearly smooth with a moderately broad
median longitudinal brown band between eyes almost reaching apex of pronotum,
center area with a yellow spot; pronotum smooth to shallowly punctate (Fig. 451);
scutellum smooth; hemelytra reticulate punctate with whitish pits; propleuron straight
without tubercle at hind of coxa I; mesopleural lobe weakly rounded cephalad and
widely procurved at mid-half . . . . . . . . . . . . . . . . . . . . . . . . . . Paraplea liturata (Fieber)
234(232') Eyes strongly convergent, touching each other postero-dorsally or with a very narrow
gap (0.03 mm) in between (Fig. 452); head rounded in lateral view; terminal antennal
segment pale yellow (Fig. 453); tibia I slender, thin and strongly flattened, without
spines; apical 1/2 of tibiae with a brown median band; tarsus I 0.6 times length of tibiae
(Fig. 454) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anisops sp.
234' Eyes moderately convergent with a clear separation (0.07 mm) postero-dorsally (Fig.
455); head pointed in lateral view; terminal segment of antenna brown (Fig. 456); tibia I
slightly concave and cylindrical with spines; apical half of tibia without band; tarsus I 0.7
times length of tibia (Fig. 457) . . . . . . . . . . . . . . . . . . . . . Anisops kuroiwae Matsumura
235(231') Tarsi 2-segmented; apex of abdomen with a pair of flat, retractile airstraps; margins of
pronotum and wings yellowish, thin, and rather flap-like; head snake-like in front and
triangular dorsally (Fig. 458); eyes small, slightly convex in the inner posterodorsal side,
elongate and 3 times longer than height seen laterally; proboscis hidden beneath; venter
of abdomen elevated medially, basal 2 segments narrow and slightly converging mid-
ventrally; clavus and corium brownish and finely punctated; membrane hyaline with 6
rectangularly closed cells (Fig. 459); apical lower end of corium with globular dense
mass of short brown setae [Belostomatidae] . . . . . . . Diplonychus rusticus (Fabricius)
235' Tarsi l-segmented; eyes prominent and transverse; proboscis visible in front (Fig. 460);
apex of abdomen with a non-retractile, cylindrical siphon; venter with static sense
organs; body long, narrow and cylindrical with nearly contiguous coxae II and III; front
leg sickle-shaped with a strong midfemoral tooth (Fig. 461) [Nepidae] . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ranatra diminuta Montadon
236(228') Tarsal claws anteapical (especially on forelegs); tip of last tarsal segment more or less
cleft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
236' Tarsal claws apical; tip of last tarsal segment entire . . . . . . . . . . . . . . . . . . . . . . . . . 243
237(236) Ocelli present; long-legged; hind femora extending beyond apex of abdomen; mid-
coxae closer to hind- than forecoxae; medial margins sinuate [Gerridae] . . . . . . . 238
237' Ocelli absent; short-legged; hind femora not reaching apex of abdomen; mid-coxae
equidistant between hind- and forecoxae; medial margins straight [Veliidae] . . . 242
238(237) Body comparatively short and oval; inner margin of eyes convex; posterior end of
abdomen produced to a spine-like point; body entirely black dorsally and ventrally
except orange-brown band at midapex of pronotum, eye margins, and hind vertex; basal
half of femur I yellow (Fig. 462) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagadotarsus sp.
238' Body long and thin, inner margins of eyes concave; pronotum with a yellow median
longitudinal stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
239(238') Hindtibia at least 4 times as long as the first tarsal segment; 1st antennal segment longest
but shorter than segments 2 + 3 (Fig. 463); head and pronotum with whitish markings,
venter light brown with 2 dark brown patches at anterior 1/2 of mesosternum; pronotum
with a mid-longitudinal band (Fig. 464); hemelytra brown with dark brown embolial
region; mid-leg with brown femora at anterior l/2 and dark brown at posterior 1/2 and
with a spine at distal end; 1st tarsal segment of leg II 4 times as long as the 2nd . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerris adelaides Dohrn
239' Hidtibia not as above; 1st antennal segment equal or longer than segments 2 + 3 . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
240(239') Anterior lobe of pronotum with a large subglobular to quadrangular yellow spot
(Fig. 465) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnogonus parvulus (Stål)
240' Anterior lobe of pronotum with a pair of longitudinal yellow-brown spots . . . . . . 241
241(240') Posterior lobe of pronotum with a longitudinal yellow line (Fig. 466). . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnogonus fossarum (Fabricius)
241' Posterior lobe of pronotum without a longitudinal yellow line (Fig. 467). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnogunus nitidus (Mayr)
242(237') Apical ventral sternite of male abdomen angled at corner, base not evenly rounded;
paramere thin immediately above curved point and broadens just below it (Fig. 468);
general shape short but broad, usually black with small grey areas (Fig. 469); abdomen
of females laterally compressed at base . . . . Microvelia douglasi atrolineata Bergroth
242' Apical ventral sternite of male abdomen rounded at corner, base evenly rounded,
paramere nicely sickle-shaped, still wide at curvature point and tapers apically, uniform-
ly broad towards base (Fig. 470); greyish to brown (Fig. 471); abdomen of females not
markedly compressed laterally at base . . . . . . . . . . .Microvelia douglasi douglasi Scott
243(236') Head long and slender, as long as thorax and dilated distally; elongate thorax with
nonsulcate mesosternum and mid-coxa closer to fore-coxa than to the hind-coxa; eyes
situated a little below mid-length of head; body long, slender and cylindrical (Fig. 472)
Fig. 472 [Hydrometridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
243' Head not more than twice as long as wide, not dilated distally; eyes near base of head;
bodyshortandstout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
244(243) Tergite X cylindrical and longer than IX, with a longer apicomedian spine; wing tips
extended to 2/3 of tergite VII; wings with the white band below 2nd longitudinal vein
with few patches inside 2 closed rectangular cells, outer closed cell with an oblong white
spot at midbase; vein touching costal area short, 0.54 times basal length of outer cell
(Fig. 473) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrometra orientalis Lundbland
244' Tergite X nearly as long as IX, with apex widened laterally and relatively short
apicomedian spine; wing tips a little beyond segment VIII; white band of wing above the
2nd longitudinal vein; outer cell with a white band along entire basal half, vein touching
costal area long, as long as bottom length of outer cell (Fig. 474). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrometra lineata Eschscholtz
245(243') Antennae 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
245' Antennae 4-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
246(245) Tarsus 2-segmented; hemelytra with clavus and membrane similar in texture and
without veins; venter of head with a longitudinal groove for beak reception; pronotal pits
in a transverse row; sides of pronotum constricted at mid-half producing an elongated
lobe; body and wings with white hairs; scutellum with a median longitudinal carina
forming 2 lobes; legs without ventral spines, corium veins hardly visible, membrane light
brownish black with 3 white spots (Fig. 475) . . . . . . . . . . . . Hebrus bergrothi Horvath
246' Tarsus 3-segmented; hemelytra distinct from clavus and usually with veins . . . . . 247
247(246') Tibiae with numerous stout spines, usually in 2 rows (Fig. 476); apices of mid and hind
coxae fringed with nearly contiguous rigid setae; forelegs fossorial; tarsi 3-segmented;
usually small, slightly depressed, black oval to elliptical; burrower bugs [Cydnidae]. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
247 ' Tibiae witbout numerous strong spines (Fig. 477); apices of mid and hind coxae not
fringed with closely set rigid setae; abdominal venter sometimes moderately elevated
but not prominently keeled or tectiform to flattened; frenum, if present hardly extended
beyond 3/4 marginal length of scutellum [Pentatomidae] . . . . . . . . . . . . . . . . . . . . . 249
248(247) Small sized bugs, 5 mm long; vertex rounded and hairy, peg-like spines along apical
margins of vertex absent (Fig. 478) . . . . . . . . . . . . . . . . . Geotomus pygmaeus (Dallas)
248' Medium sized bugs, 6-7 mm long, vertex notched mid-anteriorly and with 10 peg-like
spines along margins (Fig. 479) . . . . . . . . . . . . . . . . . . . . . . Aethus indicus (Westwood)
249(247') Phytophagous bugs; all femora with a dorsal spine apically (Fig. 480); tip of abdomen
with 4 spines (Fig. 481); lateral angle of pronotum with laterally projected spines (Fig.
482); tylus longer than juga; head with 6 rows of longitudinal punctations; 2nd antennal
segment as long as 3rd; African species . . . . . . . . . . . . . . . . . . . . . . . Diploxys fallax Stål
249' Phytophagous bugs without dorsal spine at apex of femora . . . . . . . . . . . . . . . . . . . 250
249'' Predatory bugs without dorsal spine at apex of femora . . . . . . . . . . . . . . . . . . . . . . 274
250(249') Pronotal spine more or less prominent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
250’ Pronotal spine absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268
251(250') Tylus slightly beyond juga; head long; body dorsally flat and convex underneath; 3rd
antennal segment longer than the 2nd; 2nd abdominal segment impressed medially not
obtusely convex, devoid of a tubercle; mesosternum not carinated; lateral margins of
head prominently trenchant; frenum not passing midscutellum; tibiae feebly planosul-
cate; fulvous species (Eysarcoris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
251' Tylus and juga in the same line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
252(251) Lateral angle of pronotum prominent or partially developed; head blackish; tylus and
juga with nearly contiguous punctuations; body length 5.8-6.7 mm . . . . . . . . . . . . 253
252 ' Lateral angle of pronotum not prominent; head brownish with punctations rather sparse
and separated; body length 4-5.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
253(252) Lateral angle blackish, robust, and with a blunt end, lateral to slightly upward; tylus with
a mid-longitudinal yellow dagger-like band; base of scutellum with large ovoid and
obliquely set spots; midscutellum with a transverse groove; a pair of black spots above
calli present (Fig. 483) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eysarcoris montivagus Distant
253' Lateral angle less developed to sharply pointed but never projected upward; tylus
without a yellow dagger-like band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
254(253') Lateral angle of pronotum not sharply pointed; tylus almost entirely black, calli slightly
concave and bear no black spots behind; midscutellum not deep; second antennal
segment shorter than the 3rd segment; yellow spots on the basal outer edge of scutellum
not as oblique as in E. montivagus; body length 5.82 mm (Fig. 484). . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eysarcoris guttiger (Thunberg)
254' Not as above; pronotum’s lateral angle blackish and acute or sharply pointed, directed
laterally; tylus and juga in same line producing a broad tip; body length 6-6.5 mm
(Fig. 485) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eysarcoris panvus Uhler
255(252') Body 5-5.8 mm long and 3 mm wide; head darker than E. capitatus with blackish
punctations along jugal margins, area between ocelli and compound eyes, and edge of
tylus; tylus with a thin yellow longitudinal stripe; calli triangular-like with black outer
base; second antennal segment shorter than 3rd; base of scutellum with 3 yellow spots,
median slightly concave and tip relatively narrow but rounded (Fig. 486). . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eysarcoris ventralis Distant
255' Body barely 4.3 mm long; head yellowish brown with light blackish brown well separated
punctations; margins of juga black; calli not well defined; tip of scutellum broadly
rounded with nearly parallel sides (Fig. 487) . . . . . . . . . . . Eysarcoris capitatus Distant
256(251') Scutellum short, not as long as abdomen; body length 9-12 mm dorsally flat and ventrally
convex; American species (Oebalus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257
256' Scutellum very long, reaching tip of abdomen, almost rectangular but not covering
lateral portion of forewing; anterolateral process of pronotum toothed or elongately
spinelike with tip projected in different directions; body length 6.5-11 mm; Asian and
African species (Scotinophara) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
257(256) Tylus exerted slightly beyond juga (Fig. 488); margin of pronotum not clearly separated
from disc; each pronotal corner with small teeth-like tubercles; second antennal seg-
ment shorter than 3rd; scutellum dark yellow along margins forming a Y-shaped ap-
pearance (Fig. 489); body elongated, body length 9 mm . . . Oebalus poecila (Dallas)
257' Tylus and juga in the same line; head long; pronotal shoulders with long latero-forward
directed spines; scutellum much longer than wide at base with knob-like yellow bands in
the outer basal half margin, apex yellow; body length 12 mm (Fig. 490). . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oebalus pugnax (Fabr.)
258(256') Anterolateral margins of pronotum between anterior and humeral processes serrate or
dentate; anterior portion of pronotum irregularly rounded but with a distinct transverse
depression behind . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259
258' Anterolateral margins of pronotum between anterior and humeral processes not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
259(258) Midanterior part of pronotum not swollen, anterolateral margins serrate or dentate with
a small anterior process shorter than the humeral processes; head and anterior part of
pronotum same color as the rest of the body (Fig. 491); femora without a pale annulation
apically; clasper with a slight hook at apex and straight margin below it (Fig. 492). ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scotinophara serrata (Vollenhoven)
259' Pronotum tumescent at the midanterior part; anterolateral margins minutely serrate;
anterior processes distinct and usually longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260
\
260(259') Transverse depression on pronotum obscure; anterior part less swollen; brownish
ochraceous, anterior pronotum, head, and base of posterior tibiae black; rest of
posterior tibiae dark brown; pronotal anterior processes straight and spinous (Fig. 493)
..................................................... Scotinophara parva Yang
260' Transverse depression between humeral processes distinct; anterior part of pronotum
swollen; humeral process triangular; anterior processes of pronotum distinctly curved
forward (Fig. 494); tips of clasper inwardly curved, base with acute tubercle (Fig. 495);
black or rarely reddish brown, only posterior tibiae black. . . . . . . . . . . . . . . . . . . . . . . . . .
.................................................. Scotinophara scotti Horvath
261(258') Postocular anterior margins of pronotum broad and oblique; anterolateral margins
insinuate .............................................................. 262
261' Postocular anterior margins of pronotum straight to slightly sinuate; anterolateral

margins of pronotum anteriorly excavate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263
262(261) Pronotal disk not excavated anteriorly and a transverse depression behind excavation
weak; scutellum without a pair of distinct oblique depressions; tip of anterolateral spine
of pronotum projected posterad but not extended beyond anterior angle of pronotum
(Fig. 496); clasper with a large tubercle medially (Fig. 497) . . . . . . . . . . . . . . . . . . . . . . . .
................................................... Scotinophara coarctata (F.)
262' Disk of pronotum excavated anteriorly and a transverse depression behind it prominent;
scutellum with a pair of oblique depressions before midlength; tylus as long as lateral
lobes (Fig. 498); 3rd antennal segment slightly less than twice as long as the 2nd (Fig.
499); apical segment as long as segments II & III combined. . . . . . . . . . . . . . . . . . . . . . . .
............................................ Scotinophara inermiceps (Breddin)
263(261') Anterolateral margins of pronotum straight and laminate; anterior processes of
pronotum long and directed forwards; dorsum longly pilose; anterior pronotum rather
gibbous (Fig. 500) ............................. Scotinophara horvathi (Distant)
263' Anterolateral margins of pronotum totally or partially carinate, excavate on anterior
half and insinuate below; anterior pronotum not swollen; dorsum shortly pilose . . 264
264(263') Anterior processes of pronotum projected horizontally ...................... 265
264' Anterior processes of pronotum projected anteriorly ........................ 267
265(264) Black species; humeral processes of pronotum distinct, as long as or slightly shorter than
the apical ones, the latter spinous and extended beyond outer margin of eye; rostrum
reaching midpoint of hind coxae; scutellum twice as long as width at narrowest point
(7:6:12); apex of antenniferous tubercle slightly bifid (Fig. 501); clasper slightly emar-
ginate apically (Fig. 502) . . . . . . . . . . . . . . . . . . . . . . . Scotinophara lurida (Burmeister)
265' Brownish or ochraceous species; other characters not as above . . . . . . . . . . . . . . . . 266
266(265') Tylus shorter than juga (Fig. 503); femora black with a pale ring near tip; rostrum
extended beyond hind coxae to abdomen; clasper acute-triangular in the inner apex
(Fig. 504); scutellum nearly 1/3 times longer than width at narrowest point; apex of
antenniferous tubercle blunt . . . . . . . . . . . . . . . . . . . . . Scotinophara obscura (Dallas)
266' Tylus as long as juga (Fig. 505); femora without subapical ring; scutellum not reaching
abdominal tip with apex subtruncate; hemelytra without distinct pale line (Fig. 506);
antennae piceous, body beneath widely black . . . . . . Scotinophara latiuscula Breddin
267(264') Anterior processes of pronotum long, nearly as long as 1st antennal segment; humeral
processes as long as anterior processes of pronotum; pronotum gibbous anteriorly with
a pale central carinate line and a transverse impression (Fig. 507); clasper slightly broad
distally, base with a small tooth-like structure (Fig. 508) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scotinophara bispinosa (F.)
267' Anterior processes of pronotum short; less than the length of the 1st antennal segment;
humeral process shorter than anterior (Fig. 509); ochraceous with black head;
transverse impression on pronotum wanting; clasper narrow medially, wide on ends
(Fig. 510) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scotinophara ochracea (Distant)
268 (250') Abdominal sternite III with an anteriorly projected tubercle reaching midcoxae (Fig.
511) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
268' Venter of abdominal segment III without a tubercle . . . . . . . . . . . . . . . . . . . . . . . . 272
269(268) Body yellow, surface semi-translucent and smooth, with fine to minute black punctures;
2 nd antennal segment as long as 3rd; scent gland spout long and thin (Fig. 512); spiracles
black; pronotum with a transverse white to reddish band (Fig. 513); body length 8-
11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Piezodorus hybneri (Gmelin)
269' Body more or less oval, distinctly colored yellow to orange-yellow in the pronotum and
scutellum; corium with yellow spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270
270(269') Tylus and juga in same line; head with 3 pale ochraceous longitudinal stripes and a small
spot of same color before the compound eye; scutellum extended to 5th abdominal
segment, apex narrow, frena reaching near apex; callus near each other and with a black
longitudinal band in between; pronotum with a wider yellow patch behind calli (Fig.
514); last ventral segment moderate in males and abdominal segments not contracted
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Menida formosa (Westwood)
270' Tylus slightly exserted beyond juga; head with 5 pale ochraceous longitudinal stripes;
6th abdominal segment contracted medially; last ventral segment very large in male and
bell-shaped (Pygomenida) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
271(270') Top of scent gland with a broad black band; posterior margin of metapleuron black
between yellow metacoxae and lateral margin of abdomen (prominent in males); ventral
yellow bands on abdomen relatively thin and narrow; black bands along margins of
scutellum in the apical 1/3 and midbasal 1/3 less prominent (Fig. 515)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pygomenida varipennis (Westwood)
271' Top of scent gland with a broad yellow band; posterior margin of metapleuron yellow
from metacoxae to lateral margin of abdomen; ventral yellow bands on the abdomen
long and broad; black spot along margins of scutellum in the apical 1/3 and midbasal 1/3
somewhat broad and distinct (Fig. 516) . . . . . . . Pygomenida bengalensis (Westwood)
272(268') Green coloration with or without yellow marks on the head and anterior portion of
pronotum; scutellum devoid of prominent spots; humeral area of pronotum rounded
(Fig. 517); apical 3 segments of antennae with light brown tinge (Fig. 518); scent gland
spout short; rostrum fine reaching hind coxae; moderately large 11-18 mm long . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nezara viridula (Linnaeus)
272' Allied to N. viridula; humeral area (Fig. 519) of pronotum rather pointed and produced
beyond bases of wings; apical 3 segments of antennae (Fig. 520) blackish except distal
and proximal ends of segments V & VI and basal 2/3 of segment IV; East Asian species
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nezara antennata Scott
272" Yellowish orange to brownbodied bugs; scutellum with yellow or black spots . . . . 273
273(272") Pronotum (Fig. 521) with 4 black longitudinal bands, interrupted along apical 1/3;
scutellum with 2 black median longitudinal bands with interruptions along basal 1/3;
corium with 3 elongated black spots; tylus longer than juga; 2nd antennal segment as
long as segment III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antestia anchora (Thunberg)
273' Pronotum and scutellum without black bands; lateral angle of pronotum more produced
than A. anchora; tylus and juga in the same line, mid-juga more heavily punctated than
tylus; head and pronotum punctated blackish brown; antennal segment III twice as long
as segment II (Fig. 522); tip of scutellum ivory yellow (Fig. 523) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Starioides degenerus (Walker)
274(249") Uniformly metallic blue with luster except brownish membrane; lateral angle rounded
without spine; pronotum, scutellum, clavus, corium and embolium with widely
separated punctations; 4th antennal segment shorter or as long as the distance between
eyes, segment II > III (Fig. 524) . . . . . . . . . . . . . . . . . . . . Zincrona caerulea (Linnaeus)
274' Brownish with reddish brown legs, blackish antennae, and white tip of scutellum (Fig.
525); and entire margin of embolium; humeral spine very distinct (Fig. 526); pronotum,
scutellum, corium and clavus with punctations close to each other; 4th antennal segment
noticeably longer than the distance between eyes; segment II nearly as long as III . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Andrallus spinidens (Fabricius)
275(245') Ocelli absent; beak 4- segmented; hemelytra with or without cuneus; membrane with 1
or 2 closed cells or 2 to 3 basal cells where 7 or more longitudinal veins originate . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
275' Ocelli present, if absent prosternum bears a median transverse stridulatory groove; beak
3- or 4-segmented; hemelytra1 membrane different . . . . . . . . . . . . . . . . . . . . . . . . . 282
276(275) Cuneus absent; membrane of hemelytron with 2 or 3 basal cells from which 7 or more
longitudinal anastomosing veins originate; black corial spot touching inner posterior
margin of cornium (Fig. 527); body elongate; antennae inserted at middle of head;
femora slender, anterior femora with 1-2 small spines; collar as thick as anterior lobe of
pronotum; rostrum thin and long; hind margin of pronotum with black fascia, scutellum
reddish; African species [Pyrrhocoridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysdercus superstitious (Fabricius)
276' Cuneus present; membrane of hemelytron with 2 closed cells [Miridae] . . . . . . . . 277
277(276') Rostrum reaching mid-coxae only; 1st antennal segment as long as vertex; vertex without
bristles; eyes slightly far from anterior margin of pronotum . . . . . . . . . . . . . . . . . . 278
277' Rostrum reaching hind-coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
278(277) Second antennal segment about twice as long as pronotal width as base (1.5 : 0.8); apex
of 1st antennal segment pale, rest black; hemelytra and legs brownish yellow; bases of
tibiae black; median line of scutellum black (Fig. 528); aedeagus spiked including
rounded vesical spiculum (Fig. 529); right clasper branched, one lobe with setae, the
other with serrate margin (Fig. 530) . . . . . . . . . . . . . . . . . . . . Cyrtorhinus fulvus Knight
278' Second antennal segment slightly longer than pronotal width at base; III as long as
pronotal width at base; end of 1st antennal segment greenish yellow; hemelytra and legs
greenish; longitudinal median line of scutellum black (Fig. 531); all tibiae yellow;
aedeagus less spiked with vesical spiculum not evenly rounded (Fig. 532); right clasper
with a subapical spine in the under lobe (Fig. 533) . . . Cyrtorhinus lividipennis Reuter
278'' Second antennal segment nearly as long as pronotal width at base; forewings in part
brownish yellow with black clavus, outer wing edges pale, hyaline to sub-hyaline, closed
wings appearing to be blackish down middle with white borders; 1st antennal segment
yellow (Fig. 534); 2nd segment, head, thorax, and abdomen usually entirely black . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyrtorhinus mundulus (Breddin)
279(277') Arolia present, divergent toward their apices; claws toothed at base; head pointed in
front; pronotum punctate; 1st antennal segment cylindrical; body with short pubes-
cence; hemelytra opaque, abdomen and membranous wings not seen from above . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
279' Arolia absent, replaced by pair of straight hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
280(279) Pronotum lustrous black with minute hairs, yellow collar and light brown postero-lateral
margins; head blackish brown with yellow areas along eye margins, areas near eyes and
juga; first antennal segment reddish brown (Fig. 535); proboscis reaching coxa III (Fig.
536); abdomen reddish pink; apex of scent gland plate flat; corium and clavus glossy
blackish brown, corium with whitish yellow bands at base, dorsal margins and before
apex opposite apical end of clavus . . . . . . . . . . . . . . . Proboscidocoris tibialis Poppius
280' Pronotum dull blackish brown with brown shoulder, collar and behind collar; head
entirely yellow-brown; first antennal segment brown (Fig. 537); abdomen dull straw-
colored; apex of scent gland plate strongly bulged; hemelytra dull brown with a broad
yellow band at base of corium, dorsal margins of corium whitish yellow . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proboscidocoris sp.
281(279’) Pseudarolia present arising from base or inner margins of claw; vertex with round green
spots contiguous to eyes; pronotum with greenish tinge apically; collar absent; head and
pronotum black (Fig. 538); 2nd antennal segment shorter than head width across eyes;
aedeagus with a short pointed outer theca with basal half lightly sclerotized (Fig. 539);
left clasper rounded and with long hairs basally, cleft medially and each lobe with acute
structures (Fig. 540); general color pale yellow-brown to stramineous with black head,
pronotum, pleura and genital segment; legs whitish yellow . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tytthus chinensis (Stal)
281’ Pseudarolia absent; claws (Fig. 541) toothed or thickened at base; vertex without spots;
pronotum punctated, collar present; antennal segment blackish brown, segment I1 more
hairy than I and 3.4 times longer than 1st, twice longer than 3rd, and longer than head
width taken across eyes (Fig. 542); head, pronotum, scutellum and thorax yellowish
brown; wings yellowish with corium and clavus finely punctated; sides of abdomen
reddish brown; legs yellow with brownish bands in the dorso-apical areas of femora . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deraecoris vittatus (Reuter)
282(275‘) Beak 3-segmented; hemelytra with cuneus and embolium, cuneal fracture well
developed, membrane without closed cells; adults with well developed metapleural
scent gland openings; male genitalia strongly asymmetrical; small (1.48-3.6 mm), elon-
gate oval species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .283
282‘ Beak 3- or 4-segmented; hemelytra without cuneus; 5 mm or more in length . . . .286
283(282) Metapleural scent gland opening curved forward, often reaching anterior margin of
metapleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .284
283’ Metapleural scent gland opening directed backward, continued as a fine carina which
curves forward to anterior margin of metapleuron . . . . . . . . . . . . . . . . . . . . . . . . . . .285
284(283) Pronotal collar “absent” (hidden), lateral margins of pronotum wide; head and
pronotum dull punctate; pronotal disc with shallow callosities separated by few punc-
tures (Fig. 543); anterior male tibiae with a row of peg-like teeth; 4th antennal segment
hardly longer than 3rd; aedeagus (Fig. 544) with 3 branches of flagellum; brown to dark
brown except corium, tibiae and femora pale white and cuneus and connexivum light
brown; 1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Orius tantillus (Motschulsky)
284’ Pronotal collar prominently lying completely in front of anterior apical angles of
pronotum, lateral margins narrowly carinate and not broadly reflexed; 1st and 2nd
antennal segments very stout, apical 2 segments thinner than scape and pedicel; inner
portion of corium, apical half of clavus and innermost base and dorsal part of membrane
light colored (Fig. 545) . . . . . . . . . . . . . . . . . . . . . . . Montandoniola moraquesi (Puton)
285(283') All tibiae straight, anterior femora with slender hairs; tibia II and III without long hairs,
pubescence not longer than diameter of segment; beak reaching anterior coxae; prono-
tal collar wide, disc without prominent callus; 2 nd antennal segment much longer than
interocular width; apices of corium and embolium oblique and subequal in width (Fig.
546); apex of metasternum with a bifurcate spine concealed by coxa III . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphiareus sp.
285' Tibia I curved inwards (bowed), anterior femora armed with spines; beak short, not
reaching anterior coxae; pronotal disc with a W-shaped callus; apical width of corium 2
times or more wider than apical width of embolium; apical margin of corium and
embolus strongly curved outward-upward (Fig. 547) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physopleurella armata Poppius
286(282') Prosternum with a median transversely striated stridulatory groove extending anterior
of fore-coxae; head usually with a transverse suture near eyes; beak 3-segmented
[Reduviidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
286' Prosternum not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
287(286) Ocelli absent; tibiae I concave almost at mid-ventral half and with rows of inner lateral
spines; femora I with ventral spines; scutellum with a long basal and short apical
upwardly directed spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 288
287 ' Ocelli present, without combination of above characters . . . . . . . . . . . . . . . . . . . . 291
288(287) Lateral spines on the apex of basal one-third of pronotum nearly as long as pronotum;
apical spine of scutellum parallel with the basal spines . . . . . . . . . . . . . . . . . . . . . . 289
288' Lateral spines on the apex of basal one-third of pronotum about half the length of
pronotum; apical spine of scutellum outwardly and upwardly curved . . . . . . . . . . 290
289(288) Legs yellow except apices of femora brownish black with red-orange tinge (Fig. 548);
bases of tibia lightly red-orange, apical end blackish similar to tarsi; pronotum 1.9 times
longer than head, brownish black medially and reddish marginally in dorsal view;
pronotal lateral spines yellow with brownish tips (Fig. 549) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polytoxus fuscovittatus (Stål)
289' Legs blackish with basal half of femur I (Fig. 550) and base of II and III yellowish;
pronotum 2 times longer than head, bluish postero-medially and with a pair of brown
longitudinal bands in the antero-median area, margins reddish; pronotal lateral spines
yellow basally and blackish in the apical half . . . . . . . . . Polytoxus selangorensis Miller
290(288') Hemelytra with vein M at apical margin of discal cell roundly bent near base, (Fig. 551);
proboscis not reaching apex of prosternum; 2nd antennal segment slightly longer than
basal spine of scutellum and anterior lobe of pronotum . . . . . . . . . . . Polytoxus sp. A
290' Hemelytra with vein M at apical margin of discal cell angulate, small vein arising at angle
and extending for short distance into cell (Fig. 552); proboscis reaching apex of proster-
num; 2nd antennal segment as long as basal spine of scutellum, subequal to length of
anterior lobe of pronotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polytoxus sp. B
291(287') Femora, head, pronotum, scutellum, and abdominal margins spinose (Fig. 553); head
with 2 dorsally long horn-like spines on bases of antennae; antero-dorsal half of
pronotum with 2 long erect spines, posterior half with 6 (2 laterals each behind
transverse groove and 4 in a row before posterior margins); scutellum with 3 spines; leg
I barbed with long and short spines dorsally and ventro-laterally in femora and ventro-
laterally in tibiae, brownish yellow . . . . . . . . . . . . . . . . . . . . Polididus armatissimus Stål
291' Not as above: abdominal margins without spines . . . . . . . . . . . . . . . . . . . . . . . . . . . 292

292(291') Apex of scutellum broad with 2 or 3 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
292' Apex of scutellum triangular or subtriangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294

293(292) Apex of scutellum with 2 converging spines (Fig. 554), constricted along lateral mid-half
and with a deep cavity at center in between lateral carinae: head with a ridge-like tylus
and small and low juga; body black except whitish eyes and ocelli and band on the 2nd
basal one-fourth antennal segment III, pinkish red to pale orange-red base of embolium
and lateral margins of abdomen, venter of abdomen with 4 pinkish red transverse bands
on sternites II-V; tibiae I and II with very short spongy membrane . . . . . . . Scadra sp.
293' Apex of scutellum with 3 spines, inner one very small and outer ones nearly parallel with
each other (Fig. 555); pronotum clearly bilobed with a cross groove at middle, posterior
lobe with sublateral longitudinal grooves (Fig. 556); body black except reddish ab-
dominal margins, entire abdominal venter I to V red except black bases, rest black; legs
I and II blackish brown including coxae and trochanter, apices of tibiae slightly swollen
and bent downwards, ventrally with a moderately concave spongy membrane; ventral
apical one-third of all femora with a small blunt tubercle . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectrychotes crudelis (Fabricius)
294(292') Hemelytra with a quadrangular cell at the inner end of corium near base of membrane
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
294' Hemelytra without a quadrangular cell at the inner end of corium near base of
membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298
295(294) Head with 6 anteocular spines (Fig. 557) and 6 postoculars; anterior lobe of pronotum
with 4 long and a number of small spines; postero-median margins with short spines,
overhanging on top of scutellum; femora I hirsute with a long and 2 short blunt spines
dorsally; venter of tibiae I with 2 rows of short blunt teeth; yellow-brown to brownish
orange with blackish membrane and hindpart of head . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scipinia horrida Stål
295' Head, pronotum, and legs without such spines; body coloration commonly red and
black and structure of hemelytra different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
296(295') Abdomen with broad lateral and thin apico-sternal white bands; red with black legs,
antennae, proboscis, membrane, claws and base of corium, scutellum, ante-ocular area
up to base of antennae and post-ocular except for a red and yellow band in between
ocelli, antero-median part of anterior lobe and apical half of posterior lobe of pronotum
(Fig. 558), meso-pleuron and part of meta-pleuron; ocelli small, set closer to eyes than
to each other; pronotal humps small; cells of membrane almost equal in diameter
(Fig. 559) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinocoris fuscipes (Fabricius)
296' Abdomen, head (except white underneath) and legs entirely black; anterior lobe smooth
and with humps, posterior lobe coarse; pronotal humps large and more rounded; inner
cell larger than the dorsal one . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
297(296') Entire pronotum black dorsally and laterally, humps of anterior lobe wide apart (Fig.
560); scutellum with yellow carina and apical spine; quadrangular cell (Fig. 561) small
twice longer than wide (10:5); prosternum and all coxae black; 1st antennal segment 2.7
times longer than 2nd . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinocoris sp. A
297' Pronotum red or with posterior lobe blackened in between the 2 longitudinal grooves
near humeri (Fig. 562); scutellum red including apical tip; quadrangular cell (Fig. 563)
large 1.5-1.7 times longer than wide; prosternum and coxae I reddish; 1st antennal
segment 2.8 times longer than 2nd . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinocoris sp. B
298(294') Hemelytra with a 6-sided discoidal cell; body depressed to moderately flat . . . . . . 299
298' Hemelytra without a discoidal cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
299(298) Segment I of proboscis longer than combined length of II and III slightly beyond
hind-margins of eyes; head one-third shorter than 1st antennal segment, ventro-lateral
and postero-lateral areas of post-ocular region lined with spines and postero-median
with a pair of tubercles directed posterad; pronotum nearly as long as 1st antennal
segment and subtriangular with a broad V-shaped anterior part (Fig. 564); dull brown
with femora I-III and antennal segment I mottled yellowish white; tibiae brownish along
mid-half and both ends but yellow in between; body elongate and depressed . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pygolampis sp.
299' Segment I of proboscis not reaching hind-margins of eyes, short or just as long as
combined length of II and III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
300(299') Femur I with a single row of 31-34 (11 moderately large and 20-23 small ones) ventro-
apical teeth; discoidal and outer cell of membrane, central part of clavus, and inner part
of corium castaneous (Fig. 565); first antennal segment length less than half of segment
II; eyes indented behind at mid-half (Fig. 566). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
300' Femur I with 2 rows of ventral spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
301(300) Subgenital plate markedly narrowed in the basal one-third, concave basally and cleft
apically, plate with about 30 hairs (Fig. 567); paramere with subtruncate flat inner
process, subapical longest setae slightly longer than paramere diameter (Figs. 568-569);
inner process slightly serrated; tip of scutellum curved upwards; 1st antennal segment
0.42 length of 2nd (Fig. 566); outer distal margin of discoidal cell bisected by the middle
vein of membrane a little below mid-half (Fig. 570) . . Oncocephalus pacificus Kirkaldy
301' Subgenital plate broadly recurved medially, gradually narrowed basally and nearly
straight posteriorly, plate with about 40 hairs (Fig. 571); paramere’s subapical longest
seta as long as paramere diameter (Fig. 572); inner process flat with angled sides (Fig.
573), evenly serrated with minute teeth; tip of scutellum projected forward; 1 st antennal
segment 0.46 length of 2nd; outer distal margin of discoidal cell bisected by the middle
vein of membrane at mid-half (Fig. 574) . . . . . . . . . . Oncocephalus impudicus Reuter
302(300') Pronotum 1.8-1.9 times longer than its greatest width, 1.3 times longer than 1st antennal
segment and 1.2 times longer than head (Fig. 575); head with a pair of tubercles in
between antennal bases, and spines along ventrolateral and posterolateral portion of
head; back of head with 2 small tubercles in between the groove; proboscis without
spines; femur I moderately and uniformly swollen with 2 outer and 3-6 inner large teeth
and a number of small ones in between; apex of femur III almost at tip of abdomen;
discoidal cell rectangular; straw-colored except legs with brown striae . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sastrapada entomophaga Miller
302' Pronotum about as long as broad (Fig. 576), 2 times longer than 1 st antennal segment
and one-sixth longer than head; without tubercles between antennal bases, 3 spines on
each ventro-lateral part of ante-ocular area (Fig. 577); back of head smooth without
spines; proboscis with spines in segments I and II; femur I broadest medially with 3 outer
and 4 inner prominently long spines and a number of small ones in irregular row in
between the big spines; apex of femur I about 1-2 segments short at tip of abdomen;
discoidal cell subglobular yellow-brown. . . . . . . . . . . . . . . . . . . . . . . . Staccia diluta Stål
303(298') Pronotum constricted near mid-half, swollen anterior part with blackish band medially,
laterally and near margins; posterior half brown and with a black humeral spine; head
with a porrect tubercle in between bases of antennae and a pair of setae near antennal
base; lustrous eyes strongly diamond-like; ocelli with orange ring (Fig. 578); legs banded
dark brown in apical half and yellow in basal half; venter of femur with 2 spines in leg I
and one each in II and III; tarsus I with simple claw (Fig. 579) and 2-segmented; margins
of abdomen spinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lisarda sp.
303' Pronotum constricted below mid-half, anterior lobe and subglobular and posterior lobe
more or less transverse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304
304(303') Tibia I with spongy furrow, II normal; head long and porrect; antennae far in front of
slightly oblique eyes by a little less than 1 eye diameter (Fig. 580); anterior lobe of
pronotum with 2 longitudinal grooves on each side of the median groove, submedian
grooves fused medially before transverse groove forming elongate mark; lateral grooves
curved towards pronotal margins (Fig. 581); outermost cell of membrane large;
brownish yellow with black eyes, posterior lobe of pronotum, scutellum, forewings
except yellow apex of membrane and clavus and base of corium, and reddish brown
sides of head, thorax and abdomen (Fig. 582) . . . . . . . . . . . . . . . Sirthenea flavipes Stål
304' Tibiae I and II with spongy furrow; head of moderate length and slightly curved
downwards; general color black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305
305(304') Spongy furrow (Fig. 583) occupies barely half length of tibiae; forewings without ovoid
or subglobular yellow spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306
305' Spongy furrow occupies almost apical three-quarters of tibiae; forewings with ovoid or
subglobular yellow spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
306(305) Apical half of corium yellow (Fig. 584) except blackish brown base; base of forewings
blackish brown; inner area of cells of membrane brownish black with slight violet tinge,
large middle cell with a bifurcated vein arising at its distal end with a common point;
basal cell small and long, width of cell at distal end 0.5 times that of cell above it;
antennal segments I and II black, segment II slightly shorter than anterior lobe of
pronotum; proboscis black (Fig. 585) . . . . . . . . . . . . . . . . . Pirates atromaculatus (Stål)
306' Corium, embolium, and base of clavus yellow; inner area of cells of membrane light to
dark brown, large cell of membrane bifurcated distally not at a common point, distal
oblique vein 5 times longer than distal lateral width of cell below (Fig. 586); antennal
segments I and II brownish black, segment II a little longer than anterior lobe of
pronotum; proboscis reddish brown (Fig. 587) . . . . . . . . . . . . . . Pirates sparsus Miller
306'' Basal half of corium, clavus and scutellum dark cinnamon brown; membrane with an
arched (bow-like) transverse band near base and a rounded subapical spot (Fig. 588);
segment II of antennae about as long as anterior lobe of pronotum. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirates arcuatus Stål
307(305') Corium with a large ovoid yellow spot and a small light yellow spot in the subtriangular
inner basal cell of the membrane area; 2nd antennal segment slightly shorter than length
of anterior lobe of pronotum; lateral edge of abdomen with large black areas in between
yellow spots (Fig. 589); proboscis segment II one-fourth longer than ante-ocular area.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectomocoris biguttulus Stål
307' Corium without large yellow spot; inner basal cell of membrane with a large yellow spot
at center (Fig. 590); 2nd segment of antenna one-third longer than anterior lobe of
pronotum; proboscis segment II 1.14 times longer than anteocular area (Fig. 591);
lateral edge of abdomen with narrow black spot in between yellow spots . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectomocoris atrox Stål
308 Beak 3-segmented and long; front legs not raptorial; arolia absent . . . . . . . . . . . . 309
308' Beak 4-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 310
309(308) Membrane of hemelytra with 4 elongated closed cells; head not pointed viewed dorsally;
eyes converging frontally; pedicel longer than segment III or IV; pronotum convex
above scutellum and narrowed anteriorly; scutellum large and pointed; black except
whitish membrane (Fig. 592) [Saldidae] . . . . . . . . . . . . . . . . . . Saldula ornatula Reuter
309' Membrane of hemelytra without closed cells; corium with dark brown thickened veins
forming 3 whitish cells; head acute frontally; eyes not convergent; pedicel shorter than
segment III or IV; pronotum straight at base with shallow suture in the apical one-third
scutellum small, truncated apically and with a cavity before apex; light brown with dirty
white membrane (Fig. 593) [Mesoveliidae] . . . . . . . . . . . . Mesovelia vittigera Horvath
310(308') Arolia absent; front leg enlarged, raptorial and with spines; hemelytra with many
marginal veins [Nabidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311
310' Arolia present; front legs and hemelytra variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
311(310) Mid- and hind-femora with spines much longer than femora width (Fig. 594); femora I
and II with 5 ventral and 5-6 inner lateral spines; tibiae I and II with 9-10 long thin spines,
venter of II with 13-18 black short spines in a row; head with a pair of long trichobothria
between ocelli and inner margins of eyes, black with yellowish white marks behind base
of antennae, sides and front of ocelli; ocelli elevated and near each other; segment II &
IV of proboscis as long as segment II of antenna (Fig. 595); collar black dorsally and
white ventrally; generally blackish brown with three longitudinal white to yellow marks
6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arbela nitidula (Stål)
311' Mid- and hind-femora without such type of spines; tibiae I and II with 2 close rows of
short, dark and tooth- like ventral spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312
312(311') Basal one third of pronotum with prominently transverse and zig-zag punctations;
connexivum beneath abdomen not ventrally deflexed, not demarcated by deep lon-
gitudinal furrow; segment I & II of antenna slightly longer than head and prothorax;
anterior lateral side of prothorax pointed posteriorly (Fig. 596); ocelli not elevated and
wide apart; scutellum with a brownish black median longitudinal band; femora I with
9-10 brown patches laterally; tibiae I and II with two rows of black small spines ventrally;
dull straw-colored to pale brown-yellow with pleuron brownish red. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenonabis tagalicus Stål
312' Posterior lobe of pronotum weakly and shallowly punctate; connexivum ventrally
deflexed beneath, very distinctly demarcated from abdominal sterna by a deep lon-
gitudinal furrow; sterna without ventro-lateral rows of shiny hairless areas . . . . . . 313
313(312') Shaft of males’ left genital clasper subglobular with a narrow cleft opposite base of
blade; tip of clasper with a protrusion (Fig. 597); tegmina uniformly pale; antennal base
near eyes; ocelli behind eyes, tip of head (juga and tylus) short; membrane cells almost
of equal level at apex (Fig. 598) . . . . . . . . . . . . . . . . . . . . . . . . . . . Nabis sinoferus Hsiao
313' Shaft of males’ left genitalia clasper elongate to subglobular but without protrusion at
apex of blade; blades acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
314(313') Tegmina nearly uniformly pale straw in color; legs feebly marked; shaft of males’
claspers subglobular with a shallow and narrow concavity opposite base of blade; tip of
blade moderately acute (Fig. 599) . . . . . . . . . . . . . . . . . . . . . Nabis capsiformis Germar
314' Tegmina slightly darker straw in color; membrane cells at different levels towards apex
(Fig. 600); shaft of males’ claspers elongate with a wide concavity opposite base of blade,
tip of blade strongly acute (Fig. 601) . . . . . . . . . . . . . . . . . . . . . . Nabis stenoferus Hsiao
315(310') Membrane of hemelytra with only 4 or 5 veins [Lygaeidae] . . . . . . . . . . . . . . . . . . . 316
315' Membrane of hemelytra multi-veined; scent glands present . . . . . . . . . . . . . . . . . . 330
316(315) Head triangular; eyes moderately small; front femora usually with spines; phytophagous
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317
316' Head transverse; eyes very large; front femora without spines; entomophagous . . 327
317(316) Abdominal suture between sternites IV and V ( 3rd visible inter-segmental suture)
curved anteriorly close to the lateral margin but not completely reaching the margin.
(Subfamily Rhyparochrominae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
317' All ventral abdominal intersegmental sutures straight and extended to the lateral mar-
gins; abdominal spiracles of segment VII laterally or ventrally located, the rest situated
dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
318(317) Sides of postocular portion of head parallel for some length; segment I of antennae
exceeding tylus by more than half its length; head rarely longer than wide across eyes,
dorsal surface with 4-7 pairs of scattered trichobothria; corium without a costal spot
posterior to middle; pronotal collar narrower at sides; pronotum impunctate anteriorly
(Fig. 602); femora solid color, with 2 strong ventral teeth; tibiae with only one (Fig. 603)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pamerana nigritula (Walker)
318' Sides of postocular part of head narrowed or rounded proximally; head width across
eyes equal to or greater than length; segment I of antennae exceeding tylus by less than
half its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319
319(318') Hindwing with vein R strongly curved anteriorly, without reaching margin, head,
pronotum, and abdomen usually yellow-brown to dark brown; aedeagus without spines
and processes except for holding sclerites; total length 3-3.5 times maximum width . . .
....................................................................... 320
319' Hindwings vein R reaching anterior margin; aedeagus with spines and processes but
without holding sclerites, total length 3.5-5.6 times maximum width . . . . . . . . . . . 323
320(319) Pronotum uniformly black; inner angle of corium with distinct whitish or pale spot;
clavus without the regular rows of punctures; apex of scutellum yellow (Fig. 604);
proboscis reddish brown except yellow segment II, short and barely beyond coxa I;
femora reddish brown in apical half-one-fourth, bear 7 spines (4 inner and 3 outer) (Fig.
605) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachybrachius pacificus Stål
320' Pronotum never uniformly black; spot in the inner angle of corium not distinct . . . 321
321(320') Pronotum black except the dark brown posterior one-third; spot in the inner angle of
corium not clear; femora yellow; antennae dark yellow; apex of scutellum brown, clavus
with 3 regular rows of punctures (Fig. 606); proboscis long and yellow nearly reaching
coxa II; femora yellow with 5 teeth (Fig. 607); female venter yellow. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachybrachius inortatus (Walker)
321' Pronotum uniformly brown or black reddish brown; femora and antennae yellowish;
femur I with or without band; proboscis may or may not reach coxa II; spermathecae
with or without apical bulb and multi-coiled; female venter never yellow . . . . . . . 322
322(321') Collar yellowish brown; scutellum brownish along sides; pronotum dark reddish brown;
hind part with 4-5 black patches and evenly punctured surfaces (Fig. 608); fore-femora
with or without a broad, complete fuscous band in the apical half; segments II and IV of
antenna subequal; basal segment of proboscis usually ending noticeably before base of
head, 3rd segment about one-fourth longer than 4th (Fig. 609); coloration; dark brown
to castaneous; spermatheca with 8-10 and 4-6 loose coils; apical bulb absent . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . Pachybrachius nigriceps (Dallas)
322' Collar dark brown; pronotum blackish brown, hind part without black patches and less
irregular punctures (Fig. 610); fore-femora yellow without fuscous band; segment II of
antenna longer than IV, basal segment of labium reaching head; 3rd segment of labium
rarely longer than the 4th (Fig. 611); spermatheca short, coiled portion with 4 loose
coils, apical bulb present . . . . . . . . . . . . . . . . . . . . . . . Pachybrachius sobrinus (Distant)
323(319') Corium with a prominent transparent to white spot in between its brown band (Fig.
612); fore-tibiae unarmed in males with more evenly scattered spines (Fig. 613); fore-
femora solid color either castaneous or black for at least distal half or meso- and
meta-thoracic femur black; sides of ante-ocular part of head between eye and antennal
base parallel; head surface between eyes flat; posterior lobe of pronotum with 4 luteus
spots, 2 medians and 1 each near lateral angle; fore-coxae armed with 1 spine in both
sexes; fore-trochanter spineless in males . . . . . . . Paraeucosmetus pallicornis (Dallas)
323' Corium with 2 nearly interconnected transparent spots; costal margin of corium narrow-
ly pale with a post-median inter-costal grayish brown spot and entirely black venter (Fig.
614); head about as wide as long across eyes; pronotal collar distinct with a deep
constriction; head, pronotum, and scutellum sparsely covered with long erect hairs;
posterior lobe of pronotum solid black; fore-tibiae in males shallowly curved towards
proximal end and armed with variously developed spines around mid-half; femora with
6 spines at apex (Fig. 615) . . . . . . . . . . . . . . . . . . . . . . Horridipamera nietneri (Dohrn)
324(317') Flat bugs with macropterous and brachypterous forms; scutellum whitish but not head
and thorax; corium and clavus not pitted or at most with weak, scattered punctures;
apical coria1 margin concave; dorsal abdominal gland openings at the posterior margins
of terga IV and V only; fore-femora (Fig. 616) with 2 paired large tubercular process;
fore-coxal cavities open; antennae clavate, segment II as long as III (Fig. 617) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dimorphopterus cornutus novaequineae Ghauri
324' Not as above, corium and clavus pitted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
325(324') Compound eyes (Fig. 618) borne on a tubercle; wing (Fig. 619), corium with a blackish
brown costal band, punctured basal and lateral margins, and a long and short row of
punctures below it; membrane with 2 median and 1 laterally projected dark brown
bands; antennal segments long except base, IV longer than rest of segments, II as long
as III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ninus insignis Stål
325' Eyes not as above; corium transparent without dark brown bands . . . . . . . . . . . . . 326
326(325') Head, thorax, and antennae hairy; ridge on pronotum and scutellum absent (Fig. 620);
corium with a row of punctures; antennal segment III shorter than II, IV slightly swollen
(Fig. 621); body dark brown . . . . . . . . . . . . . . . . . . . . . . . . . Cymonius turaensis (Paiva)
326' Head, thorax, and antennae not hairy; mid-anterior half of pronotum and mid-scutellum
with a whitish longitudinal ridge; tylus sharply pointed and much longer than juga
bisected by base of antennae; segment III of antennae twice as long as segment II (Fig.
622); corium (Fig. 623) uniformly punctured throughout . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymodema basicornis (Motschulsky)
327(316') Head with an inverted broad T-shaped black to brown band; antennae dorsally whitish
yellow with blackish base of IV, basal half of III, basal two-thirds of II and basal half of
I; pronotum with a small yellow apico-median and postero-median spot; yellow area of
humeri extended to hind level of eyes (Fig. 624); enclosing 16 brown punctures mostly
on margins; posterior half of pronotum brownish; scutellum, pronotum, and head
almost equal in length; body length 4.4 times longer than pronotum or head length . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocoris jucundus (Fieber)
327' Head almost uniformly yellow to brownish yellow with a black patch between ocelli and
hind part of eyes or entire hind part of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
328(327') Posterior part of head black; high above ocelli and concave in between (Fig. 625) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocoris ochropterus (Fieber)
328' Posterior part of head black only in between ocelli or hind part of eyes . . . . . . . . . 329
329(328') Scutellum (Fig. 626) with a transverse dark yellow band at base; pronotum yellowish
around margins and broadly brownish red medially; yellow area of humeri extended to
the entire oblique apical margins of pronotum enclosing 5-8 punctures almost medially
(Fig. 627); eye height 0.5 times eye length and as long as 3rd antennal segment . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocoris sp.
329' Scutellum uniformly black (Fig. 628); narrow posterior margins and broad lateral area
of protonum yellow; humeri area yellow extended to lower half of the oblique antero-
lateral area of pronotum enclosing 13 punctures (Fig. 629); eye height 0.6 times eye
length and slightly longer than 3rd antennal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocoris flaviceps (Burmeister)
330(315') Head narrower and shorter than pronotum; bucculae long, behind level of antennifers;
hind coxae more or less rounded or square [Coreidae] . . . . . . . . . . . . . . . . . . . . . . 331
330 ' Head nearly as wide and as long as pronotum, bucculae very short, rarely reaching level
of antennifers; hind coxae more or less elongated [Alydidae] . . . . . . . . . . . . . . . . . 332
331(330) Pale brown with a pair of tiny white spots on the forewings; humeri longer, protruded
laterally and sharply pointed; 10-12 mm long (Fig. 630) . . . . . Cletus punctiger (Dallas)
331' Dark to light brown with a pair of small white spots on the forewings; body more
coarsely punctate; humeral spine sharply pointed; 8-8.5 mm long (Fig. 631) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cletus trigonus (Thunberg)
332(330') Hind femora somewhat stout, with several or many spines on ventral surface; humeral
angle drawn out into a spine; rostrum reaching mesothoracic coxae (Riptortus) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
332' Hind femora elongated, unarmed, never stout, head long, jugae porrect produced into
front of tylus and longer than it; pronotum long; lateral angles unarmed (Leptocorisa)
....................................................................... 335
333(332) Meso- and meta-pleuron with paired semi-globular white spots (Fig. 632); tips of
antennal segments I, II and III brownish (Fig. 633); body somewhat rust brown . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Riptortus pedestris (Fabricius)
333' Meso- and metapleuron with rectangular-like whitebands . . . . . . . . . . . . . . . . . . . 334
334(333') Antenna solid brown (Fig. 634); body rust brown . . . . . . Riptortus pilosus (Thunberg)
334' Apical half of antennal segments II and III and entire length of segment I dark reddish
brown (Fig. 635); lateral side of whole insect with a broad longitudinal white band
(Fig. 636) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Riptortus linearis (Fabricius)
335(332') Rostrum extended beyond apices of 2nd coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
335' Rostrum never beyond 2 nd coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
336(335) Lateral dark line well developed (Fig. 637); venter of abdomen with a median dark
streak extended from 1st to 6th segment of rostrum but reaching only midway between
2nd and 3rd coxae (Fig. 637); basal quarters of apical antennal segments never pale;
parameres large (Fig. 638); body length 15 mm . . . . Leptocorisa solomonesis Ahmad
336' Lateral black line faintly represented by 2 blackish brown dots; one on sides of anten-
niferous tubercles and on collar, never beyond pronotal collar (Fig. 639); disc of
pronotum never blackly punctured, sometimes with greenish tinge; anterior tip of head
cleft (Fig. 640); paraclypeae distinctly elongated, pointed and cylindrical . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa palawanensis Ahmad
337(335') Endocorium and clavus black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338

337' Endocorium and clavus pale to light brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339
338(337) Abdominal tergites solid color; hemelytra dark black except the costal margins above
the membrane; claspers (Fig. 641) with truncated apices and reaching well beyond
pygophore; paraclypeae short, thick basally and pointed at apices . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa biguttata Walker
338' Abdominal tergites V-VI1 black (or basal one-third) (Fig. 642); apical segments of
antennae pale distally and proximally; paraclypeae cylindrical, pointed and knob-like at
apices; base of clasper narrow, strongly and obliquely truncate (Fig. 643) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa costalis (Herrich Schaffer)
339(337') Ventro-lateral brownish black spots present in the abdomen (Fig. 644); claspers basally
broad, tapering to a flat point with curved extremities; in females, proximal margin of
abdominal sternite VII deeply notched . . . . . . . . . . . Leptocorisa oratorius (Fabricius)
339' Ventrolaterals of the abdomen and claspers not as above . . . . . . . . . . . . . . . . . . . . 340
340(339') Body length almost 13 mm; paraclypeae short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
340' Body length usually more than 14 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342
341(340) Tibiae and antennae dark; paraclypeae thin medially and robust basally; apices of
parameres pointed to each others’ base at resting position (Fig. 645); proximal margin
of abdominal tergum VII truncated . . . . . . . . . . . . . Leptocorisa pseudolepida Ahmad
341' Tibiae and antennae pale to light brown; paraclypeae (Fig. 646) thick medially and thin
basally, blunt and straight; claspers with strongly pointed apices, blades very short
(Fig. 647) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa luzonica Ahmad
342(340') Segment I of antennae longer than 3 times the length of 1st rostral segment; body pale
yellow-brown; claspers (Fig. 648) with very long cylindrical and pointed blades . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa tagalica Ahmad
342' Antennal segment I shorter; parameres different . . . . . . . . . . . . . . . . . . . . . . . . . . . 343
343(342') Callosities near pronotal posterior angles large and dark; basal joint of antennae dark
brown to black; lateral black line runs from bases of antenniferous tubercles to collar
sides; claspers truncated apically (Fig. 649); body length 16.8 – 18.0 mm ............
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa chinensis Dallas
343' Callosities small and pale; basal joint of antennae pale and solid color; lateral black line
only a black dot on the sides of collar; claspers bifid at apices (Fig. 650); body length
15-16 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocorisa acuta (Thunberg)
344( 180) Cerci sclerotized, unsegmented and modified into terminal forceps; forewing reduced
to small tegmina, hindwing large, membranous, semicircular, and fan-like; abdomen
long and freely movable; legs short and cursorial with 3-segmented tarsus (DERMAP-
TERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
344' Cerci normal; wings and abdomen not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348

345(344) Tarsal segment 2 prolonged into a narrow lobe beneath the distal segment; usually
blackish to reddish species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 346
345' Notasabove . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
346(345) Black to dark reddish brown; antenna black except segments XIV-XVII (Fig. 651);
yellow; elytra black, inner apical part of 1st pair protruded; pronotum slightly transverse
(Fig. 652); branch of male forceps broad and with inner margins toothed or arched;
pygidium of females transverse, distal part not narrowed . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelisoches morio (Fabricius)
346' Yellowish brown to reddish light brown; antenna solid light brown (Fig. 653); elytra
yellow with dark reddish brown bands on both lateral sides; pronotum longer than broad
(Fig. 654); 3rd abdominal segment with a black small tubercle; pygidium of females
short, distal part narrowed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proreus simulans (Stål)
347(345') Hind-margin and angles of pronotum broadly rounded (Fig. 655); sides of abdominal
segments slightly blunt and not keel shaped; parameres with external apical angle sharp
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euborellia philippinensis Srivastava
347’ Hind-margin of pronotum rather square (Fig. 656); sides of 7th and 9th abdominal
segments keel shaped; external apical angle of parameres broadly rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euborellia annulata (Fabricius)
348(344') Two pairs of wings subequal; sterna II and III of males with complex accessory genitalia
(ODONATA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
348' Wings not subequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362
349(348) Large dragonflies, 27-42 mm long; discoidal cell divided longitudinally into triangles and
supratriangles; wings held horizontally in repose, hind-wings broader than fore-wings
and rounded anally in both sexes; triangles of wings different, pointed rearward in the
fore-wings, and elongated lengthwise and nearer to arculus in the hindwing; eyes
globular, touching or moderately separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
349' Moderately small 18-32 mm long, slender bodied damselflies; fore- and hind-wings
similar in shape and venation; discoidal cell quadrangular, not longitudinally divided;
eyes wide apart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 358
350(349) Hind-wings very greatly broadened basally; 2 cubito-anal cross-veins present; 2nd
cubito-anal of the hind-wing forms a prominent subtriangle; stigma trapezoidal; vein M2
strongly wavy; entire basal area of hind-wings somewhat yellow (Fig. 657); slightly
swollen basal segments of abdomen with extra transverse carinae . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pantala flavescens Fabricius
350' Hind-wing moderately broadened basally; other characters not as above . . . . . . . . 351
351(350') Distal ante-nodal nervure of forewings complete; A2 opposite anal crossvein; prothorax
with a very large, erect notched posterior lobe fringed with long hairs . . . . . . . . . 352
351' Distal ante-nodal nervure of fore-wings incomplete . . . . . . . . . . . . . . . . . . . . . . . . . 353

352(351) Triangle of hind-wing crossed (Fig. 658); wings smoky transparent with light brown
coloration at apices, basal wing spot large reaching 2 nd ante-nodal and tornal angle of
wing in males; pterostigma dark reddish brown covering 2 1/2 cells; frons reddish
frontally; prothorax and thorax reddish brown similar to legs; distal ends of femora and
inner surfaces of tibiae blackish brown; abdomen basally broad, tapers apically, strongly
keel shaped and bright scarlet red; lamina of male genitalia naked or with few long
golden hairs on its outer surface; borders of segment VIII markedly dilated and bor-
dered broadly with black apical border with small ventral plate; short triangular vulvar
scales bent slightly ventrally and apically to body axis in females’ genitalia . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthetrum testaceum (Burmeister)
352' Triangle of hind-wing free from crossveins; wings smoky transparent with slight marks
at apices and borders of wings; pterostigma black with yellow-brown median, covers 2
cells (Fig. 659); frons yellow, very deeply notched to form two triangular facets in front;
prothorax bright yellow, thorax and abdomen greenish yellow with black marks; seg-
ments IV to VI with a broad oval dorsal black spot on basal one-third continued finely
along mid-dorsal carina to join with very broad apical black rings on IV and V; males’
genitalia with a mass of long hairs; females’ vental plate of segment VIII with a small
convex notch the ends of which project slightly ventral wards; 9 th ventral plate keel
shaped basally and swollen at apex . . . . . . . . . . . . . . . . . . . . Orthetrum sabina (Drury)
353(351') Prothoracic lobe large and fringed with long hairs; eyes contiguous for a short distance;
frons non-metallic dorsally; costal border offore-wing (Fig. 660) straight, discoidal field
begins with a row of 2 cells; fore-wing triangle crossed; subtriangle with 3 cells; hindw-
ings discoidal cell entire . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplacodes trivialis Fabricius
353' Prothoracic lobe small and usually naked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
354(353') Borders of discoidal field strongly converging at wing margins; antenodal crossveins of
1
fore-wings and hind-wings with 12-2 and 10, respectively; discoidal cell of fore-wing
broader, with its costal side 0.5 the length of basal; 2 rows of cells above radial planate;
no supplementary nervure; IR2 between R2 and R3; A2 smooth; stigma reddish, short
with 2 crossveins below (Fig. 661); frons red dorsally . . . Trithemis aurora Burmeister
354' Borders of discoidal field parallel or widely divergent at wing margins . . . . . . . . . 355
355(354') Wings with small basal yellow marking; usually 1 row of cells above radial planate;
stigma long with 2 crossveins underneath; discoidal field widens at edge of wing;
fore-wings with 9-21 – 10 -12 ante-nodal nervures; eyes shortly contiguous (Fig. 662); face
and frons red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crocothemis servillia (Drury)
355' Wings broadly black or dark reddish brown from basal half - two-thirds with half — two-
thirds transparent area before or after stigma in males; females with a large spot at
nodus and apex broadly darkened as in N. tullia tullia . . . . . . . . . . . . . . . . . . . . . . . 356
356(355') Basal half of male fore-wings broadly black and opaque area in apical one-half (Fig.
663); females with large black spot at nodus and apex broadly darkened (Fig. 664) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neurothemis tullia tullia (Drury)
356' Wings dark reddish brown from base up to part of stigma . . . . . . . . . . . . . . . . . . . 357
357(356') Hind-wings with one cubito-anal crossvein; brown area in both wings reaching pteros-
tigma (Fig. 665) abbreviated in hind-wing on posterior side and rounded externally . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neurothemis palliata Rambur
357' Hind-wings with 2 cubito-anal crossveins; brown area broadly squared at the base of
stigma in both wings (Fig. 666) . . . . . . . . . . . . . . . Neurothentis terminata terminata Ris
358(349') Arculus well beyond Ax2; labrum metallic blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
358' Arculus slightly beyond Ax2 ............................................. 360
359(358) Superior anal appendages shorter than inferiors and 3-spined (2 apically and 1 basally)
(Fig. 667); inferior part parallel-sided laterally and with hairs at inner side (Fig. 668);
lower sides of prothorax bluish green and blue on the anterior and posterior lobe;
median of posterior lobe slightly developed; fore-wing with 5-6 postnodal nervures and
4 in the hind-wing . . . . . . . . . . . . . . . . . . . . . . . . . Agriocnemis femina femina (Brauer)
359' Superior anal appendages longer than inferiors and without spines or tubercles (Fig.
669), inferior part with inwardly curved tips (Fig. 670); anterior lobe, lower sides and
extreme part of posterior lobe apple green; posterior lobe trilobate with midlobe
produced backwards; fore-wings (Fig. 671) with 6-7 postnodal nervures, and 5-6 in hind
wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agriocnemis pygmaea (Rambur)
360(358') Anal bridge arising from the hind border of wing, more or less below the point where it
meets anal crossing (a short nervure at base of wing extending from Cu2 + IA (1st anal)
to the anal bridge or hinder part of wing); pterostigma of male fore-wings and hind-
wings different in shape and size with 8 postnodals in the fore-wings (Fig. 672); segment
X of males with a pair of apical tubercles dorsally (Fig. 673); postocular spots present;
thorax with distinct black markings; body black and green, segment VIII of male
abdomen bright blue and reddish to dull green in females; abdominal length 25 mm . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ischrura senegalensis (Rambur)
360' Anal bridge arising from the hind border at the point where anal crossing meets it . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
361(360') Segment VIII of females with an apical ventral spine; pterostigma of fore-wings almost
twice the size of that in the hind-wings but both blackish grey; costal and distal borders
broadly white covering three-quarters of cell; fore-wings with 9 postnodal nervures and
8 in the hind-wings; abdomen of males extremely slender and comparatively long, I-III
pale azure blue, IV-VII pale yellow and VIII-X blue; X with an X-shaped black dorsal
spot and broadly blue on sides; anal appendage black, superiors 0.5 length of segment
X and apex inwardly curved (Fig. 674); inferior part small, rounded tubercles (Fig. 675);
females' anal appendages very small, conical and black; vulvar scales not conspicuous
and pale blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aciagrion occidentale Laidlaw
361' Segment VIII of females without an apical ventral spine; pterostigma of forewings very
oblique, diamond shape, brownish black with outer angle and costal border narrowly
white, tinted with blue on the upper surface of wing; fore-wings with 14 postnodal
nervures (Fig. 676) and 12 in the hind-wings; abdominal dorsum dark metallic blue,
venter brown to brownish yellow; abdominal segment IX and X with inverted yellow
V-shaped marks laterally . . . . . . . . . . . . . . Pseudagrion pilidorsum pilidorsum Brauer
362(348') Wing veins with marginal branching; wings broad with blunt to slightly rounded apices;
Rs slightly wavy and inner series of gradates of both wings converged with Rs; subcosta
and radius in hind-wings of male fused below stigma; male stigma heavily developed,
narrow in the fore-wings, and wider and often brown-pigmented, never fused with
radius in the hind-wings (Fig. 677); adults green with conspicuous orange-yellow mid-
dorsal stripe; larvae trash-carriers, head with 6 prominent reddish brown stripes, middle
4 forming Y-shape; body with 3 longitudinal brownish stripes (NEUROPTERA)
[Chrysopidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chysopa basalis Walker
362' Wing veins without marginal branching; larvae not trash-carriers . . . . . . . . . . . . . 363
363(362') Fore-wings form a highly chitinous or leathery non-folded, rigid plate called elytra
(COLEOPTERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364
363' Fore-wings elongate membranous without hamuli and shed off by basal sutures, or
fore-wings and hind-wings coupled by hamuli (hook structures) during flight; some
species wingless with characteristic node forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . 485
364(363) Notopleural suture in the prothorax absent; wings without an oblongum; hind-coxae
movable, separated from the meta-sternum and never dividing basal abdominal sternite
into 2 lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365
364' Notopleural suture present; wings with an oblongum; hind-coxae fused to the meta-ster-
num dividing the basal abdominal sternite into 2 lateral lobes . . . . . . . . . . . . . . . . 426
365(364) Posterior coxae not reaching elytra; meta-sternum with a transverse suture; terrestrial
beetles with legs adapted for crawling or running . . . . . . . . . . . . . . . . . . . . . . . . . . . 366
365' Posterior coxae reaching elytra; meta-sternum without a transverse suture; hind-leg
hairy and adapted for swimming; coxa III large without longitudinal plates, metasternal-
metacoxal suture arched [Dytiscidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419
366(365) Antennae arising in front of head above bases of mandibles; head width across eyes
always wider than thorax; inner lobe of maxillae with movable hook at its end; clypeus
extended laterally beyond the base of antennae; elytra with white to light yellow mark-
ings or spots; tiger beetles [Cicindellidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
366’ Antennae arising more laterally on sides of head between eyes and bases of mandibles;
head width across eyes always narrower than thorax; inner lobe of maxillae without a
movable hook; clypeus not extended laterally beyond base of mandibles; elytral pattern
variable; ground beetles [Carabidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
367(366) Elytra with 6-8 white to yellow sub-globuiar spots; middle of elytra with 3-4 spots
arranged in a longitudinal row parallel to each other (also parallel transversely); humeri,
lateral, and sutural margins without marks or spots; sides of pronotum with or without
white hairs; scape with 2 setae near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368
367’ Elytral marks variable, never in a longitudinal row parallel to each other; humeri, lateral
and sutural margins with marks or spots, if not dots present and very close to each otlier
in the humeri lateral and sutural areas; apex of scape with 1-2 setae . . . . . . . . . . . 373
368(367) Sub-apical area of labrum with 7 long setae; sides of pronotum without white hairs;
elytra with 6 globular to sub-globular spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
368’ Sub-apical area of labrum with 6 long setae; sides of pronotum with or without hairs;
elytra with 6-8 sub-globular spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370
369(368) Right side of labrum with 3 sub-apical and 1 sub-median long hairs, sub-basal area with
a narrow transverse yellow band; head and pronotum slightly coarse, striped near eyes;
longitudinal mid-groove almost 0.5 times width of sub-posterior area of pronotum,
posterior one-fifth with longitudinal stripes (Figs. 678-679); median elytral yellowish
spot markedly oblique and larger than the rest, sub-apical spot smallest in males (Fig.
680) and sub-humeral spot the smallest in females (Fig. 681); length 15-17 mm . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela sp. A
369’ Left side of labrum with 4 sub-apical long hairs in a row; each side of the labrum with a
broad rectangular yellow spot, apex of labrum with 3-5 teeth in some females; head
striped farther behind eyes; front pronotum shallowly V-shaped, posterior one-fifth
without longitudinal stripes (Fig. 682); elytral white spots usually globular and equidis-
tant (Fig. 683); length 13.5-16 mm . . . . . . . . . . . . . . . . Cicindela sexpunctata Fabricius
370(368’) Sides of pronotum with white setae; males with 8 spots on elytra, basal spot very small
roughly one-third of middle spot and transverse (Fig. 684); females (Fig. 685) with 6
spots without the subbasal; subhumeral spots in both sexes slightly oblique; labrum with
3 teeth, black with a rectangular yellow spot on each side in females; yellow band of
labrum in males almost contiguous except for a moderately broad blackish brown apical
and basal median band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela sp. B
370‘ Sides of pronotum without white hairs; both sexes with 6 spots . . . . . . . . . . . . . . . 371
371(370’) Apical spots on elytra noticeably large almost equal in size to the median spots (Fig.
686) but larger than the subhumeral spots in the females; median spot the largest in
males; distances between spots unequal; posterior part of pronotum with transverse
rope-like stripes; labrum bluish with light yellow marks on each side, broad and
prominent in males (Fig. 687) . . . . . . . . . . . . . . . Cicindela near sexpunctata Fabricius
371' Apical spots on elytra smaller than the median, equal to or a little smaller than
subhumeral spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372
372(371') Elytral spots same distance from each other, subhumeral slightly oblique and oblong
(Fig. 688); pronotum as long as wide and with 2 transverse ridges posteriorly (Fig. 689);
next to last sternite of males with a moderately wide and deep median apical cleft;
longitudinal light groove of terminal sternite near the left side of the apical cleft of the
preceding sternite; length 16 mm . . . . . . . . . . . . . . Cicindela sp. C. (sexpunctata grp.)
372' Median spots closer to the subhumeral than to the subbasal spots (Fig. 690); pronotum
with midlongitudinal part with short transverse striae and oblique in the posterior part
(Fig. 691); penultimate sternite of males with a moderately deep cleft medio-apically;
longitudinal groove of terminal sternite slightly on the right side of the apical cleft of the
preceding sternite; length 14.8 mm . . . . . . . . . . . . . Cicindela sp. D (sexpunctata grp.)
373(367') Small to moderately small tiger beetles, 7-12 mm long . . . . . . . . . . . . . . . . . . . . . . . 374
373' Large tiger beetles, 13-20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380

374(373) Humeri without yellow marks or spots; sides of pronotum with white hairs, apical width
slightly less than median width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375
374' Humeri with yellow mark or spot; sides of pronotum with or without hairs; apical width
prominently narrower than median width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 376
375(374) Elytra black with 2 small yellow spots in each elytron, a pale sub-median closer to the
sutural margin and an oblique apical (= apical lunule without a leg), distinctly
punctated but not confluent, some discal areas not pitted; apical spine of elytra
developed and slightly beyond level of apical teeth (Fig. 692); labrum black, raised
medially with 7 sub-apical setae (4 in the left and 3 in the right), apico-median with a
tooth and wavy towards sides producing a small inner and wider outer concavities;
sub-apex of scape with only 1 seta (Fig. 693); pronotum (Fig. 694) cylindrical and as long
as wide, slightly rough striae converging to the median line; length 8 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela nr. kaleea Bates
375' Elytra dull black with 3 small spots near middle of elytra and an apical lunule, in some
specimens the spot in the midbasal one-third absent but a wide unpitted area around the
globular spot in the apical one-third present (Fig. 695); median tooth of apex of labrum
in a deep cavity, blunt elevations on each side of the tooth and moderately obliquely
square towards lateral sides; subapex of labrum with 6 white hairs; middle of frons with
fine zigzag stripes running backwards to the vertex; supraorbitals with 13-14 well
separated longitudinal ridges; length 8-9 mm . . . . . . . . . . . . . . Cicindela kaleea Bates
376(374') Basal half of elytra with 2 pairs of parallel yellow stripes near sutural margins, base with
a pair between humeri and margins near scutellum; humeri with a yellow band extended
up to the apical lunule, sometimes interrupted along mid-half but usually connected to
an oblique band directed to the sutural margins (Fig. 696); labrum yellow with apical
area brown, 3-5 teeth with 4 white hairs subapically (Fig. 697); length 9-11 mm . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela striolata Illiger
376' Subsutural area without parallel yellow stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
377(376') Marginal band with a “foot” and a “knee” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
377’ Marginal band reduced to spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
378(377) Humeral lunule and its slightly oblique, broad and strongly C-shaped spot occupying
more than half of width of elytron, distinctly separated from the marginal lunule; band
connecting marginal lunule to its “knee” deeply concave towards base and slightly
pointed on opposite end (apex); foot of marginal lunule square, enlarged at tip and
concave towards the knee; marginal lunule narrowly separated from apical lunule; arm
of apical lunule directed towards “foot” broadly connected to it; apex of elytron distinct-
ly higher than sutural spine (Fig. 698); length 8-11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela elisae Motschulsky
378' Humeral lunule and its spot normal C-shaped with tips almost on top of each other
occupying about half of elytra1 width and thinly connected marginally to the marginal
lunule; rounded “knee” connected to the marginal lunule by a straight transverse band
and J-shaped from “knee” to “foot”; tip of foot rounded; marginal lunule well separated
from the apical lunule; arm of apical lunule water-drop like and very thinly to almost
unnoticeably connected to it; apex of elytra lower than the sutural spine, elytra of
females broadened along marginal lunule (Fig. 699); labrum (Fig. 700) yellow except
black apex and base, apico-median with a sharp moderately long black tooth and 15
sub-apical white hairs in 2 rows; length 10-12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela sumatrensis Herbst
379(377') Elytron dull black with 6 sub-globular yellow spots (Fig. 701); 5 along lateral margins
and 1 small globular spot at mid-half close to the sutural margin; humeral spot well
isolated from the sub-humeral spot; apex of elytra with serrations small and very close
to each other towards the prominent sutural spine and apical lunule short (Fig. 702);
labrum pointed apico-medially and lined with 14 hairs (Fig. 703); females with a more
pronounced apex of elytra and oblique toward the sharply pointed sutural spine,
pronotum dark blue, without hairs along lateral margins, nearly glabrous and rounded
latero-medially (Fig. 704); length 9-10 mm (note: this is a new Philippine tiger beetle
very similar to C. ferriei) . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela nr. ferriei Fleutiaux
379 ' Elytron with 3 ovoid yellow spots in an oblique row towards the humeral spot, middle
spot thinly connected to the subapico-median spot; lateral margins with 2 smaller thinly
connected spots parallel to the median and subapico-median spots (note: these 4 spots
are well separated in the males); apical lunule complete with enlarged tips and relatively
thin connections (Fig. 705); labrum largely yellow with brownish apical margins, 3 teeth
with a large middle tooth and slightly concave towards lateral margins; sub-apical
margins with 6 long white hairs; pronotum hairy laterally, subapical and subbasal
transverse groove deep (Fig. 706); length 12 mm . . . Cicindela lacrymosa (Fabricius)
380(373') Apical lunule complete and inverted J-like; marginal and humeral lunule present or
reduced to spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
380' Apical lunule absent with only a spot near base of apical serrations; marginal without
distinct knee or reduced to spots; humeral and sub-humeral spots distinctly separated
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
381(380) All lunules complete; humeral spot slightly oblong; basal part of inverted H-like mar-
ginal lunule sharply pointed and apical lunule small Y-shaped with a moderately
enlarged inner arm directed towards the “foot” (Fig. 707); length 13 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela specularis Chaudoir
381' Elytra with 6 prominently yellow spots and an apical lunule; 3 spots in a mid-diagonal
row towards humeral spot and 2 along lateral margins (a small dot below middle
towards base and an oval large one above middle towards apical lunule); lower humeral
spot strongly oblique and the spot in the apical one-third near sutural margin transverse-
oblique (Fig. 708); both ends of apical lunule square, margins of elytra opposite lunule
evenly toothed (Fig. 709); pronotum smooth between transverse grooves and stripes
near eyes with a narrow smooth area in between (Fig. 710); labrum with a very sharp
tooth near mid end (Fig. 711); length 13-13.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela nr. lacrymosa (Fabricius)
382(380') Elytra with 3 oval or round spots and a large key-shaped marginal lunule with an
expanded band towards lateral margin and an oblique band towards the sutural margin
(Fig. 712); length 15-18 mm . . . . . . . . . . . . . . . . . . . . . . . Cicindela japana Motschulsky
382' Elytra pattern rather different; elytra with 5 spots, middle area usually with 2 spots
opposing each other in an oblique position . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
383(382') Discal spot slightly above mid-half towards apex near the sutural margin, small and
globular about 2-2.5 times its diameter away from marginal band; rest of the spots
unequal in size except the biggest marginal band; lower humeral spot longitudinal in
position; humeral and subapical (in the apical one-fourth) transverse and discal spot
slightly oblique (Fig. 713); length 17 mm . . . . . . . . . . . . . . . . Cicindela ferriei Fleutiaux
383' Disd spot ovoid-elongate and very close to the marginal middle band; length 18-20 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
384(383') Marginal middle band transversely oblong with a pointed inner tip directed towards
subglobular discal spot; subhumeral spot button-like; subapical spot subovate and
transverse (Fig. 714); length 20 mm . . . . . . . . . . . . . . . . . . Cicindela japonica Thunberg
384' Marginal middle band subglobular without a pointed inner tip; discal mark elongated
with a pointed outer tip directed towards middle marginal band; subhumeral spot
oblongate; subapical spot slightly pentagonal (Fig. 715); length 18 mm . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cicindela chinensis De Geer
385(366') Abdomen with 7 or 8 sterna visible; mandibles with a setigerous puncture in scrobe (Fig.
716); elytron truncate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
385' Abdomen with 6 visible sterna; mandibles without a setigerous puncture in scrobe;
elytra usually not truncate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391
386(385) Costae on elytra weak, faint and vaguely connected before apex (Fig. 717); black except
brownish yellow pronotum with some brown marks apicomedially and medially;
pronotum and elytra hairy; head moderately hairy, few white bristles behind eyes
latero-ventrally, and with closely punctate to network-like punctations in the frons and
vertex; labrum slightly rounded apically with 5 bristles (Fig. 718); length 6-7 mm . . . . .
............................................................... Brachinus sp.
386' Costae on elytra strong, distinct and separate apically; length 13-22 mm ....... 387
387(386') Elytra without transverse yellow spots or marks at midhalf and humeri; pronotum with
or without a pair of lateral longitudinal bands . . ... . . . . . . . . . . . . . . . . . . . . . . . . . 388
387' Elytra with prominent transverse yellow bands at midhalf and at humeri, pronotum with
a pair of yellow bands or entirely yellow with anterior and posterior margins black
interconnected by a median longitudinal black band . . . . . . . . . . . . . . . . . . . . . . . . 389
388(387) Moderately small species; length 13-15 mm; pronotum entirely black; black band of
vertex entire extended to rearside of eyes and dorsally a little above level of supraorbital
setae (Fig. 719); knees or apices of femora little, brownish red, and groove 3 with a long
subapical setae; costae I straight basally; all costae of elytra very prominent producing
clear grooves (Fig. 720); proepisterna uniformly black . . . . . . . . Pheropsophus sp. A
388' Relatively large species, length 19-22 mm; pronotum black with a pair of longitudinal
yellow bands; vertex with a black median band, its anterior part high in front of
supraorbital setae reaching midhalf level of eyes (Fig. 721); knees with prominent black
band, entire ventro-apical half blackish brown and inner apical one-fourth dark brown
in males; groove 2 with a long subapical seta; costae I slightly diverging basally; all costae
of elytra prominent producing shallow grooves (Fig. 722); central area of proepisterna
yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pheropsophus sp. B
389(387') Median transverse band of elytra zigzag-like and not reaching costae 2 or very small and
narrow reaching only groove 2 and interrupted between costae 4, 5, 7 and 8; humeral
spot extended to groove 5 (Fig. 723); entire length of epipleura yellow; pronoturn black
with a broad oblong yellow band on each side of median longtitudinal groove;
propleuron entirely yellow except black spot near coxa I, anterior and dorsal margins;
vertex with a broad median black band, its apical end strongly concave and above
midlevel of eyes (Fig. 724); length 17-20 mm . . . . . . . Pheropsophus javanus (Dejean)
389' Median transverse band of elytra broad reaching costae 2 . . . . . . . . . . . . . . . . . . . 390
390(389') Humeral spot large and subglobular containing the subbasal setae in groove 5 (Fig. 725);
pronotum with a narrow yellowish lateral band not reaching anterior and posterior
apices, widens only medially opposite midlateral setae (Fig. 726); propleuron reddish
brown with a yellow band along dorsal margin becoming subtriangular subposteromar-
ginally; vertex with a broad median band reaching entire median of frons, constricted at
middle and subbasal area (Fig. 726); length 12-21 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pheropsophus occipitalis (MacLeay)
390' Humeral spot small and elongate, below the subbasal setae in groove 5 (Fig. 727);
pronotum brownish yellow with a black I-shape band; vertex with a median band
narrowed at basal half, not reaching frons (Fig. 728); length 16 mm. ................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pheropsophus jessoensis Morawitz
391(385') Prothorax long, swollen at middle, narrowed anteriorly and posteriorly with or without
setae in the lateral groove; elytra reddish similar to prothorax; with a broad black or blue
transverse band along apical half or before apex with a pair of white spots on the apical
margins of the band (some species with another pair along the posterior margin of band
or with blue to black band on base of elytra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392
391' Prothorax subcylindrical with almost parallel sides and dorsal surfaces glabrous; shining
black to dull and punctated; prothorax usually wider than long . . . . . . . . . . . . . . . 399
392(391) Fourth tarsal segment simple, not bilobed; head impunctate; prothorax smooth or
transversely striated; head, prothorax and elytra blackish . . . . . . . . . . . . . . . . . . . . 393
392' Fourth tarsal segment bilobed; head slightly to heavily punctate; pronotum strongly
punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
393(392) Prothorax without lateral setae, apical third less punctate than the rest, lateral side
ungrooved and not bordered; head punctate except smooth vertex and hind area of eyes;
subapical hair of scape longer than entire length of scape (Fig. 729); shiny black species
with brownish epipleura and apices of elytra; intervals 5 and 6 with a longer blackened
area closer to apex; apical portion of elytra obliquely truncate (Fig. 730); length 5.5-
6mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucolliuris sp.
393' Prothorax with 1 or 6 setae on each side; elytra with a pair of white spots subapically
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
394(393') Prothorax red and fairly long with transverse striations, swollen below midhalf but
constricted before base; lateral side with 1 seta; apical half or more of elytra black with
silvery white spots below midhalf of black band in intervals 5 and 6, basal half red (Fig.
731); head black and straightly narrowed up to neck; femur I black except yellow basal
third and with few scattered spines; length 7-8.3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
....................................... Archicolliuris bimaculata (Redtenbacher)
394' Prothorax black and moderately short, shiny and smooth with 6 setae on each side, base
with 4 transverse grooves (Fig. 732); elytra shiny black with reddish brown tinge: and a
silvery white spot in interval 4 before apex of each elytron (Fig. 733); femur I light brown
and uniformly clothed with many hairs around its entire length; length 6 mm . . . . . . . .
.............................................................. Odacantha sp.
395(392') Prothorax without a pair of lateral setae; elytra with 1-2 bluish black bands and 4 white
spots ................................................................. 396
395' Prothorax with a pair of lateral setae; elytra with 2 bluish black bands and 2 white spots
........................................................................ 397
396(395) Elytra with 2 blackish blue bands, a small one at base and a broad one right below
midhalf with band length of each elytron as long as width of elytron; all 4 spots along
margins of the band; anterior spots on interval 5 twice as large as posterior spots on
intervals 4 and 5; prothorax somewhat rugose on apical and basal areas and apical
angles rectangular (Fig. 734); length 6.6-7.8 mm . . . . . . . Ophionea indica (Thunberg)
396' Elytra with 1 black band near middle, basal band absent; white spots on elytra nearly
similar in size; half of anterior spots sunken in the band and interval 5 widens along that
area to accommodate the spot; posterior spots on intervals 4 and 5 within the band;
prothorax distinctly and closely transversely rugose forming small transverse meshes
(Fig. 735); length 7-7.1 mm . . . . . . . . . . . . . . . Ophionea nigrofasciata Schmidt-Goebel
397(395') Elytra (Fig. 736) with 2 pores each on interval 5 of each elytron and 10 on interval 3,
white spot on interval 4 slightly bean-shaped and small; sides of back of head straight
and with few hairs; apical part and margins of prothorax above setae with short fine
hairs; 2 long setae noticeably below hind margins of eyes (Fig. 737); length 7 mm . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ophionea interstitialis Schmidt-Goebel
397' Elytra without pores on interval 5 and with less than 10 pores on interval 3 . . . . . . 398
398(397') Silvery white spot long, more than 0.5 times length of posterior band; interval 3 with 9
pores; back of head behind eyes moderately rounded (Fig. 738); prothorax swollen a
little above the posterior constriction; length 7 mm . . . . Ophionea ishii hoashii Habu
398' Silvery white spot small, noticeably less than half length of posterior band; interval 3 with
8 pores (Fig. 739); back of head behind eyes less rounded but not straight; prothorax
swollen below the setae (Fig. 740); length 6.4-7 mm . . . . . . . Ophionea ishii ishii Habu
399(391') Prothorax subcylindrical or quadrate but with a small tooth in postero-lateral margin
opposite the last pair of setigerous punctures; tibia I normal or with laterally developed
claw-like teeth ......................................................... 400
399' Prothorax broad and usually wider than long ............................... 404
400(399) Tibia I with laterally extended claw-like teeth; tarsal segment IV simple; pronotum with
bordered lateral margins and a median longitudinal groove . . . . . . . . . . . . . . . . . . .401
400' Tibia I simple without laterally extended claw-like teeth; tarsal segment IV bilobed;
pronotum not bordered but heavily punctured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
401(400) Pronotum quadrate with a small tooth in the postero-lateral margin transversely op-
posite the last pair of setigerous punctures (Fig. 741); frons with 2 deep longitudinal
grooves and a moderately elevated lateral lobe between eyes and sides of clypeus;
clypeus truncate with a small rounded lobe on each lateral side; supraorbital groove with
2 setae; antennae bead-like and slightly beyond level of posterior setigerous puncture;
elytra with rectangular meshes, ridge 3 with 4 pores (Fig. 742); apex of tibia I with 3
claw-like teeth, middle tooth with a subtooth at base; reddish brown, length 5.5 mm . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clivina sp.
401' Pronotum slightly cylindrical, uniformly black, glabrous and shiny, and wider than
abdomen; frons with a small rounded horn in the middle; uniformly black species . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
402(401') Subapical transverse groove of pronotum concave medially and narrowed down to the
apicolateral angle, posteromedian margin almost unnoticeably indented; base of
pronotum without a short longitudinal groove on opposite sides of the median lon-
gitudinal groove (Fig. 743); elytra with moderately shallow and more widely separated
grooves; apices of ridges 2-6 punctated similar to ridge 1 and enclosed by ridges 1 and
7; ridge 5 reaching base of elytra (Fig. 744); labrum with a longitudinal median groove;
apico-median margins of clypeus straight; length 16 mm . . . . . . . . . . . . . Scarites sp. A
402' Subapical transverse pronotal groove gradually narrowed down to the apicolateral
angle with more convex-like pits; posteromedian margin with deep cleft and with a short
longitudinal groove on opposite sides of the long median longitudinal groove (Fig. 745);
elytral grooves deep and closer to one another, apices with few but wide pits; ridge 5
aborted before base and closed by ridge 6 and 7 (Fig. 746); labrum with a median
longitudinal groove; apicomedian portion of clypeus slightly concave; length 12 mm . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scarites sp. B
403(400') Head, prothorax and abdomen including elytra metallic green, closely punctated and
finely hirsute (Fig. 747); brown labrum trilobe-like with 2 setae in each lobe; apices of
femora I-III black; apical one-fifth of scape blackish and larger than the slightly bent and
brownish basal half; margins of frons in front of eyes parallel-sided; pronotum 2 times
longer than broad; apico-lateral corner of elytra with a small acute tooth (Fig. 748);
length 9-11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drypta gerliculata (Mug)
403' Head and prothorax brown, elytra black along sutural, basal, and lateral margins with a
brown band covering ridge 3-7; head less punctate behind the eyes; pronotum slightly
longer than broad and densely punctate (Fig. 749); elytral punctations on the intervals
and none on the ridges; brown labrum trilobed with lateral lobes rather acute lateral;
apices of femora I-III reddish brown; apical one-half to one-third of scape blackish
brown; margins of frons in front of eyes converging apically; apico-lateral corner of
elytra rounded (Fig. 750); length 7-9 mm . . . . . . . . . . . . . . . . . Drypta japonica (Bates)
404(399') Labrum strongly notched apicomedially; mentum emarginate with 2 setae near mid-
emargination; clypeus straight to moderately emarginate; margin of elytra without an
internal plica towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405
404' Labrum and clypeus truncated apically or labrum strongly rounded at apex; mentum
with tooth, straight or emarginate; margins of elytra with or without internal plica
towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406
405(404) Supraorbital setae one pair; clypeus straight to weakly emarginate; pronotum glabrous
and strongly bordered laterally; latero-apical one-third with a pair of setae, basal
one-third with a pair of long longitudinal grooves (Fig. 751); elytra uniformly black,
groove 5 and 6 fused before apex and not reaching base of elytra; groove 7 fused with
groove 8 becoming part of apical groove up to tip of groove 4; groove 1 with a setigerous
pore basally and converging towards groove 2 (Fig. 752); labrum with 4 setae; length
15-20 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplocheila polita (Fabricius)
405’ Supraorbital setae 2 pairs; clypeus moderately emarginate; pronotum very finely
punctate with light transverse striae and shallow longitudinal lateral grooves (Fig. 753);
elytral grooves 1 and 2 fused basally to a setigerous pore, grooves 3 and 4 and 5 and 6
separately fused at their apices before apical groove; groove 7 separated from 8 towards
apex (Fig. 754); labrum with 6 setae; length 14 mm . . . . . Submera latifrons (Dejean)
406(404') Midlateral sides of pronotum angular and bear a lateral seta on each side; pronotum
triangular posteriorly; head with 2 pairs of supraorbital setae . . . . . . . . . . . . . . . . 407
406' Lateral sides of pronotum more or less rounded; pronotum truncate posteriorly; head
with 1 pair of supraorbital setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408
407(406) Entire body black with lateral margins of pronotum and elytra yellow; head, prothorax,
and elytra very finely reticulated (Fig. 755); antennae blackish brown except brown
segments III and IV; hirsute except basal 4 segments; elytra subtruncate apically and
striae lightly and sparsely punctulate (Fig. 756); length 5.3 mm . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentagonica nr. erichsoni Schmidt-Goebel
407' Body black except red pronotum; antennae black except brown to dark brown basal 4
segments (Fig. 757); elytral striations faintly punctulate (Fig. 758); length 6 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentagonica ruficollis Schmidt-Goebel
408(406') Elytral margins without an internal fold toward apex; antennal segments III-XI hirsute
with a long seta toward apex, basal one-third of segment III and entire segment I and II
not hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
408' Elytral margins with an internal fold toward apex; antennal segments IV-XI with
pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 410
409(408) Anterior marginal line of pronotum deep and entire, posterolateral corner angulated
and posterior one-third with dense close fine pits (Fig. 759); head reddish brown
between eyes; labrum with 6 subapical setae and hairs apico-laterally (Fig. 759); groove
2 of elytra with a setigerous pore basally, entire groove complete, grooves 3 and 4 fused
towards apex similar to 5 and 6, 7th groove extended up to tip of 3 and 4; apices of elytra
brownish with slight emargination; scutellar striae absent; shiny black beetles with
margins of pronotum and elytra yellow (Fig. 760); length 8.6 mm . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoplogenius microgonus Bates
409' Anterior marginal line of pronotum fine and interrupted at middle, posterolateral angle
rounded and moderately punctated in between the angle and the median longitudinal
groove (Fig. 761); groove 2 very short reaching level of bulging of groove 1; elytra
brownish black concolorous to the head and pronotum with 4 yellowish spots, in grooves
5-8 near the humeri and base and 7-9 subapically (Fig. 762); scutellar setae present;
length 6.5 mm . . . . . . . . . . . . . . . . . . . . . . . . Egadroma quinquepushdata (Wiedemann)
410(408') Elytra with yellow marks or lateral margins of elytra yellow . . . . . . . . . . . . . . . . . . 411
410' Elytra uniformly black without yellow marks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417
411(410) Elytra with 3 yellow marks, in the humeri, 2 transversing groove 4 to midlateral margins
of elytra and 1 along sutural margin subapically covering suture up to groove 4 (Fig.
763); pronotum black rough medially with brownish red margins; head metallic blue-
green with brownish clypeus and yellow-brown rectangular labrum (Fig. 764); length
13 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius xanthospilus
411' Elytra with 2 subapical marks or entire lateral margin yellow . . . . . . . . . . . . . . . . . 412
412(411') Entire elytral margins yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413
412' Elytral margins black with yellow marks subapically . . . . . . . . . . . . . . . . . . . . . . . . 414
413(412) Yellow elytral margins broad covering groove 7 to margins of elytra and epipleura, the
band slightly narrows apically to the sutural margin (Fig. 765); pronotum (Fig. 766)
broadest medially and constricted just above lateral setae, disc slightly rough with widely
spaced moderately shallow pits; head (Fig. 766) moderately fine with network-like
marks behind eyes dorsally and 2-3 fine convex grooves between eyes radiating from
supraorbital area; labrum rectangular and yellow with 6 subapical setae (Fig. 766); head
and pronotum metallic blue-green, elytra black and legs yellow except black to reddish
brown coxae; length 15 mm (Fig. 767) . . . . . . . . . . . . . . Chlaenius circumdatus Brulle
413' Yellow elytral margins narrow not reaching groove 7, the band broadens towards apex;
lateral margins of pronotum gradually rounded with thin and narrow yellow band (Fig.
768); length 11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius inops Chaudoir
414(412') Apical one-third of elytra with a broad V or U-shaped yellow band, top of band with a
foot extended to groove 3 (Fig. 769); elytra almost entirely clothed with short brown
hairs; pronotum with reddish brown lateral margins, disc clearly punctate with the
posterior area more closely punctate than the anterior; head very finely punctate;
labrum slightly emarginate at midapex and with a black spot posteromedially (Fig. 770);
length 9-14 mm (Fig. 771) . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius virgulifer Chaudoir
414' Subapical band of elytra not U-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
415(414') Elytron with a moderately large subapical yellow spot covering grooves 3-8 (Fig. 773),
and entire elytra punctate; yellow spot short between grooves 3-5 and longest between
5-6; pronotum with a deep lateral longitudinal groove on each side of median groove
posteriorly; basal third of pronotum more punctate than the anterior; head smooth
between eyes except for some very fine transverse striae; anterior and posterior area
with fine punctations; median area of clypeus with moderately dense fine punctations
(Fig. 772); antennae blackish brown except yellow-brown scape; elytra black, head and
pronotum metallic blue-green; legs with black tibiae and knees, reddish brown tarsi and
yellowish brown to red femora; trochanters dark reddish brown; length 12-13 mm . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius bioculatus Motschulsky
415' Subapical mark on elytron small and represented by short longitudinal bands or 2
contiguous bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 416
416(415') Yellow marks of elytron on intervals 4, and 6 to 8 with the 6th interval having the longest
band (Fig. 774); pronotum with moderately deep, evenly distributed and uniform
punctations (Fig. 775); head smooth behind, finely punctated and relatively smooth up
to frons except oblique striations near supraorbital area; clypeus smooth and with very
fine pits; labrum reddish brown, narrows behind, and widens opposite latero-subapical
setae; antennae IV-VIII black, IX-XI brown, III brown except blackish ends, scape and
pedicel brownish yellow; elytra dull black, prothorax black with metallic blue-green
luster laterally and head lustrous blue-green; length 14.5 mm . . . . . . . Chlaenius sp. A
416' Yellow marks of elytron on intervals 4, and 7 to 8, with the 6th interval without band or
reduced to 3 small dot-like forms (Fig. 776) or contiguous on interval 4 to 6 and a spot
on interval 8 (Fig. 777); pronotum with more prominent carina between pits; head with
light punctations, short median longitudinal striae between eyes and oblique striae on
top of it present; clypeus with small evenly scattered pits; labrum black and reddish
apical margin (Fig. 778); antennal scape and pedicel yellow-brown, segment III dark
brown with black ends, IV to VI blackish and VII to XI brownish yellow (Fig. 778);
femora and dorsum of tibiae yellow, knees, venter of tibiae and entire tarsi black; length
13mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius sp. B
417(410') Pronotum noticeably punctated, punctations dense apically and posteriorly and sparse
or well separated medially, lateral groove wide and less elongated (Fig. 779); head
smooth with sparse fine punctations, labrum moderately emarginate with setae almost
midway its length; elytra1 striae 3 and 4, and 5 and 6, fused, separated at apices (Fig.
780); length 12.6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius sp. C
417' Pronotum with very few scattered fine punctations; striae 6 separated from 5 which is
fused to 7 apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 418
418(417') Pronotum lustrous metallic green with 2 rows of sparse punctations aligned to the
median longitudinal groove, discal area with light, widely scattered shallow punctations
and about one-fourth wider than long (3:4); head concolorous with pronotum, smooth
and finely punctated behind head, and with 2-3 light grooves along supraorbital area;
labrum red (Fig. 781); elytra with striae 5 zigzagging towards apex, separated from 6 and
7 (Fig. 782); length 15 mm . . . . . . . . . . . . . . . . . . . . . . Chlaenius quadricolor Fabricius
418' Pronotum metallic blue to shining black, almost as long as wide (19.5 : 21); pronotal disc
moderately punctated and with light transverse striations along the median groove; head
concolorous with pronotum, finely punctated up to clypeus and slightly rough behind
eyes (Fig. 783); labrum red with 3 small black spots basally; elytra1 striae 3 and 4 fused
apically with tip joining striae 7, striae 5 and 6 separated and vaguely connected to striae
7 (Fig. 784) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlaenius sp. D
419(365') Small, 2.75-4.3 mm long; elytra testaceous, marking consisting of zigzag double, clear
lines, sometimes the lines becoming thick and coalescent . . . . . . . . . . . . . . . . . . . . 420
419' Moderately large to large; 10-37 mm long; elytra mostly black with yellow longitudinal
markings; if pale yellow-brown, pronotum and elytra with transverse black band . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421
420(419) Elytral markings (Fig. 785) fine undulating, zigzag double lines, clear and thick but
never coalescent; pronotum (Fig. 786) testaceous with transverse black markings in the
middle along the anterior and posterior margins, and a narrow median transverse black
patch on the disc; male penis with a broad, twisted plate, broader in front of the arch
and deeply emarginate at apex ..................... Laccophilus sharpi Regimbart
420' Elytral markings as above but generally thick and coalescent; pronotum (Fig. 787)
testaceous without any black markings anteriorly and posteriorly; penis moderately
strongly curved medially, almost uniformly wide and notched at the apex on the ventral
side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laccophilus parvulus Aube
421(419') Prolateral sides of tarsi III armed with a double layer of comb-like teeth, retrolateral
sides clothed with golden brown fringe of hairs; body black with yellow bands on sides
of pronotum, elytra, or apical part of frons; body length 24-40 mm . . . . . . . . . . . . 422
421' Prolateral and retrolateral sides of tarsi III lined with golden yellow-brown fringes of
hairs; body black or yellow with black punctations; body length 10-14 mm . . . . . . 424
422(421) Anteapical inward dilatation of the yellow band of the elytra drop-like (Fig. 788); its
inner rounded part separated by half its length from the main yellow band; two lon-
gitudinal rows of shallow and faint punctations in the black part of elytra evenly
separated and rather sparse, median punctures within 2 rows made up of four punctures
square form; body length 35-40 mm . . . . . . . . . . . . . . . . . . . . . Cybister japonicus Sharp
422' Anteapical inward dilatation hooked inward (half arrow-like); puncture arrangement
not in line with each other transversely, but in oblique pattern four points form a
rectangle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423
423(422') Elytra and pronotum very smooth; pronotal lateral yellow band one-third narrower than
the basal yellow band of elytra; lateral band of elytra (Fig. 789) trifurcated slightly below
midhalf, space in between trifurcation reddish brown, anteapical inward dilatation half
arrow-like and connected to the tip of the lateral band before the suture; row of
punctures adjacent to suture with 17 pits, row closer to the yellow band with 12;
anteromedian part of pronotum with a very shallow (almost unnoticeable) cleft,
posteromedian at base of scutellum slightly procurved; body length 26 mm . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cybister trifunctatus orientalis Gschwendtner
423' Elytra with broken fine longitudinal striae except smooth unstriated area near suture
and proximal one-third; punctations near suture with 23 pits and 14 in other row
adjacent to the yellow band excluding two more pits in the anteapical inward, broad
hook-like band; elytral yellow band narrowed proximally and broad towards base of
elytra, two times broader than that of lateral pronotal band; punctures form an oblique
rectangle transversely (Fig. 790); pronotum with irregular network-like irrorations;
anteromedian of pronotum with a small noticeable cleft, posteromedian with a very fine
indentation (almost unnoticeable); body length 38 mm . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cybister trifunctatus subsp. A
424(421') Lateral border (Fig. 791) of elytra serrated below midhalf but not reaching apex of
suture; tip of suture tapers to a point; elytra with deep black punctations, a black spot
lateromedially laterad of a small longitudinal groove; and a transverse zigzag-like band
on the top of the posterior one-third; base of elytra reddish brown towards scutellum;
pronotum (Fig. 792) rebordered laterally with a transverse black median band broken
at middle by eight minute globular to ovate blackish brown spots; head yellow except for
black back and a transverse oblong black spot dorsad of black hindpart of head; apex of
frons strongly recurved medially . . . . . . . . . . . . . . . . . . . . . . . . Eretes sticticus Linnaeus
424' Lateral border of pronotum not serrated; lateral area of pronotum not rebordered;
posterior claws unequal, apical spurs of hindtibiae pointed; suture between the
metapisternum and metasternal wing straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 425
425(424') Elytra smooth and without punctate-like marks, black with yellow humeral and sub-
humeral stripes narrow and joined together posteriorly after the middle (Fig. 793);
pronotum with a broad yellow band laterally, as broad as the distance from elytral
margin to subhumeral stripe; frontal half of head yellow; antennae yellow except VI-XI
blackish brown; mesotarsi of male with two rows of sessile palletes; tarsal claws of legs
I and II always widely diverging; tibiae I enlarged submedially to as much as half its
length, with three long setae apicolaterally, five (3 moderately large and 2 hair-like
setae) at venter of enlarged tibial area and spineless in between the two groups of setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydaticus vittatus (Fabricius)
425' Elytra brown with black interconnected punctate-like marks, contiguous medially (Fig.
794); lower humeral area and lateral margins yellowish; scutellum black; pronotum (Fig.
795) brownish red with blackish shades midanteriorly and midposteriorly; head con-
colorous with pronotum; antennae uniformly brownish yellow; mesotarsi as in H. vit-
tatus; tarsal claws usually parallel to each other; tibiae I with its width of enlarged area
a little over half of tibial length, apicolateral with three setae, venter of enlarged area
with 4 (3 moderately large and 1 hair-like setae) and with small spines in between the
two groups of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydaticus rhantoides Sharp
426(364') Head prolonged into a distinct beak or snout; basal segment of geniculate antennae
often received in grooves on snout; labrum absent; tarsi 4-4-4; front tibiae usually
without series of teeth externally or prolonged distally into a stout spur; size and shape
variable [Curculionidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
426' Not with the above combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
427(426) Tarsi simple; body covered with waterproof coat of scales; rostum long; larvae are root
feeders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 428
427' Tarsi bilobed; body without prominent waterproof coat of scales; rostrum short or long;
1arvae may or may not be root feeders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
428(427) Pronotum (Fig. 796%) longer than wide, almost rectangular; body covered by varnish-like
coat of scales, bead-like scales along eye margins, back of head, apical one-third of
rostrum, coxae I to III, venter of prothorax, and between coxa II and last 4 abdominal
sternites except the penultimate segment; eyes partly hidden dorsally, twice longer than
broad; space between eyes convex; antennal groove straight; space not reaching eye
(Fig. 797); tibia II (Fig. 798) medially curved with a prominent upcurved spine and a
small one opposite it; ash-grey; adults 3-3.5 mm, temperate species ................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lissorhoptrus oryzophilus Kuschel
428' Pronotum wider than long, rounded along lateral sides; body covered by small leaf-like
scales on elytra and somewhat circular scales in pronotum; eyes prominently visible
dorsally, one-fourth higher than wide; space between eyes with a pit (Fig. 799); elytra1
grooves fine and straight; tibiae straight, II curved towards tip with a single spur (Fig.
800); black adults 5 mm or more, Oriental species . . . . Hydronomidius molitor Faust
429(427') Rostrum short, as long as wide or a little longer than wide; larvae not root feeders . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430
429' Rostrum much longer than wide and more or less cylindrical; larvae are root feeders .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
430(429) Rostrum short, a little longer than wide, dorsomedian longitudinal ridge present and
highest above the area between antennal base and anterior eye margin; mandible with
deciduous cusp leaving a scar; antennal groove curved below front of eye margins; scape
long almost reaching collar of pronotum; eyes visible above; globular (as wide as high);
pronotum cylindrical longer than wide (Fig. 801); antennae dark reddish brown, seg-
ment II as long as III; black . . . . . . . . . . . . . . . . . . . . . . . Tanymecus nr. boettcheri Voss
430' Allied to T. nr. boettcheri; prothorax less cylindrical; dorsomedian longitudinal ridge
highest above antennal base or a little above it; scape of antennae not reaching hind
margin of eyes (Fig. 802); antennae entirely black, segment I1 longest next to pedicel,
and 2.2 times longer than segment III (Fig. 803) . . . . . . . . . . Tanymecus indicus Faust
430'' Rostrum nearly as wide as long, deeply grooved medially from apical one-fourth to basal
one-eighth, side grooved as in Tanymecus; head and thorax covered with circular scales,
glossy and hairy; eyes subspherical, slightly bulging, rimmed with white scales; antennae
12-segmented with 4 compact into club, 1st funicular segment one-third shorter than
combined length of segments II and III; prothorax about one-fourth wider than long,
wider behind than in front; elytra tapers apically with 6 dotted lines up to shoulder (Fig.
804); tips of tibiae without spur except brush of hairs (Fig. 805) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypomeces nr. squamosus Fabricius
431(429') Elytra with a pair of whitish oblong spots on the apical one-third of stria 2 and 3 (Fig.
806) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinocnemus squameus Billberg
431' Elytra without any white spots (Fig. 807); rostrum shorter than E. squameus . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinocnemus oryzae Marshall
432(426') Antennae lamellate (at least three); with ten or fewer segments; front coxae prominent;
pygidium exposed in most species [Scarabaeidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . 433
432' Antennae not lamellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 440
433(432) Tarsal claws of leg III unequal in length, inner claw smaller; tibiae III with 2 long spurs
at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
433' Tarsal claws of leg III equal in size, toothed or bifid; foretibiae with 2-3 outer tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438
434(433) Black beetle with shiny pronotum and elytra; pronotum very smooth; lateral and
anterior parts strongly margined; head roughly carinate, lightly punctated in between
eyes; anterior clypeus with a pair of upcurved “horns” (Fig. 808); 6 large mandibular
teeth visible dorsally; tibiae I strongly serrated with 7 teeth in the outer part 1,2 and 4
the longest, inner apical part with one long tooth; outer claw of tarsi very broad and
scoop-like in males; elytra (Fig. 809) each with 11 fine striae, striae 4, 5, and 6 enclosed
by 3 and 7 striae; scutellum not punctated, triangular apically; length 17 mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteronychus lioderes Redtenbacher
434' Brownish red or blackish brown beetles; body may or may not be hirsute; clypeus
without horns at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
435(434') Uniformly hirsute; head, thorax, and elytra clothed with spike-like decumbent white
hair, hair underneath rather fine; moderately flat beetles; clypeal (Fig. 810) margins
strongly round, thin edges upcurved; pronotum partly rough, almost rectangular, 2.3
times wider than long; scutellum not triangular; semi-punctated, lateral edges curved
and basal end curved medially; basal end of elytra deep between shoulder and a hump
before scutellum; outer edge of tibiae I with 3 blunt tubercles, 1 and 2 nearer each other,
2 and 3 wide apart, and a moderately long spur in inner side (Fig. 811); legs dark
brownish red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adoretus ranunculus Burmeister
435' Not hairy as above; elytra brown; pronotum reddish brown . . . . . . . . . . . . . . . . . . 436
436(435') Head black except dark brown clypeus; vertex slightly raised; clypeus punctated,
rounded with elevated edges, basal area in front of eyes straight or shortly angled;
transverse ridge above clypeus prominent, elevated near eyes; pronotum with black
patches anteriorly, smooth, 2.2 times wider than long, margined with a broader
anteromedian than rest, laterally rounded, slightly straight to concave anteriorly and
concave posteriorly (Fig. 812); elytra brown, punctated with 2-3 longitudinally low
ridges between 3-5 irregular rows of black striae; scutellum rounded apically with base
nearly straight; outer part of tibia I (Fig. 813) with 3 black-tip blunt tubercles, inner tip
with a short spur; outer claw of leg I bifurcated; legs yellow with blackish brown patches
on femora, and tibial tubercles . . . . . . . . . . . . . . . . . . . . . . Adoretus luridus Blanchard
436' Reddish brown beetles with brownish yellow to reddish brown elytra; body finely
punctated; clypeus broadly angled posteriorly; venter of body clothed with long fine
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
437(436') Elytra reddish brown, same color as head and thorax; glossy bodied beetle, head
subtruncate anteriorly; pronotum much broader behind the anterior, midlateral half
slightly bulging (Fig. 814); tibiae I with one long blunt tubercle on outer end and a small
inner one (Fig. 815); tibiae III with at least 3 rows of spurs in the outer median half . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anomala sulcatula Burmeister
437' Elytra brown or brownish yellow, dark reddish brown on shoulder extended below to
lateral margins and on the apical hump (Fig. 816); tibiae I with 2 tubercles (Fig. 817);
tibiae III with 2 rows of spurs in the anterior and posterior end of the outer median half
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anomala humeralis Burmeister
438(433') Anterior and posterior margins of pronotum margined (Fig. 818); pronotum less con-
vex, apico-lateral half widely margined and higher than “eye bar”, median half deeply
lobed and bulged posteriorly viewed from above (Fig. 819); clypeus with upcurved
margins, cleft median area (Fig. 820); tibiae I with 3 tubercles on the outer side and a
spur on the inner end opposite 2nd and 3 rd tubercles; stridulatory file present in the
anterior edge of the elytra1 epipleuron; ridge of shoulder incurved posteriorly (Fig. 821)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holotrichia spp.
438' Pronotal margins not margined except lateral edges . . . . . . . . . . . . . . . . . . . . . . . . 439
439(438') Body black and slightly shiny, surfaces covered with spherical to ovoid whitish yellow
spots (Fig. 822); clypeus partly cleft anteriorly and broadly angled posteriorly (Fig. 823);
lateral end of pronotum lobed behind midhalf and posterior tip pointed (Fig. 824); body
length barely 30 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucopholis irrorata (Chevrolat)
439' Body brownish red, not glossy, uniformly clothed with spike-like setae (Fig. 825);
clypeus broadly margined and raised medio-apically (seen frontally) (Fig. 826), not
cleft, lateral margin of pronotum lobed a little behind median, posterior tip rounded
(Fig. 827); body length more than 30 mm . . . . . . . . . . . . . . . . . . . . . . . . Leucopholis sp.
440(432') Aquatic insects; maxillary palpi antenna-like, long, as long as or longer than the short
antennae; vertex with a Y-shaped impressed line, antennal cupule preceded by five
segments; front coxal cavities open behind; procoxae not concealing prosternum; tarsal
segmentation 5-5-5; with or without median sternal keel [Hydrophilidae] . . . . . . 441
440' Terrestrial insects; maxillary palpi short; vertex without Y-shaped impression; antennae
8-11 segmented without cupule; pronotum variable either hexagonal, constricted on
both ends or widest to globular at middle; head subglobular to strongly transverse; tarsal
segmentation 4-4-4,5-5-4 or 5-5-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445
441(440) Large aquatic beetles, 28-35 mm long; head including eyes, pronotum, scutellum, and
elytra entirely black; elytra with two rows of longitudinal pits narrowing on both ends;
median sternal keel very long reaching grooved prosternum . . . . . . . . . . . . . . . . . 442
441' Small aquatic beetles, 5-12 mm long; elytra never entirely black; median sternal keel
absent, if present not beyond coxae I and not contained in the prosternum; groove in
prosternum absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 443
442(441) Bases of two longitudinal rows of punctures widely separated, distance between them as
long as distance from side of scutellum to subsutural punctations (Fig. 828); median
sternal keel reaching midhalf of sternite III; venter of body including head and thorax
black with few red-brownish tinges on apices of femora and sides of abdomen; junction
of Y-shaped impression about in level midhalf of eye, leg of Y shorter than eye length
taken laterally (14:18) (Fig. 829); length 32-35 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrophilus acuminatus Motschulsky
442' Bases of two longitudinal rows of punctures much nearer each other than the distance
of subsutural punctations to side of scutellum (Fig. 830); median sternal keel reaching
almost apex of sternite III; venter of body brownish red; junction of Y-shaped impres-
sion above midhalf of eyes, leg of Y longer than eye length taken laterally (20:17) (Fig.
831); length 28 mm . . . . . . . . Hydrophilus nr. bilianeatus cashmirensis Redtenbacher
443(441') Median sternal keel completely absent; head, pronotum, scutellum and elytra
punctated; junction of Y-impression in line with apical third of eye, leg of Y long and as
long as eye length taken laterally; back of head behind eyes with a transverse groove;
elytra with 10 long punctated grooves and 6th and 7th not fully reaching basal margin
across humeri and a small incomplete one after the first near sutural line; in between
long grooves 1-2 lines (usually one) of punctures; head longer than pronotum; generally
yellow-brown with black abdomen and metapleuron; length 5.5 mm (Fig. 832) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Berosus sp.
443' Median sternal keel present but short, not reaching apical half of coxae I; prosternum
entire and without a groove; pronotum with two rows of punctations along lateral sides,
discontinued medially; apical half of clypeus reddish brown; frons with punctations near
eye margins; lustrous black beetles; length 9-12 mm . . . . . . . . . . . . . . . . . . . . . . . . . 444
444(443') First antennal segment short, only reaching anterior eye margins (Fig. 833); prosternum
gradually produced or narrowed forward with a straight floor (seen laterally); median
sternal keel slightly notched subapically and without bristles; apex of clypeus moderate-
ly notched medially; frons with two transverse concave punctations and almost medially
erased; scutellum 1.22X longer than wide; body form less convex; length 12 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sternolophus rufipes Fabricius
444' First antennal segment beyond midhalf of eye (Fig. 834); prosternum rounded forward
with its floor slightly oblique at apex and straight posteriorly; subapex of median sternal
keel with bristles in the notch; apex of clypeus straight; frons with transverse punctations
medially, concave on booth ends with closer punctures and sparse medially but
prominent; scutellum nearly as wide, as long; body strongly convex; length 9 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sternolophus sp.
445(440') Tarsal formula 5-5-4, heteromerous with bipartite claws; a sclerotized blade-like
process beneath each claw; body cylindrical and moderately soft; eyes nearly transverse
with emarginate margins near antennae [Meloidae] . . . . . . . . . . . . . . . . . . . . . . . . . 446
445' Tarsal formula variable 5-5-5, 5-5-4, 4-4-4 but without such claws . . . . . . . . . . . . . 447
446(445) Body uniformly black, clothed with thick white hair; eyes oblique with base of antennae
below top level (Fig. 835); antennae same color as body except brownish scape, pedicel,
and dorsal half of segment III all covered with white hair dorsally and black hair laterally
and ventrally; scape nearly 3 times as long as pedicel, segment III much longer than
segment IV, one-third shorter than combined length of IV and V (Fig. 836) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cylindrothorax sp.
446' Body black except orange-brown pronotum; eyes transverse with antennal base almost
in line with anterior level of eyes (Fig. 837); edges of pronotum markedly grooved
elevating outer rims, lateral and posterior edges dark brown, anterior one-third with a
transverse deep region (Fig. 838); elytra with bluish lustre; antennae entirely black,
scape twice as long as pedicel, segment III as long as IV . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cylindrothorax melanocephala Fabricius
447(445') Tarsal segmentation 3-3-3 (each tarsus with 3 segments, actually 4-segmented each with
the basal two segments expanded and densely pilose beneath, 2nd produced into a long
lobe underneath, 3rd minute and concealed, 4th long); head deplexed strongly or deeply
sunken into the prothorax; body broadly ovate and highly convex in form; entire ab-
domen covered by elytra; 5-6 visible sternites, basal sternites with a curved femoral line
running backwards from inner end of coxae; antennae 8-11 segmented with a 3-seg-
mented club [Coccinellidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 448
447' Tarsal segmentation 5-5-5 or 5-5-4; body form usually elongate, not strongly convex;
head not deeply sunken into prothorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 460
448(447) Clypeus expanded laterally; antennae 8-segmented; postcoxal line complete, tarsal
claws simple, slightly thickened at base; body oval to rounded oval and moderately
convex above; elytra with three longitudinal black vitta and a yellow longitudinal vitta in
between . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 449
448' Clypeus not expanded laterally; anterolateral angle of clypeus produced forward; anten-
nae 11-segmented; postcoxal line incomplete; tarsal claws usually cleft; genae extending
into eyes; mandibles bifid at apex; body broadly oval to elongate oval with or without
spot or broad longitudinal bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 450
449(448) Discal and sutural black vittae both broad, reach base of elytra and confluent basally,
discal band distinctly more than 2 times wider than the width of yellow and sutural bands
taken separately; head yellowish or red testaceous with blackish area on vertex in the
male and entirely blackish in the female; pronotum yellowish brown; punctations in the
head and pronotum fine and sparse and muchstronger and closer in the elytra (Fig. 839)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brumoides lineatus (Weise)
449' Sutural and discal black vittae both narrow; discal band rarely more than twice wider
than the yellow and sutural bands and never confluent with the sutural band; only sutural
band reaches base of elytra; coloration similar to B. lineatus (Fig. 840) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brumoides suturalis (Fabricius)
450(448') Tibial spurs absent in all legs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 451
450' Tibiae II and III with spurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456
451(450) Elytral epipleura wide, concave and strongly descending externally . . . . . . . . . . . . 452
451' Elytral epipleura moderate, feebly concave and almost horizontal; body broadly elon-
gate oval and less convex; prosternum without carinae; abdominal lines incomplete;
pronotum reddish yellow with one or two pairs of quadrate to subovate black spots;
elytron normally with a pair each of transverse black humeral, median, and a spot each
of post-median and subapical; sutural band narrow and broadens postmedially (Fig.
841); paramere short and moderately stout, about as long as basal piece; infundibulum
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harmonia octomaculata (Fabricius)
452(451) Elytra with longitudinal streaks; pronotum with black marks . . . . . . . . . . . . . . . . . . 453
452' Elytra uniformly yellow to reddish yellow without black marks . . . . . . . . . . . . . . . . 455
453(452) Sutural band black; thin and narrow on both ends but gradually widened medially; discal
band absent; posterior one-third of pronotum black medially and a pair of spherical
spots above it (sometimes entire basal half black without spots above but with 2-3
dentations anteriorly); elytra orange-brown (Fig. 842); siphonal capsule moderately
stout, its outer arm moderately stout and inner arm short and truncated at tip; sipho
relatively curved and shorter than M. hirashima and M. yasumatsui; paramere noticeab-
ly subglobular in apical half; apex of medium lobe broadly truncate . . . . . . . . . . . . . . . .
................................................. Micraspis discolor (Fabricius)
453' Sutural and discal bands prominent ...................................... 454
454(453') Discal band very broad and curved, 2-3 times broader than sutural band, and not
reaching distal and proximal ends of elytra; posterior half of pronotum black and apical
half yellow; general color yellowish brown (Fig. 843) . . . . . Micraspis inops (Mulsant)
454 ' Discal band narrower than median area of sutural band; both bands narrower than the
reddish yellow background of elytra, reach proximal end but not distal end of elytra;
pronotum blackish except yellow lateral margins; general color reddish yellow (Fig. 844)
................................................... Micraspis vincta (Gorham)
455(452') Body short oval, about four-fifth times as wide as long; head slightly narrower than half
the pronotal width and interocular distance slightly shorter than 0.5 head width; apical
side of terminal antennal segment strongly obliquely truncate (Fig. 845); prosternal
carinae more broadly separated; sipho relatively stout and weakly curved with a slender
and longer sclerite in the apical end; siphonal capsule very long and slender inner
process, the latter rarely shorter than the outer process (Fig. 846); tegmen comparative-
ly slender (Fig. 847); apex of median piece strongly bending dorsally and sharply
pointed at apex in side view, very narrow, parallel-sided with a truncated tip; reddish
with yellowish brown head, antennae, pronotum, scutellum, legs and abdomen except
for a black median area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micraspis hirashimai Sasaji
455 ' Body nearly hemispherical, slightly longer than wide; head distinctly broader than half
the pronotal width; interocular distance nearly half as broad as the head width; apical
side of 11th antennal segment weakly obliquely truncate (Fig. 848); sipho (Fig. 849)
relatively slender and weakly curved; siphonal capsule slender with outer process much
longer than the inner one; tegmen (Fig. 850) slender and long, about two-fifths as long
as sipho; median piece slender, gradually narrowed apically with a truncate tip; lateral
lobes of tegmen also long and slender, more than three-fifths of tegmen weakly arcuate
laterally; orange-brown, head yellowish brown to orange-brown with darker antennae
and mouthparts, underside of thorax and abdomen entirely brown and legs yellow-
brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micraspis yasumatsui Sasaji
456(450') Body slightly depressed and ovate (Fig. 851); prothoracic hypomeron with distinct
fovea; pronotum yellow with a pair of subquadrate spots separated by a thin yellow line
posteromedially; elytra yellow with external margination complete and uninterrupted,
and with two interconnected sometimes interrupted anchor-like black bands (Figs.
852-853); humeral black markings never divided into two separate markings; proster-
num with distinct narrow lateral ridge reaching the apex; postcoxal line incomplete; legs
yellow to yellowish brown without black markings . . . Propylaea japonica (Thunberg)
456 ' Body moderate to strongly convex, rounded or broadly elongate oval; patterns on elytra
entirely different ...................................................... 457
457(456') Femur III long extending well beyond the outer margin of elytra; tarsal claw with a
slender tooth at middle; tibiae II and III with two spurs; femoral line on first abdominal
segment complete; body elongate oval; elytron with 6 spots 4 above median (1 each in
discal and suturo-basal, humeral and above humeral) and 2 below median (1 large
knob-like submedian and a smaller subovate subapical); pronotum largely black except
yellow lateral and anterior margins, apical part of black band slightly 4-lobed with 3
clefts and a pair of small ovate yellow spots (Fig. 854); sipho with four hooked-lobes on
sides of ventral and dorsal part; median piece of tegmen slender with a point apex and
longer than paramere; length 4-4.6 mm . . . . . . . . . . . . Hippodomia variegata (Gocze)
457' Femur III short, not reaching the outer margin of elytra; body rather rounded and
convex; frons narrow, about 0.5 as long as head; pronotum covering large part of eyes
and widest near base; prosternal process with parallel carinae . . . . . . . . . . . . . . . . 458
458(457') Black basal band of pronotum transversely long and narrow and connected by a short
very narrow longitudinal constriction to a transverse oval black discal spot; elytra with a
black sutural stripe dilated behind scutellum and before apex; each elytron with two
wavy post-humeral and median black bands, and a rounded black subapical spot; all the
bands and spot not connected to the sutural stripe (Fig. 855) . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Menochilus sexmaculatus (Fabricius)
458' Pronotum almost entirely black except for a yellow patch along its antero-lateral half;
elytra together with 7 spots, each elytron actually with 4-5 spots . . . . . . . . . . . . . . 459
459(458') Elytra with a black trilobed or slightly Y-shaped subhumeral marking, postmedian black
band wavy and confluent with sutural margin, sutquadrate black spot at three-fourths
the external margin, and one black spot each on both ends of the sutural margin (Fig.
856); each elytron with 5 spots; prosternum with a pair of carinae which hardly extend
beyond the level of the front coxa; abdominal lines forked V-shaped-like in the external
part; tibia III with distinct apical spur . . . . . . . . . . . . . . . Coccinella repanda Thunberg
459 ' Elytra1 spots globular to subglobular and well separated from each other except almost
contiguous spots behind scutellum; each elytron with 4 spots; sutural band absent except
at base (Fig. 857) . . . . . . . . . . . . . . . . . . . . . . . . . . .Coccinella septempunctata Linnaeus
460(447') Elytra very short exposing a considerable portion of abdomen, exposed abdominal
tergites laterally margined; tarsal segmentation 5-5-5; generally orange-red with blue-
black elytra, blackish head and apical two segments of abdomen . . . . . . . . . . . . . . 461
460' Abdominal tergites entirely covered by the long, well developed elytra . . . . . . . . 462
461(460) Pronotum subglobular (Fig. 858); all legs blackish brown; femur uniformly blackish
brown (Fig. 859); terminal segment of palpi blackish brown; black basal area of elytra
not very close to the pronotum (Fig. 860) . . . . . . . . . . . . . . Paederus tamulus Erichson
461' Pronotum elongate-ovate (Fig. 861); all legs brownish yellow except apex of femur III
(Fig. 862); terminal segment of palpi brown; brown basal area of elytra close to
pronotum (Fig. 863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paederus fuscipes Curtis
462(460') Tarsal segmentation 5-5-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463

462' Tarsal segmentation 5-5-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
463(462) With click mechanism formed by a long prosternal process; labrum visible; head black;
small and transversely ridged between eyes; pronotum with an inverted W-shaped
brown band, posterolateral angle with a sharp tooth (Fig. 864); scutellum slightly oblong
and deeply sunken between elytra; elytra blackish brown basally, yellow-brown towards
margin with black patches in the discal area; median base of elytron protruded basally
(Fig. 865); antennae extended to the tooth of pronotum; body clothed with yellow hairs;
length 7 mm [Elateridae] . . . . . . . . . . . . . . . . . . . . Aeoloderma brachmana (Candeze)
463' Without click mechanism, posterolateral angle of pronotum without a sharp tooth;
11-segmented antennae shorter than body; tarsus pseudotetramerous with 4th segment
small and hidden in the bilobed 3rd segment; leaf beetles (Chrysomelidae] . . . . . . 464
464(463') Head with frons inflexed, mouthparts hypognathous; pronotum and elytra without
broadly expanded margins Hispinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 465
464' Head with frons not inflexed, mouthparts prognathous . . . . . . . . . . . . . . . . . . . . . . 471
465(464) Body cylindrical, narrow, elongated, and parallel-sided, unarmed with distinct spines;
head irregularly punctured; basal joint of antenna long, thick, and apically compressed
and dilated externally; prothorax subquadrate longer than broad, sides straight and
parallel, posterior angles acute; scutellum black, impunctate and apically rounded;
elytra parallel-sided, more than 4 times longer than broad with ten regular rows of striae
and finely serrated towards tip; small beetles 4.5-5 mm long, dorsally metallic green with
lustre, black ventrally (Fig. 866) . . . . . . . . . . . . . . . . . . . . . . . . . . Leptispa pygmaea Baly
465' Body margins with spines dorsally and laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . 466
466(465') Anterodorsal margin of pronotum spineless; lateral edges each with 4 short spines, 2
anterolaterals nearly contiguous basally and 2 more below middle widely separated (Fig.
868); pronotum (Fig. 868) constricted behind 4th lateral spine, cylindrical to somewhat
rounded dorsally, not quadrate as in Dicladispa and Dactylispa; elytra with 4 small, short
spines in laterobasal half (Fig. 867); segment III of antennae nearly as long as basal and
4th segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichispa sericea (Guerin)
466' Anterodorsal margin of pronotum without spines; prothorax with a group of long and
prominent spines behind anterolateral angles and a single spine behind middle; elytral
margin not produced; antennae 11-segmented; segment III longest; body narrow and
subparallel (Dicladispa) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 467
466'' Not as above, anterior margin with a spine or group of spines on each side . . . . . 469
467(466') Oriental species; basal segment of 11-segmented antennae with a short ventro-apical
tooth; pronotum with 4-branched stout spines laterally before middle and a simple one
below middle; posterolateral angle without a prominent thin seta; spines more or less
straight; elytral punctations in regular rows, extreme apex without a long spine; lustrous
blue-black hispid except dark brownish to black legs and antennae, and reddish femora;
body length 3.7-4.8 mm (Fig. 869) . . . . . . . . . . . . . . . . . . . Dicladispa armigera (Oliver)
467' African species; each side of pronotum with 4-6 robust spines . . . . . . . . . . . . . . . . . 468
468 Pronotum with 4 spines, one 3-branched before middle and a simple one below middle,
lightly punctated, posterolateral angle with a short thin setae; elytra with relatively deep
irregular punctations, spine on apico-lateral area large with a broad base, extreme apex
with a prominent spine at median (Fig. 870) . . . . . . . Dicladispa viridicyanea (Kraatz)
468' Pronotum with 5 long spines, one 4-branched before middle and a single spine just
behind the tetramerous spine; a small, short, horizontal spine present posterolaterally
(Fig. 871); antennae without spines; elytra coarsely punctated, body oblong, bluish-
black including legs and antennae, shiny except opaque pronotal disc
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicladispa striacollis Gestro
469(466') Antenna lacking spines, segment III a little shorter than length of segment IV and V
together; lateral margins of pronotum with 3 spines, median apical two very close to
each other basally, 3rd spine below middle; anterodorsal part of pronotum with 2
double-branched spines; elytra coarse and clothed with numerous spines, lateral ones
longer and more slender than the short but robust inner ones (Fig. 872); African species
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylispa bayoni Gestro
469' First antennal segment with a long simple or 4-branched spine, segments 11-VI with or
withoutspines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 470
470(469') Basal segment of antennae with a 4-branched spine, apicodorsally, 2nd with two, 3rd to
6th with one each (Fig. 873); pronotum constricted behinde simple spine, with 10 spines,
4 double-branched with 2 each of anterodorsal and lateral margin, and 2 simple ones
below the laterals; head rugose with grey hairs around eyes; black with glossy punctate-
striate elytra (Fig. 874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hispa stygia (Chapuis)
470' Basal segment of antennae the largest, with a simple spine (Fig. 875), segment II small
and rounded, increased in size thereafter with 5 apical segments thick and elongate
club-like covered by brown pubescence; prothoracic spine close to H. stygia except for
a subacute tooth in the anterolateral edge; head broad and rugose, convex eyes rimmed
with silvery hairs; bluish black hispid . . . . . . . . . . . . . . . . . . Hispellinus moestus (Baly)
471(464') Pronotum broader than long or as broad as long, sides strongly constricted behind
middle, narrower than elytra without distinct lateral margins; head usually constricted
behind emarginate eyes; antennae filiform, insertions separated by entire width of frons,
elytra rarely more than 3 times as long as broad, punctures at least regular with
interspaces between rows at least as wide as punctures; occiput abbreviated
(Criocerinae) (Oulema) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 472
471' Pronotum as wide as elytra, usually with lateral margins; prosternum without antennal
grooves; antennal insertions nearly approximate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473
472(471) Basal segment of antennae yellow, rest of segments black; pronotum as broad as long,
lateral sides medially constricted, disc with a double row of coarse punctures and
impunctate basally; basal groove prominent (Fig. 876); dorsum yellowish brown except
black lateral margins of elytra and scutellum . . . . . . . . . . . . Oulema atrosuturalis (Pic)
472' Basal segment of antennae reddish; pronotum (Fig. 877) reddish brown, punctures
moderately dense and scattered, relatively dense at middle forming 2 light longitudinal
rows, basal grove not prominent; elytra blue with yellow thorax . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oulema oryzae (Kuwayama)
472'' All segments of antennae uniformly black; pronotal punctures sparse, central ones
arranged in 3 rows, rest impunctate (Fig. 878); body entirely blue or green with metallic
green elytra; pronotum, femora, tibiae, and tarsi reddish yellow, apices of tibiae and
tarsi darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oulema melanopa (Linnaeus)
473(471') Femora III slender, adapted for walking (Galerucinae) . . . . . . . . . . . . . . . . . . . . . 474
473' Femora III thicker than I and II, enlarged for jumping (flea beetles) (Alticinae) . . . . . 477
474(473) Pronotum longer than wide, slightly constricted below middle producing a minute
process, gena slightly shorter than smaller diameter of eye; anterior coxal cavities open
behind; elytral epipleuron wide basally, elytron (Fig. 879) yellow-brown with a midlon-
gitudinal black band not reaching apices and outer margins black; head black; antennae
dark brown to black except segments VIII and IX yellow . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medythia nigrobilineata (Motschulsky)
474' Pronotum wider than long, anterior and posterior lateral angles with thin setae; anterior
coxal cavities closed behind . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
475(474') Anterior and posterior pronotal margins concave or procurved, not subquadrate dor-
sally; elytral epipleuron suddenly narrowed at end of basal one-third and distinctly
narrower at middle; basal segment of tarsi III long and slender, one-fifth longer than
remaining segments combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
475' Pronotum subquadrate or rectangular, anterior margin straight, midposterior area
strongly indented; basal segment of tarsi III short and not slender, nearly as long as 2nd
segment (Fig. 880); African species . . . . . . . . . . . . . . . . . . Sesselia pusilla Gerstacher
476(475) Elytra (Fig. 881) black with 2 broad subglobular yellow spots before and behind middle;
abdomen reddish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monolepta signata Olivier
476' Elytra (Fig. 882) yellow with 2 transverse blackish brown bands, basal one runs from
suture to suture, the submedian band not reaching lateral wing margins; body and legs
yellow except brownish black head, apical antennal segments, mesopleuron and
mesosternum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monolepta bifasciata (Hornstedt)
477(473') African species of Chaetocnema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478
477' Oriental species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
478(477) Body length more than 3 mm; head and pronotum moderately punctated; space be-
tween antennal base as long as or slightly longer than scape; antennal segments III-VII
far more slender than the apically dark colored four segments; pronotum rounded on all
sides; inner striae of elytra rather oblique (Fig. 883) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocnema kenyensis Bryant
478' Body length, a little over 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
479(478') Pronotum approximately one-third wider than long, densely punctured, narrows toward
anterior and broadens below middle, anterior and posterior lateral angles more or less
rounded; vertex lightly punctated; antennae without very slender segments similar to C.
kenyensis ; elytral striae in regular rows, 1st or 2nd stria incomplete (Fig. 884) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocnema abyssinica Jacoby
479' Pronotal punctures moderately dense and coarse to almost smooth; posteromedian part
of pronotum margined or unmargined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
480(479') Posterior edge of pronotum margined, middle strongly concave; head almost smooth
(Fig. 885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocnema pulla Chapuis
480' Posterior edge of pronotum not margined; middle mildly concave; head punctated
(Fig. 886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocnema pallidipes Fairmaire
481(477') Interantennal space carinate or swollen medially, not punctated; vertex impunctate or
with few punctures; elytron with interstices flat mesally and weakly swollen laterally;
punctations rather irregular; pronotum with a punctured, faintly impressed line along
basal margin (Fig. 887); anterior femora reddish to pitchy brown with a pale apex . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocnema basalis Baly
481' Interantennal space flat and strongly punctured; vertex prominently punctured, elytral
punctations in 15 regular rows (Fig. 888) . . . . . . . . Chaetocnema concinnipennis Baly
482(462') Claw toothed with a fleshy appendage beneath; body clothed with long erect setae; head
quadrate, strongly transverse; pronotum reddish yellow and hexagonal; neck absent;
antennae 10-segmented, brown except three reddish yellow basal segments; legs I and
II brown and III bluish; elytra deep blue with a transverse reddish yellow band medially,
apical end of elytra with a small reddish yellow band; apex of sutural area of elytra open
V-shaped (Fig. 889) [Melyridae] . . . . . . . . . . . . . . . . . . . . Apalochrus rufofasciatus Pic
482' Claw simple; body without long erect setae; pronotum not hexagonal .......... 483
483(482') Head strongly deflexed and moderately globular, free from anterior margin of
pronotum; neck distinct; pronotum globular medially and constricted anteriorly and
posteriorly; elytral humeri obsolete; metathoracic wings vestigial; femur clubbed;
penultimate tarsal segments bilobed (Anthicidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 484
483' Head not deflexed, wider than long and deeply sunken in the concave anterior pronotal
margins; sides of frons strongly produced over the antennal insertions; eyes deeply
indented in between frontal flange and anterolateral part of pronotum; pronotum
subquadrate with a wide lateral margin and double convex posterior margin (Figs.
890-891); clypeus with a V-shaped cleft anteromedially (Fig. 890); body clothed with
yellow-brown clavate setae arranged in 9 bands in each elytron with 3-4 rows in each
band (Figs. 892-893), area in between bands smooth and blackish; basal 3 abdominal
sternites immovably connected [Tenebrionidae] . . . . . . . . . . . . . . Gonocephalum spp.
484(483) Elytra without any oblique white markings (Fig. 894); entire body and legs uniformly
brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Formicomus sp.
484' Elytra with oblique white markings (Fig. 895); basal one-half of antennae brownish and
blackish towards apex; basal portion of femur II and III pale; body black . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Formicomus braminus La Ferte Senectere
485(363') Front and hindwing similar in size, shape, and venation and extended well beyond tip of
abdomen at rest, hamuli and crossveins absent; wing scales ( = stumps) present; veins
M and Cu multibranched; fontanelle usually present; antenna moniliform; tarsi 4-seg-
mented (ISOPTERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486
485' Frontwing different from the hindwing in size, shape, and venation, and as long as or
slightly beyond tip of abdomen at rest, hamuli present; with or without crossveins in the
4 membranous wings; wing scales absent; veins M and Cu not multibranched; fontanelle
absent; antenna usually elbowed, clubbed or filiform; tarsi 3-5 segmented (HYMENOP-
TERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 488
486(485) Anterior wing scales covering at least the base of the posterior scales; wings slightly
reticulate, veins Sc + R and radial sector with 2 thin longitudinal lines near apex, radial
sector with 5-7, medius trifurcate apically; cubitus of frontwing with 8 branches and 11
in the hindwing (Fig. 896); pronotum large and cleft at its median anterior margin;
ocelli present [Rhinotermitidae] . . . . . . . . . . . . . . . . . Coptotermes formosanus Shiraki
486' Anterior wing scales short, not reaching to base of posterior scales; wings not entirely
reticulate [Termitidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487
487(486') Radial and median sector 4-branched at apices, cubitus multidichotomously branched
(Fig. 897); head ovoid with ocelli and a small fontanelle; small clypeus moderately
truncate apically, labrum dome-shaped; pronotum very wide along anterior margin and
narrow posteriorly (Fig. 898); African species . . . . Macrotermes natalensis (Haviland)
487' Radial and Sc + R (Fig. 899) almost fused at wing apex, unbranched, medius with 3
oblique branches in apical one-third; cubitus with 15 branches, tridichotomous at
middle; ocelli and fontanelle prominent; prothorax cleft along midanterior and mid-
posterior part (Fig. 900); Oriental species . . . . . . . . . . . . Macrotermes gilvus (Hagen)
487' Sc + R and radial sector fused towards apex and merged with tip of medius, medius
branch at midhalf with a pentadichotomous branch, cubitus with 14 single-branch veins
in the frontwing (Fig. 901); head subovoid with indistinct fontanelle, clypeus moderately
large, slightly cleft along apical margin to straight, deeply sunken in the concave frons;
labrum longer than wide, partly dome-shaped; pronotum (Fig. 902) subtruncate with a
minute cleft at midanterior margin, anterolateral angle slightly elevated, posterior
margin moderately indented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microtermes sp.
488(485') Hind tibial spur or spurs present, one (the medial when there are 2) modified into a
calcar through development of a comb of hairs or teeth on its inner tarsal margin; a
corresponding strigil or brush is developed on the slightly emarginate basitarsus; if both
spurs are non-simple or lacking, then at least some body hairs are plumed and the hind
basitarsus widened, or first 1 or 2 segments of metasoma forming a node . . . . . . 489
488' Hind tibia with 1 or 2 spurs (absent in a few Ichneumonidae), but none modified for
pruning; sometimes a dense brush of enlarged hairs developed on the basitarsus and
apex of tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519
489(488) Lateral pronotum and mesopleuron (or prepectus) overlapping and with considerable
free movement, lower portion of pronotal lobe rounded (pronotum and prepectus fused
in Dryinidae, but the antennae 10-segmented), spiracle cover lobe of pronotum not
margined with close fine hairs (prepectus visible); hindwings without closed basal cells
...................................................................... 490
489' Lateral pronotum and mesopleuron meeting with carinate margins and with very little
free movement between them, lower portion of pronotal lobe tapering to a point . . . . .
....................................................................... 503
490(489) Antennae 10-segmented in both sexes; females usually wingless with chelate foretarsi
[Dryinidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491
490' Antennae 13-segmented, apical segments globular; foretarsi of both sexes simple, not
chelate, apterous forms absent; head depressed to slightly convex; lateral ocelli near
posterior margin of head; face with a ridge in between antennal sockets; thorax without
humps or node-like patterns, smooth propodeum quadrate and margined laterally and
posteriorly; legs short, blackish brown with yellow tarsus; wings hyaline, pubescence
outside closed cells, stigma prominent (Fig. 903) [Bethylidae]; larval parasite of
Cnaphalocrocis and Marasmia leaffolders . . . . . . . . . Goniozus nr. triangulifer Kieffer
491(490) Both sexes fully winged; notaulices completely distinct, and jointed posteriorly in male;
female with testaceous head, antennae, and pronotum; abdomen similarly testaceous
except tergites 1, 2 and 4 partly brown; propodeum and petiole black; forewing
transparent without dark transverse bands; maxillary palpi 3-4 segmented and labial
palpi 2-segmented; segment 1 and 4 of foretarsus equal in length; enlarged claw with
subapical tooth and 4-5 lamellae; segment V of foretarsus with a single row of 9-12
lamellae, apex with a group of 6-11 lamellae (Fig. 904); aedeagus of male with a deeply
cleft apicomedian process (Fig. 905); hosts - N. lugens, S. furcifera, L. striatellus, P.
saccharicida, and Nephotettix spp. . . . . . . . . . . . . . . . Echthrodelphax fairchildii Perkins
491' Female apterous; male winged; other characters not as above . . . . . . . . . . . . . . . . 492
492(491') Enlarged claw without subapical tooth, with or without small tooth at the end of the
longitudinal furrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493
492' Enlarged claw with subapical tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 496
493(492) Pronotum not crossed by a transverse depression, if so, impression very weak; labial
palp 2-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494
493' Pronotum with a prominent transverse depression . . . . . . . . . . . . . . . . . . . . . . . . . . 495
494 Maxillary palpi 2-segmented; enlarged claw with 3 bristles at end of longitudinal furrow;
segment V with a row of 6 minute lamellae on distal half, apex with 6 lamellae (Fig. 906);
segment I of foretarsi longer than segment IV (11:9); black except testaceous antennae
and yellow legs (Fig. 907); hosts — Nephotettix cincticeps and N. nigropictus .........
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrodontochelys sakaii (Esaki & Hashimoto)
494' Maxillary palpi 3 to 4-segmented; enlarged claw as above but with 5 peg-like hairs;
segment V with a row of 16-20 lamellae and a group of 8-10 lamellae at apex, proximal
region with an inner serrate margin (Fig. 908); segment I of foretarsi as long as segment
IV; reddish testaceous with black petiole, brown vertex and abdomen; hosts - N.
virescens, N. nigropictus, and Recilia dorsalis . . . . . . . . . Tetrodontochelys lucens Olmi
495(493') Maxillary palpi 3-segmented; labial palpi 2-segmented; enlarged claw with a bristle and
3 peg-like hairs at end of longitudinal furrow; segment V of front tarsi with innerside
proximally not serrate, with 2 rows of 12 lamellae and a group of 6 lamellae at apex (Fig.
909); 1st segment of tarsi I the longest, two times longer than 4th and slightly longer than
5 th; black species except yellow malar space, clypeus, mandibles, and front of vertex,
and brownish antennae and legs; host — N. lugens . . . . . Gonatopus yasumatsui Olmi
495 Maxillary palpi 6-segmented; labial palpi 3- segmented; enlarged claw with 6 peg-like
hairs, innerside not proximally serrate but with 18-19 lamellae in 2 rows, apex with 14
lamellae in a group (Fig. 910); 1st segment of tarsi I two-thirds of 4th, 5th segment four
times longer than 1st; head brownish red with yellow mandibles, clypeus and front
vertex; antennae brown with segments 1 and 2 yellow; thorax and abdomen blackish
brown; host —Nephotettix virescens . . . . . . . . . . . . . . . . . . Gonatopus lucidus (Rohwer)
496(492') Pronotum without or very weak transverse depression; labial palpi one-segmented;
maxillary palpi 2-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497
496' Pronotum clearly with a prominent depression; labial palpi 2-segmented; maxillary
palpi 2- to 4-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 499
497(496) Female with abdomen entirely testaceous to occasionally brown; propodeum yellow
testaceous; enlarged claw with 3-6 lamellae, segment V with 2 rows of 7-10 lamellae and
a group of 2-7 lamellae at apex (Fig. 911); male with dorsal process of gonoforceps
distally broadened and serrated (Fig. 912); hosts — N. lugens, Laodelphax striatellus and
Sogatella furcifera . . . . . . . . . . . . . . . . . . . . . . . Haplogonatopus apicalis R.C.L. Perkins
497' Female abdomen fully black; male dorsal process of gonoforceps distally serrated or not
serrated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498
498(497') Dorsal process of gonoforceps distally serrated but not broadened (Fig. 913); segment
V with 12 lamellae (Fig. 914); host - L. striatellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplogonatopus atratus Esaki & Hashimoto
498' Dorsal process of gonoforceps long and slender, apex without a serrated margin (Fig.
915); segment V with 11 lamellae, enlarged claw with 4 (Fig. 916); host - L. striatellus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplogonatopus oratorius (Westwood)
499(496') Legs testaceous; thorax and propodeum red testaceous with yellow scutum; maxillary
palpi 2-4 segmented; sides of metanotum rounded; metathorax + propodeum with a
weak tract of median furrow; enlarged claw with subapical tooth and 6-7 lamellae,
segment V of foretarsus with 2 rows of 19-20 lamellae and a group of 5-8 lamellae at apex
(Fig. 917); basal segment of foretarsus as long as 4th; dorsal process of gonoforceps of
male slender, pointed at apex and with a subapical point (Fig. 918); hosts - N. lugens
and Sogatella sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogonatopus sarawaki Moczar
499' Legs mostly yellow; thorax, propodeum, and abdominal coloration more or less dif-
ferent (except red testaceous in P. nudus); sides of metanotum rounded or protruding;
dorsal process of gonoforceps of male different . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500
500(499') Sides of metanotum protruding, metathorax + propodeum without median furrow;
maxillary palpi 4-segmented; enlarged claw with a subapical tooth and 6 lamellae;
segment V of foretarsus with 2 rows of 14 lamellae and a group of 6 lamellae at apex
(Fig. 919); basal segment of foretarsi as long as 4th segment; general coloration reddish
testaceous except black petiole and brown-red abdomen (Fig. 920); hosts - N. lugens
and Sogatella furcifera . . . . . . . . . . . . . . . . . . . Pseudogonatopus nudus R.C.L. Perkins
500' Sides of metanotum rounded; body color black to brownish black . . . . . . . . . . . . . . 501
50l(500') Head, thorax, and propodeum brownish black; abdomen black; (Fig. 921); maxillary
palpi 2-segmented; mandible tetradentate (Fig. 922); metathorax and propodeum weak-
ly granulated; enlarged claw with subapical tooth and 4-5 lamellae, segment V of
foretarsus with 2 rows of 10-12 lamellae and a group of 5 more at apex (Fig. 923); basal
segment of foretarsi one-fifth longer than 4th (20:16); dorsal process of gonoforceps in
male broad and long, rounded at apex and with a lateral point (Fig. 924); hosts - S.
furcifera and L. striatellus . . . . . . . . . . . . . . . . . . . Pseudogonatopus fulgori (Nakagawa)
501' Not as above ........................................................... 502
502(501') Body uniformly black except brown head; propodeum without a yellow area at apex;
metathorax and propodeum dull and granulated (Fig. 925); maxillary palpi 4-seg-
mented; enlarged claw with a subapical tooth and 5 lamellae; segment V of foretarsi with
2 rows of 13 lamellae, apex with a group of 7 lamellae (Fig. 926); segment I of foretarsus
1.24 times longer than 4th (21:17); dorsal process of gonoforceps of male very short and
pointed (Fig. 926); hosts - N. lugens, S. furcifera, and Nephotettix spp. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogonatopus flavifemur Esaki & Hashimoto
Figs. 910-918
502' Thorax and propodeum black; propodeum with a yellow patch at apex; abdomen
brownish black and head brown; metanotum and propodeum with a tract of median
furrow and indistinctly sculptured; maxillary palpi 2-4 segmented; enlarged claw with
4-9 lamellae; segment V of foretarsus with 2 rows of 20-25 lamellae, apex with a group
7-10 more lamellae (Fig. 927); dorsal process of gonoforceps broadly long, pointed and
blade-like (Fig. 928); hosts – S. furcifera, N. lugens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
........................................ Pseudogonatopus hospes R.C.L. Perkins
503(489') Segments 1 or 2 of metasoma nodiform and sharply marked off from the rest;
posteroventral corners of thorax each with a metapleural gland [Formicidae] . . . . 504
503' Segment 1 of metasoma not node-like; metapleural glands absent . . . . . . . . . . . . . 509
504 (503) One petiole (node) present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505
504' Two petioles present ................................................... 508
505(504) Head and thoracic dorsum with prominent long grooves (deeply carinated) transverse
in the pronotum and mesoscutum and longitudinal in the mesoscutellum; anterolateral
angle of pronotum toothed (Fig. 929); right mandible with 5-teeth; clypeus with 7 blunt
teeth; frontal carina widely separated; antenna 12-segmented; veins Rs + M reaching
wing apices; first discoidal cell 5-sided; second discoidal cell trapezoidal; cubital cells 1
and 2 almost equal in size (Fig. 930); gaster attached to the alitrunk by a single node, top
of node thinly flat and concave at middle; black ants . . . . . . . . . . . . Odontopouera sp.
505' Head and thorax smooth to moderately punctated; venation variable .......... 506
506(505') Pronotum with anterolateral teeth, propodeum and petiole with a pair each of spines
(Figs. 931-932); petiolar spines projected sidewards or curved posteriorly (Fig. 933);
antennae 12-segmented (Fig. 933); first abdominal segment very broad, almost twice as
long as the total length of remaining segments; veins Rs and M diverging originating
from a common point at little beyond 7; inner tip of Cul curved downwards; Cu straight
basally, hump-like above tip of A and straight towards wing apex (Fig. 934); black ants
clothed with fine white hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polyrachis spp.
506' Not as above . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507
507(506') Head (sometimes finely carinated along sides), thorax, and abdomen (Fig. 935) clothed
with moderately long hairs, smooth and shiny black; frontal carina weakly raised; stigma
thin; veins Rs and M moderately diverging after a square plate junction at base of
crossvein r or after a small extension past base of same crossvein; apical half of Cu
convex and appears fused to tip of A above wing notch; discoidal cell very large; Cu-a
close to midhalf of M+Cu; M and Cu not reaching wing margins (Figs. 936-937);
antenna brown except black scape; petiole narrowed at tip in lateral view, subtruncate
in posterior view; abdominal segment II (T2) usually longer than either I or III (Fig. 938)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camponotus spp.
507' Head, thorax, and abdomen dull black clothed with short hairs, noticeably punctated
with transverse carinae in the propodeum and the petiole; frontal carina small but well
BARRION AND LITSlNGER 193
expanded; stigma rather large; Rs and M stems from different points; 1st discoidal cell
small and subrectangular; Cu-a close to apex of M + Cu; M and Cu slightly erased
towards their apices and parallel to each other (Fig. 939); antennae blackish red to
brown, black towards scape; petiole rounded at apex; abdominal tergite I as long as II
(Fig. 940) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diacamma sp.
508(504') First discoidal cell strongly triangular towards its base, vein M straight almost reaching
wing margin, Rs with a short extension after fusion with R1 prior to wing apex, very long
almost meeting Cu near wing notch; 1st cubital cell narrowed towards median cell,
Sc + R unparallel to Rs + M (Fig. 941); propodeal spiracle elongated; nodes subglobular
with flat top; pedicel a little longer than wide . . . . . . . . . . . . . . . . . . . Pheidologeton sp.
508' First discoidal cell trapezoidal, vein M very short and Rs not reaching wing margin and
never fused to R1; vein A short not meeting Cu; 1st cubital cell narrowed on both ends
with Sc+ R parallel to Rs +M (Fig. 942); propodeal spiracle subglobular; 1st node
divided at middle dorsally; pedicel globular . . . . . . . Solenopsis geminata (Fabricius)
509(503') Posterior lateral lobes of pronotum not reaching back to and terminate below tegula;
hairs on body unbranched; notaulices wanting or feebly developed; jugal lobe in hind-
wings; one interradial crossvein present in forewings [Sphecidae] . . . . . . . . . . . . . 510
509' Posterior lateral lobes reach above tegula, usually angulated; forewing longitudinally
folded at resting position, 3 marginal cells present; tegula with or without raised margin;
tarsal claws bifid to simple [Vespidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 512
510(509) Gaster with a long yellow petiole, longer than gaster or femur III (Fig. 943); pronotum
with a pair of yellow lobes; mesoscutellum and metanotum yellow similar to apical
one-third of femur and tibia of legs I and II, trochanter III, base of femur III, basal
one-half of tibia III and basal tarsal segment of leg III; midtibia with 2 spurs; forewing
with 6-sided 2nd cubital cell, apex of cubital protruded and rounded (Fig. 944); hindw-
ing with 2 distinct 2nd anal veins well separated from 1A, jugal lobe without notch . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sceliphron madraspatanum conspicillatum (Costa)
510' Gaster subsessile; 2 posterior ocelli more or less distorted (not round and convex),
flattened and reduced; posterior border of pronotum straight viewed dorsally; tarsal
claws without tooth; pygidium of female very polished . . . . . . . . . . . . . . . . . . . . . . 511
511(510') Abdomen and hind femur black; fore tibia without dorsal spines, 2nd antennal segment
glabrous and polished, segment III not nearly twice as long as segment IV (Fig. 945);
wings dark fuscous brown; disc of propodeum wrinkled in female; male antennal
segment III as long as or a little shorter than IV (Fig. 946); clypeus gently rounded in its
entire margin to very weakly but broadly emarginate sides; gaster smooth and polished;
pygidium broad with arcuate sides . . . . . . . . . . . . . . . . . . . . . Larra carbonaria (Smith)
511' Abdomen polished and reddish except 3 black apical segments; fore tibia with dorsal
spines; 2nd antennal segment pilose, III nearly twice as long as IV (Fig. 947); wings
clear; disc of propodeum with a faint median carina, basally with separate punctures and
densely reticulate in female; male antennal segment III noticeably longer than IV (Fig.
Figs. 929-938
948); clypeus produced medially into a polished and distally impunctate and very
shallow emarginate lobe; gaster impunctate in female and banded with silvery file in
male; pygidium very smooth with delicate punctures . . . . . . Larra sanguinea Williams
512(509') Tarsal claws bifid; tegula with raised margins, apical half dark brown; mandibles long
with visible serration, projected downwards; yellow except black vertex, mesoscutum
(Fig. 949), anterior and posterior margins of mesoscutellum, apical and median part of
propodeum, subapical area of tergite I, midhalf of II, basal two-thirds of IV and most
lateral part of thorax except yellow band in propleuron and mesopleuron; tergite II with
a pair of ovoid yellow spots behind black band (Fig. 950); discoidal cell 3 not rectangular
with an oblique recurrent vein 2 connected to base of 2nd cubital cell; antennal segment
III as long as combined length of IV and V . . . . Eumenes campaniformes (Fabricius)
512' Tarsal claws simple; tegula without raised margins . . . . . . . . . . . . . . . . . . . . . . . . . . . 513
513(512') Reddish brown; apical part of abdominal tergite II with a prominent yellow band, with
or without yellow spots towards base; mesoscutum without parallel yellow stripes;
mesopleuron with or without yellow band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 514
513' Yellowish black to brown; basal portion of abdominal tergite II broadly yellowish to
light brownish yellow or with apical margin narrowly banded yellow; mesoscutum always
with a pair of yellow parallel longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515
514(513) Propodeum with a pair of basal convergent carinae running from posterolateral angle
of metanotum toward muscle slit base; muscle slit narrow and pointed (Fig. 951); basal
portion of abdominal tergite II with a pair of prominent yellow spots obliquely cutoff at
the base (Fig. 952); tergite I swollen posterolaterally and anterodorsally (Fig. 953);
mesopleuron with a yellow band running parallel to the yellow band on sides of
pronotum . . . . . . . . . . . . . . . . . . . . . . . . . Ropalidia marginata sundaica van der Vecht
514' Propodeum without such carinae; muscle slit broad and rounded anteriorly; basal
portion of tergite II usually without yellow spots or highly reduced (Fig. 954), apical
portion with a broad yellow fascia; tergite I parallel-sided at base anterodorsally,
swollen posteriorly but narrowed towards apex in posterior one-quarter (Fig. 955);
mesopleuron without yellow band . . . . . . . . . . . . . . . . . . Ropalidia fasciata (Fabricius)
515(513') Mesoscutum without parallel longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . 516
515' Mesoscutum with parallel longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 517
516(515) Clypeus yellow V-shaped, pointed apicomedially with brown patches in the central area
and brown V-shaped mark below antenna (Fig. 956); vertex black, reddish brown
behind and below ocelli up to eye margins and black near antennal base; pronotum
reddish brown except thin yellow apical margin; hump-like mesoscutellum and
metanotum with a pair each of yellow spots; propodeum without yellow markings;
abdomen black except apices of tergite I and II yellow (Fig. 957); legs yellow to reddish
brown, blackish brown in femora II and III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ropalidia nr. philippinensis (Saussure)
516' Clypeus cleft medially producing 2 prominent teeth and entirely yellow (Fig. 958);
dorsum of head black with a deep and broad pit behind ocelli; yellow bands at the back
and front of eyes towards emargination and a triangular yellow band above antennal
base (Fig. 959); entire pronotum yellow with brown tinge along margins; mesoscutellum
and metanotum with transverse rectangular yellow bands; abdominal tergite I reddish
brown with a black base and yellow anterior, tergites III-VII yellow, II red and black
with a pair of yellow spots near basolateral and yellow entire apical margin (Fig. 960) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ropalidia sp. A
517(515') Clypeus uniformly yellow, clothed with fine hairs and a small black spot at base of
antennae; yellow band emanating from antennae directed to but not touching the ocelli;
vertex black with a thin oblique yellow stripe between eyes and lateral ocellus (Fig. 961);
yellow part of pronotum reaching tegula; mesopleuron with a large yellow band;
metapleuron with a yellow band at middle; abdomen yellowish except a broad subapical
brown band in tergite II, a thin narrow band on apex of tergite III, and entire terminal
segment (Fig. 962); coxae II and III mostly yellow on apical one-half . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ropalidia flavopicta flavobrunnea van der Vecht
517' Clypeus with a broad median longitudinal black band (Fig. 963) and sparsely haired;
yellow spot present above antennae; vertex black without yellow oblique stripe between
eyes and lateral ocellus (Fig. 963); tergites III and rest of apical segments brown, II
yellow apically and basally but black submedially (Fig. 964); mesopleuron with a small
yellow band, metapleuron and coxae II and III black . . . . . . . . . . . . . Ropalidia sp. B
517'' Not with the combination of characters above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 518
518(517'') All legs black except yellow apice of femora I-III and parts of all coxae; yellow band on
propodeum hook-shaped along spiracular area (Fig. 965); tergite I short, posterior
two-thirds high or swollen in lateral view and yellow basally with a lateral protruding
yellow band (Fig. 966); tergite II with a broad nearly medially contiguous dorsobasal
lateral spot (Fig. 967); apices of rest of tergites yellow; clypeus with a median dark spot
similar to (Fig. 961); mesoscutum with a yellow median band; metanotum yellow with a
median dark band (Fig. 968) . . . . . . . . . . . . . . . . . . Ropalidia cyathiformis (Fabricius)
518' Femora I to III yellow with black bands along the basodorsal one-half; tibia and tarsus
of leg I brownish yellow, tibia II blackish brown in basodorsal two-thirds, and tarsus II,
tibia and tarsus of leg III blackish brown; propodeum entirely yellow without a hook-
shaped area near spiracles; tergite I moderately long, yellow apicoposteriorly and
extended to the spiracles, II with alternating brown and yellow transverse bands, III
median transverse yellow band constricted in the middle (Fig. 969), metanotum entirely
yellow, mesoscutum with a pair of yellow longitudinal bands; clypeus yellow, front of
head with an ovate black band running from ocellar region to below antennal base (Fig.
970); submedian yellow band of tergite II broad and transversely uninterrupted . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ropalidia sp. C
519(488') Pronotum with at least a lateral spiracle cover lobe reaching back to tegula, or, if
wingless, lobe margined with close fine hairs; lateral pronotum vertically grooved for
reception of fore femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520
519' Pronotum not reaching back to tegula (almost so in some Leucospidinae, in which
tegula is lengthened, and in some Mymaridae); forewing with venation very reduced,
without fully formed cells; pronotum separated from tegula by prepectus . . . . . . 637
520(519) Spiracle cover lobe of pronotum without close fine hairs along its margin . . . . . . 521
520' Spiracle cover lobe of pronotum margined with close fine hairs, anal lobe of hindwing
notched; hindwing without jugal lobe; apex of wing with close pseudovenation; tibia II
with 1 spur; eyes emarginate; stout bodied and densely haired wasps [Scoliidae] . . . 596
521(520) Lateral area of pronotum without a vertical groove for reception of femora I; scutellar
cup elevated and well developed; mesonotum with parapsidal furrows; abdomen
without hairy ring on tergite 2; antennae 13-segmented in the female and 15-segmented
in male [Cynipidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 522
521' Lateral pronotum vertically grooved for reception of front femora; scutellar cup absent;
costal cell present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 523
522(521) Third antennal segment the longest, as long as combined length of scape and pedicel;
segment 4 to 9 rather stout and equally long; segments 10 to 15 slightly shorter than each
of 4 to 9; segments 9-13 light brown (Fig. 971); scutellar cup very close to the posterior
end of thorax, sides with setae and network-like subglobular to rectangular marks (Fig.
972); radial cell subtriangular with unequal “arms” (15:25), inner angle of cell rounded
opposite vein (Fig. 973); hosts – Hydrellia and Notiphila pupae . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucoilidea sp. A
522' Third antennal segment shorter than 4th, seventh antennal segment longest, as long as
each of segments 8 to 15 and 1.45 times longer than 3rd; 4th to 6th almost subequal in
size and length (Fig. 974); segments 1-4 yellowish brown; scutellar cup far from
posterior end of thorax, sides with indistinct network-like marks (Fig. 975); radial cell
triangularly thinner than Eucoilidea sp. A and with equally long arms, inner angle of cell
rounded opposite small crossvein above horizontal vein (Fig. 976); hosts – Hydrellia
and Notiphila pupae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucoilidea sp. B
523(521') Costal cells absent, but distinct marginal vein from base or a pterostigma; hindwing with
at least one closed basal cell (except in Aphidiinae) . . . . . . . . . . . . . . . . . . . . . . . . 524
523' Costal cells present, or venation greatly reduced; hindwing without closed cells . . 604
524(523) Forewing with 1 or without recurrent vein; hindwings with median cell not extending to
base of marginal vein [Braconidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
524' Forewing with 2 recurrent veins (rarely one); hindwing with median cell elongated
beyond base of marginal vein [Ichneumonidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 557
525(524) Mandibles widely separated, tips not touching when closed; head transverse with
posterior margin of vertex emarginate; wings hyaline with dark brown stigma and veins,
radial cell large, ending before apex of wing; recurrent nervure received in 1st cubital
cell at a far distance from its apex; radial cell of hindwing petiolate (Fig. 977); antenna
with 41-42 segments in both sexes; prothorax honey-yellow in females and yellowish
brown in males; host – Chlorops oryzae; Japanese species . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coelinidea oryzicola Watanabe
525' Mandibles in usual position, tips touching when closed (Fig. 978) . . . . . . . . . . . . . . . 526
526(525') Clypeus semicircularly emarginate at apex forming a circular opening with the man-
dibles (Fig. 979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 527
526' Clypeus not emarginate at apex, not forming such an opening with the mandibles; if very
rarely with a long narrow opening (e.g. some Opiinae), then first orbital and first
discoidal cells separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 535
527(526) Abdomen petiolate, basal segment with 3 constrictions, longest one occurs before
widened apex (Fig. 980); brownish yellow to dark brown; wings hyaline; legs pale yellow,
ovipositor as long as 2nd and following abdominal segments together; body length 3-4
mm; host – Chilo suppressalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spathius helle Nixon
527' Abdomen sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528
528(527') Occiput not margined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 529
528' Occiput margined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 532
529(528) Second tergite of abdomen with oblique furrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . 530
529' Second tergite of abdomen without oblique furrows . . . . . . . . . . . . . . . . . . . . . . . . . 531
530(529) Large species, 9-13 mm long; abdomen elongated, with elevated median area of tergite
II smooth and strongly defined (Fig. 981); reddish yellow with 6th abdominal tergite
black; wings yellowish, subhyaline with apical margins fuscous, forewing with three
blackish brown spots; ovipositor very long, 2 times as long as body (Fig. 982); hosts –
Sesamia inferens, Scirpophaga incertulas . . . . . . . . . Stenobracon nicevillei (Bingham)
530' Smaller species, 5-6 mm; abdomen oblong with median raised area of 2nd tergite weakly
defined and rugose with a V-shaped mark (Fig. 983); yellowish brown to brown, females
generally darker than males; wings fuscous to subhyaline; ovipositor short, as long as
half of abdomen (Fig. 984); hosts – S. incertulas, Sesamia inferens, Chilo suppressalis .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tropobracon schoenobii (Viereck)
531(529') Head and thorax yellowish red; abdomen black with a white band on each tergite; plate
of first tergite parallel-sided, 3 times as long as apical breadth (Fig. 985); ovipositor as
long as hind femur (Fig. 986); body length 3.5-4 mm; hosts similar to T. schoenobii . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bracon chinensis (Szepligeti)
531' Body reddish yellow with black markings; plate of lst tergite narrowed basally, less than
2X as long as apical breadth (Fig. 987); head almost parallel-sided behind eyes,
transverse with immargined occiput (Fig. 988); ovipositor as long as half of abdomen
(Fig. 989); body length 3.5-4 mm; hosts - C. suppressalis, S. inferens . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bracon onukii Watanabe
532(528') Second cubital cell distinctly smaller than the lst (inner) cubital (Figs. 990-992); 2nd
intercubitus prominent; submedian cell of forewing with nearly parallel margins; ner-
vulus straight, emanates from about one-third basal length of l st discoidal cell;
transverse Cu1 less oblique or slightly straight; brachial cell angled in its basoapical
border; propodeum quadrate (Fig. 990) or narrowed posteriorly (Figs. 991-992); ter-
gites 1-3 yellow brown to dark reddish brown; host – C. suppressalis . . . . Rogas spp.
532' Uniformly yellowish brown except a pair of light reddish brown longitudinal bands that
converge posteriorly in the median mesoscutum; head in top view not quadrate; width
as long as mesothorax; lateral ocelli 2 ocellar diameters away from eyes; transverse
cubital I noticeably oblique; median of propodeum with a long longitudinal carina
extending to tergites I and II; tip of abdomen more or less rounded dorsally (Fig. 992)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rogas narangae Rohwer
532'' Second cubital cell as long as first cubital; inner end of 1st intercubitus and 2nd
intercubitus usually pale; apical end of 1st brachial cell rounded and 4.7 times longer
than wide; postnervellus long; abdomen sessile with tergites 2 + 3 divided by 1 or 2
transverse furrows; ovipositor about two-thirds as long as abdomen . . . . . . . . . . . 533
533(532'') Abdomen with tergites 2 + 3 divided by 2 transverse furrows into three parts, the middle
one being short and fusciform (Fig. 993); dark brown to black; head not transverse, area
behind eyes unparallel (Fig. 994); legs honey-yellow; wings subhyaline with pale yellow
stigma; body length 3.5-4 mm; host – S. incertulas in Java and Thailand . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhaconotus schoenobivorus (Rohwer)
533' Tergites 2 + 3 divided by a single transverse furrow into 2 parts . . . . . . . . . . . . . . . 534
534(533') Antenna with 44-47 segments; mesonotum a little longer than its maximum width;
propodeum ill defined dorsally, with only fine sculpture between basal and inner carina;
hairs on vertex, pronotum, and mesonotum conspicuous; tergites evenly carinate lon-
gitudinally except the broadly transverse tergite (Fig. 995) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Rhaconotus scipophagae Wilkinson
534' Antenna with 30-32 segments; mesonotum shorter than its maximum width; dorsal areas
of propodeum very distinct provided with scaly reticulation; hairs on vertex, pronotum,
and mesonotum not conspicuous; 1st and 2nd tergites entirely carinate longitudinally,
3rd and 4th uncarinated in the apical half, 3rd sometimes smooth, 5th carinated only on
side (Fig. 996) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhaconotus oryzae Wilkinson
535(526') Forewing with 2 cubital cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536
535' Forewings with 3 cubital cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 551

536(535) Abdomen inverted bath tub-like (broadly convex), apical end slightly wide to strongly
rounded; segmentation absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 537
536' Abdomen not as above, normal in form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 538
537(536) Abdomen almost parallel-sided, with a pair of broad pale yellow markings towards base
(Fig. 997); body length 6-8 mm; host – Chilo suppressalis in East Asia ..............
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelonus munakatae Munakata
537' Entire dorso-basal segment of abdomen yellow, tip of abdomen broader than base (Fig.
998); body length 5-5.8 mm; hosts – C. medinalis & Marasmia spp. in Southeast Asia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelonus sp.
538(536) Female antennae “aberrant”, scape slender and very long with an apophysis in the basal
third, entire length a little over one-third of antennae; funicular segment VII with an
apico-ventral protrusion (Fig. 999); 1st metasomal tergite petiolate with deep dorsopes
(Fig. 1000); notauli complete, rugose posteriorly; ovipositor straight and its sheath
slender; emergence hole of cocoon subterminal and irregularly round; host – Medythia
nigrobilineata in East Asia . . . . . . . . . . . . . . . . . . . . . . . . Streblocera okadai Watanabe
538' Antennae of both sexes normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539

539(538') Radial cell closed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 540
539' Radial cell not clearly defined, radius weakly developed . . . . . . . . . . . . . . . . . . . . . 541
540(539) Radius concave; median cell large indistinct or vague medius; first discoidal cell 5-sided
and subtriangular towards large stigma; subdiscoideus incomplete (Fig. 1001); head
slightly transverse with occipital carina complete; eyes slightly convergent with scattered
minute hairs; ocellar triangle not well raised and lateral ocelli closer to compound eyes
than to each other (Fig. 1002); propodeum with a strong transverse carina; first ab-
dominal tergite sessile and striate longitudinally (Fig. 1003); ovipositor curved ventrad;
antennae with 21-22 segments; antennal sockets opposite midlength of eyes (Fig. 1004);
host - M. nigrobilineata in East Asia . . . . . . . . . . Centistes medythiae Maeto & Nagai
540' Radius straight; median cell relatively small with prominent medius; first discoidal cell
4-sided and subtrapezoidal; cubitus faint, similar radiella and cubitella of hindwing;
subdiscoideus complete (Fig. 1005); head subglobular with occipital carina dorsally
interrupted; eyes smooth; ocellar triangle nearer each other than to the compound eyes;
propodeum very finely punctated and smooth; orange bodied except hyaline wings and
blackened ocellar triangle; hosts – Cnaphalocrocis medinalis, Marasmia spp . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orgilus ashmeadi Viereck
541(539') Hypopygium relatively large with a series of striae medially; head twice as long as broad,
minutely punctate except coarsely punctate face; metanotum with median small cup-
shaped area surrounded by a ridge; propodeum with indistinct areola and costulae
(more or less obscured by coarse rugose sculptures); tergite I of abdomen twice as long
as wide basally and rugosely punctate; tergite II transversely rectangular; and much
wider than long (Fig. 1006); ovipositor sheath as long as hindtibia; black except legs with
coxae, 2nd and 3rd abdominal tergites reddish yellow; wings hyaline brown with
moderately large pterostigma (Fig. 1007); leg III black except reddish middle tibia, most
of femur, trochanter and coxa (Fig. 1008); hosts C. polychrysus, S. incertulas . . . . . . . . .
.................................. Exoryza ( = Apanteles) schoenobii (Wilkinson)
541' Hypopygium usually short, sides evenly sclerotized, without a series of longitudinal
creases near middle; tergite II at least half as long as III and subrectangular; ovipositor
sheath short and mostly concealed by hypopygium . . . . . . . . . . . . . . . . . . . . . . . . . . . 542
542(541') First tergite of abdomen very narrow, parallel-sided and 3 times as long as broad (Fig.
1009); head particularly vertex, frons, and temples smooth and impunctale; mesoscutum
with numerous and distinct punctures anteriorly; propodeum rugulose, subopaque with
weak costulae and indistinct areola; ovipositor sheath black, approximately one-third as
long as abdomen, distinctly curved downward; coxae I to III black, all femora, apical half
of hind tibia, and hind tarsus basally brownish black (Fig. 1010); hosts — Cnaphalocrocis
medinalis, Marasmia spp. . . . . . . . . . . . . . Cotesia ( =Apanteles) angustibasis (Gahan)
542' First tergite variable and without the combination of above characters .......... 543
543(542') Areola strongly V-shaped at apex (Fig. 1011); head mostly punctate; mesonotum rather
irregularly strongly punctate with rugose fine striae along sides of notauli; first tergite as
broad at base as at apex, widest in apical third, with an excavated area reaching apex of
tergite just beyond the strongly tumescent median; second and third abdominal tergites
reddish yellow; ovipositor sheaths as long as or slightly longer than the hind femur; hind
femur mostly reddish yellow with a fuscous mark at extreme apex (Fig. 1012); hosts —
C. medinalis, Herpetogramma ( = Pachyzancla) stultalis . . . . . . . . . . . . . . . . . . . . . . . . . . .
....................................... Cotesia (=Apanteles) opacus (Ashmead)
543' Areola not V-shaped .................................................... 544
544(543') Areola large, subquadrate with an open extreme base, carinae and costulae strong; first
abdominal tergite (Fig. 1013) slightly swollen at midlength and lightly rugose, unsculp-
tured and smooth apically; tergite II trapezoidal, basal area of tergite (2 + 3) only about
half as long as the rest of the segment beyond it; ovipositor sheaths (Fig. 1014) shorter
than hind femur, apical attenuation as long as 4th segment of tarsus, very thick, curved
and strongly tapering from base to apex; all femora, tibiae, and tarsi red testaceous;
antennae dark red-brown to black; costal veins testaceous; stigma light brown; host —
Spodoptera litura . . . . . . . . . . . . . . . . . . . . . Cotesia ( = Apanteles) prodeniae (Viereck)
544' Areola absent; if present not subquadrate; lst and 2nd tergite of different shape and
sculpture; ovipositor length variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545
545(544') Areola and costulae present .............................................. 546
545' Areola and costulae absent ............................................. 547
546(545) Solitary cocoon white; host – Cnaphalocrocis medinalis; vannal lobe feebly concave
beyond its widest part and without projecting hairs; tergite I of male more or less
narrowed behind, and very mildly narrowed in the female; tergite 2+3 black; stigma
hyaline 1st transverse cubital very narrow; ovipositor about three-quarters as long as
hind tibia (Fig. 1015) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cotesia cyrpris Nixon
546' Cocoons white, gregarious and clustered together to form a narrow elongate mass
covered with rather loose silk; vannal lobe evenly convex beyond its widest part and with
distinct hair fringe throughout, Cul strongly triangular apically, radius far from cubitus
(Fig. 1016); tergite I parallel-sided and without longitudinal carina, slightly turned over
and down, slightly excavated basally and partially swollen medially and apical two-thirds
indefinitely sculptured; tergite II smooth, apex evenly and strongly rounded (Fig. 1018);
ovipositor sheath (Fig. 1017) strongly down-curved, longer than hind tibia, its attenua-
tion almost as long as two-thirds of hindtarsus . . . . . . . . . . Cotesia baoris (Wilkinson)
546'' White cocoons either solitary or gregarious; tergite I (Fig. 1019) barely tumescent
medially; 2nd tergite smooth and unsculptured; propodeum smooth with open areola;
fore and mid legs, hind femur completely darkened; and basal half of hind tibiae red
testaceous; antennae dark brown; stigma and wing veins brown; wings slightly infumated
evenly throughout; ovipositor sheaths as long as or slightly longer than hind tibia; hosts
– hesperiid and rice leaffolder larvae . . . . . . . . . . . . . . . . . . Cotesia agilis (Ashmead)
547(545') Thorax dorso-ventrally flattened; head in lateral view not usually prominent below
antennae, largely impunctate; propodeum evenly rugolose; 1st tergite constricted sub-
basally and broadened apically; 2nd tergite punctated with ovoid-shaped mark; legs and
antennae red testaceous; tegulae, stigma, and costal veins (Fig. 1020) testaceous; tergite
I of abdomen widened behind, apicolateral corners weakly and roundly constricted,
posteromedian margin of tergite I slightly cleft (Fig. 1021); ovipositor sheath short . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . Cotesia flavipes Cameron
547' Thorax not dorso-ventrally flattened; head in lateral view usually prominent below
antennae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 548
548(547') Hind coxa roughened and rugose; tegulae red testaceous ................... 549
548' Hind coxae smooth, at most closely punctate; tegulae of different coloration ... 550
549(548) Solitary parasitoid; scutellum densely rugose-punctate all over without any smooth
interspaces anteriorly; fore and middle coxae invariably black; tergite III covered all
over with fine setae except for small midbasal area; tergites I and II with subquadrate
network like punctures except narrowed and smooth base (Fig. 1022); cocoons pale
cream; hosts – Plutella xylostella, Spodoptera litura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cotesia plutellae (Kurdjumov)
549' Gregarious parasitoid; punctures of scutellum at least on anterior half, wide enough
apart to leave smooth shiny interspaces fully equal to width of punctures; pterostigma
hyaline and shorter than R1; r and r-m meeting at a distinct angle (Fig. 1023); fore and
middle coxae yellowish brown; tergite I wider at apex than base, black and rugulose, II
aciculate with diverging sulci (Fig. 1024); tergite III with a single transverse row of few
fine setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cotesia ruficrus (Haliday)
550(548') Propodeum without a transverse basal carina; hypopygium acute; ovipositor very short;
first tergite gradually narrowed towards base, 1st and 2nd reticulate – rugose with
median basal one-third of 1st and median apex of 2nd smooth, apex of first somewhat
cleft and that of second uniformly pitted (Fig. 1025); legs with coxae reddish yellow and
dark brown tarsus; tegulae, stigma, and veins yellowish brown (Fig. 1026); hosts –
Mythimna unipuncta, Naranga aenescens . . . . . . . . . . . . . . Cotesia kariyai (Watanabe)
550' Propodeum with a strong basal transverse carina and a longitudinal median carina;
hypopygium obtuse; ovipositor sheath about half the length of hind tarsus; 1st tergite
smooth in the excavate basal one-third, rugulose laterally, narrowed immediately after
the base then widens apically with rounded apicolateral margins and slightly emarginate
apex; 2nd tergite with median disk sculpture, apical margins without uniform pits and
smooth laterally (Fig. 1027); legs including coxae brownish yellow except hind coxa dark
brown to black with extreme apex and ventral side reddish yellow; scape, flagellum,
tegulae and costal veins dark brown; hosts – Busseola fusca, Chilo ignefuscalis, C.
zonellus, Sesamia calamistis, S. cretica in Africa . . . . . . . Cotesia sesamiae (Cameron)
551(535') Second cubital cell small and triangular-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 552
551' Second cubital cell moderately large and subrectangular . . . . . . . . . . . . . . . . . . . . 555
552(551) Ovipositor sheath entirely hairy, arising distally from valvifer, a little shorter than half as
long as abdomen, and half as long as hind tibia (Fig. 1028); hypopygium rather large and
sclerotized evenly across middle and not folded; radius, cubitus, and subdiscoideus all
effaced, Cul strongly curved outward (Fig. 1029); black with three basal tergites yel-
lowish red; legs reddish yellow; host – Chilo suppressalis . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hygroplitis russatus (Haliday)
552' Ovipositor sheath with only few hairs distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 553
553(552') Prepectal carina present; notauli deep; radial cell open with base of radius effaced;
submediellan cell triangular-like towards base; cubitella more or less straight; abdomen
oblong elongate, black except yellow lateral patches in the basal part; hind tibia
darkened in the apical half (Fig. 1030); hypopygium short and evenly sclerotized
ovipositor sheath very short; host – S. litura . . . . . . . . Snellenius manilae (Ashmead)
553' Prepectal carina absent; notauli less developed to absent; hypopygium usually
moderately small and elongated medially; ovipositor sheath arises proximally from
second valvifer, radius almost entirely effaced; submediellan cell rounded or angled
distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 554
554(553') Nervellus strongly rounded; abdomen almost completely darkened except yellow
patches along lateral margins of basal tergites (Fig. 1031); host – S. litura . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microplitis tuberculifer (Wesmael)
554' Nervellus oblique forming an angulated corner at apex of submediellan; abdomen not
dark colored except apical three tergites and apical margins 6th tergite (Fig. 1032); host
– S. litura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microplitis pallidipes Szepligeti
554" Nervellus similar to M. pallidipes; wings hyaline; body typically black; tibia III light
colored at middle and dark on both ends (Fig. 1033) . . . . . . . . . . . . . . . . Microplitis sp.
555(551') Eyes hairy (Fig. 1034); entire body black including legs and with white hairs; forewing
with brown infuscation after stigmal vein; hindwing with similar infuscation along costal
area, basal one-fourth and apical one-fourth (Fig. 1035)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cardiochiles philippinensis Ashmead
555' Bases and wing tips of both pairs of wings brownish (Fig. 1036) . . . . . Cardiochiles sp.
555" Eyes bare; body not black; wings hyaline except brown veins and stigma . . . . . . . . 556
556(555") Ovipositor very long, almost twice as long as the abdomen; abdomen slender, brownish
black dorsally with longitudinal striae and yellowish brown edges; venter entirely
orange-brown; basal two tergites very slender and flattened; middle lobe of mesonotum
more bulging than lateral lobes and prominently separated from the latter by the deeply
pitted notauli; coxa III evenly swollen with a brown patch apico-dorsally (Fig. 1037);
hosts – Cnaphalocrocis medinalis, Marasmia spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrocentrus philippinensis Ashmead
556' Ovipositor very short, moderately exerted beyond tip of abdomen; tergite I of abdomen
rather rough with a slightly raised median longitudinal carina, carina narrow basally and
widens apically; tip of abdomen much broader than basal tergites (particularly I) and
curved slightly downwards; notauli absent; all legs including coxae yellow (Fig. 1038);
host – Hydrellia philippina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Opius barrioni Fischer
557(524') Abdomen flattened, segment III and following segments wider than deep . . . . . . 558
557' Abdomen strongly compressed in apical half, segment III and following segments
thicker (in terms of height) than wide; spiracle of tergite I distinctly distad of the middle;
ovipositor with a preapical dorsal notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 582
558(557) Spiracle of tergite I at or basad of the middle; flagella of female antennae without a
white ring medially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 559
558' Spiracle of tergite I distad of the middle; flagella of female always with a median white
ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 570
559(558) Tergite VII with a white apico-median transverse band; tergite I fused with its sternite,
sclerotized portion reaches apical one-third of tergite; second recurrent vein opposite
intercubital vein; areolet of forewing absent; sternaulus long and rather distinct . . 560
559' Tergite VII without a white marking; tergite I not fused with its sternite, sclerotized
portion of the sternite reaches basal one-third of tergite; areolet of forewing present;
sternaulus weak or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 561
560(559) Tergites II and III entirely black; tergite I, black and reddish apico laterally (Fig. 1039);
leg coloration similar to A. a. schoenobii; hosts – S. incertulas, S. innotata . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amauromorpha accepta metathoracica Ashmead
560' Tergite III and apical one-third – half of tergite II red (Fig. 1040); tergite I red,
sometimes blackish in part; mid and hind coxa red except apex narrowly black; fore coxa
and all trochanters black; hind femora black to blackish brown basally; hind tarsus black
and tibia pale basally; male with subtegular ridge and a pair of marks on face white;
tergite IV and V of male entirely red; hosts – S. inferens, C. suppressalis, S. innotata,
and S. incertulas in India, Malaysia, Thailand, Sabah, Taiwan, China . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amauromorpha accepta schoenobii (Viereck)
561(559') Ovipositor with a preapical dorsal notch (Fig. 1041); epomia absent; tergite I twice as
long as wide, II-IV without tubercles, II and III only slightly longer than wide or
subequal; tergites black except apices of I-III narrowly (in females) or broadly (in
males) brownish red; mandibles yellow; clypeus obscurely reddish in females and yellow
in males; hindlegs of female entirely blackish brown to black and reddish brown in the
males; host – C. suppressalis in Japan and China . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lampronota mandschurica (Uchida)
561' Ovipositor without a preapical dorsal notch; epomia present; tergite I as long as wide or
slightly longer than wide, II-IV each with a pair of tubercles, II and III broader than long
....................................................................... 562
562(561') Mandibles twisted apically, lower tooth hidden when mandibles are closed; propodeum
more or less areolated; body and legs yellow with or without black markings . . . . . 563
562' Mandibles not twisted towards apex; propodeum usually with only a pair of dorso-
median short longitudinal carinae; body mainly black, legs variously colored but not
mainly yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 568
563(562) Abdomen entirely without paired black spots (Fig. 1042) . . . . . . . . . . . . . . . . . . . . . 564
563' Abdomen with paired black spots; mesoscutum with black spots; propodeum with or
without black spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 565
564(563) Areola receives costulae behind center, 0.82-1.35 times as long as wide; midapical
transverse carina half the length of median longitudinal carina below level of spiracle
(Fig. 1043); ovipositor sheath 4.8 times as long as wide with base and median equal in
width; hosts – C. suppressalis, S. inferens, C. medinalis, P. mathias, Telicota augias . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthopimpla flavolineata Cameron
564' Areola receives costulae near center, 1.05-1.45 times as long as wide, midapical
transverse carina nearly a median longitudinal carina length below level of spiracle (Fig.
1044); ovipositor sheath 3.9 times as long as wide, base wider than middle; host — S.
inferens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthopimpla enderleini Krieger
565(563') Propodeum without black spots; areola with apically wide and basally narrow margin
(Fig. 1045); mesoscutum with a broad transverse trilobed black band above median;
apicomedian band with a yellow V-shaped cleft; tergite I and apical two tergites without
paired black spots (Fig. 1046); hosts — C. suppressalis, O. nubilalis . . . . . . . . . . . . . . . . .
.............................. ........... Xanthopimpla modesta modesta Smith
565' Propodeum (Fig. 1047) and mesoscutum with black spots ................... 566
566(565') Abdominal tergites I, III, V and VII with paired black spots (Fig. 1048); areola (Fig.
1047) relatively small, 2.4 times broader posteriorly than apex, apical part narrowed;
midtransverse carina almost midway level of spiracle; mesoscutum with a transverse
black band in the subapical one-half and more or less trilobe (Fig. 1048); tegula not
black; ovipositor sheath 1.8 times as long as hindtibia, its tip strongly convex above
without transverse ridges; host — C. suppressalis, P. guttata, O. nubilalis, C. sac-
chanphagus . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . Xanthopimpla punctata (Fabricius)
566' Tergites I to VII except VI with paired black spots ......................... 567
567(566') Mesoscutum with a pair of broad latero-median and smaller postero-median black
spots; tegulae without black spot; spots on tergite I outside longitudinal carina (Fig.
1049); 1st lateral area of propodeum (Fig. 1050) with an oblique ovoid black spot; hosts
- S. nivella, C. suppressalis, O. nubilalis, C. infuscatellus, C. sacchariphagus, C.
auricilius, S. inferens . . . . . . . . . . . . . . . . . . . . . . Xanthopimpla stemmator (Thunberg)
567' Mesoscutum with a black broadly hooked, claw-like lateral spot and a median lon-
gitudinal cup-like band, longitudinal above constricted median and broadly transverse
below constriction; tegulae black posteriorly (Fig. 1051); 1st lateral area of propodeum
with a black subquadrate spot (Fig. 1052); spots on tergite I half inward the longitudinal
carina; hosts — C. suppressalis, S. nivella . . . . . . . . Xanthopimpla pedator (Fabricius)
568(562') Eyes with a strong emargination opposite antennae (Fig. 1053); notauli indistinct,
mesopleural sulcus without median angulation; tergites I-IV or I-V orange-red; flagella
and pedicel of antennae reddish brown; mid- and hind coxa and hind femora red,
femora apically black, trochanters, hind tibia and hind tarsus yellowish white with bases
and apices of tibia and apices of tarsal segments blackish brown; hosts — C. suppressalis,
C. auricilius, S. incertulas, S. innotata, S. inferens, Naranga aenescens . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Itoplectis narangae (Ashmead)
568' Eyes with a weak emargination opposite antennae; notauli distinct; mesopleural sulcus
distinctly angulated medially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 569
569(568') Hindtarsal segment V longer than II; ovipositor stout, its ventral valve with vertical
ridges very near apex (Fig. 1054); basal flagellar segments and pedicel of antennae
somewhat brownish beneath; all trochanters yellowish white; host — C. suppressalis . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scambus annulitarsis (Ashmead)
569' Hindtarsal segment V shorter than II; ovipositor slender (Fig. 1055); its ventral valve
with strongly slanted ridges subapically; basal flagellar segments and pedicel yellow
underneath; coxa, trochanter, and hind femur of female reddish brown, fore and
midcoxa and all trochanters yellowish white in male; host — C. suppressalis . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gregopimpla kuwanae (Viereck)
570(558') Face occupied largely by a yellow flat to concave escutcheon-shaped plate bounded by
a carina on all sides, and a yellow thin apico-median tubercle between antennal bases
projected towards vertex or ocellar area; a broad yellow patch between antennal area
and emarginate area of eyes; scutellum brown in apical half and yellow posteriorly,
parallel-sided with small tubercles postero-laterally; wing with a brown infuscation in
the 3rd cubital part or radial cell (Fig. 1056); abdomen with black and yellow bands (Fig.
1057); coxae I and II yellow, III black to reddish brown; hosts — S. litura, H. armigera .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metopius rufus browni Ashmead
570' Face and abdomen not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571
571(570') Ovipositor not or slightly exserted behind apex of abdomen; sternaulus absent or weak;
abdomen with or without apico-median spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 572
571' Ovipositor conspicuously exserted behind apex of abdomen; sternaulus present, at least
about one half as long as mesopleuron; abdomen with an apico-median spot . . . . . . . . .
........................................................................ 573
572(571) Clypeus with a pair of apico-median teeth; head globular in front; malar space short
(Fig. 1058); spiracle of propodeum subglobular; scutellum not carinate laterally; ab-
domen without an apico-median white spot; flagella of female antenna subcylindrical
and apically broadened; thorax and tergites I-IV red; mid- and hind coxa including
trochanter and base of femur reddish brown; hind femur blue-black except basal part;
hind tibia and tarsus blackish brown, segments pale at base; host — C. suppressalis . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centeterus alternecoloratus Cushman
572' Clypeus without apico-median teeth; head transverse in front; malar space very long
(Fig. 1059); spiracle of propodeum elliptical; scutellum carinate laterally; flagella of
female antennae strongly flattened and broadened posteromedially; head, thorax,
propodeum, tergites, and legs yellowish brown to reddish brown; abdomen with an
apicomedian white spot; apical II or III tergites, apices of hindfemora and tibiae
broadly black including narrow part of basal hind tibia and apical one-third-half of
hindtarsal segment I; host — S. incertulas . . . . . . . . . Ischnojoppa luteator (Fabricius)
573(571') Areolet of forewings large; thorax including propodeum without red marks . . . . . 574
573' Areolet of forewing small; thorax at least behind the level of scutellum and propodeum
conspicuously reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 576
574(573) Mesoscutum polished; with strong punctations; tergite black, (not rufous), in male,
basal half of tergites I and II (sometimes III too) white; hind tibia white basally flagella
of female antennae reddish basally beneath; frons of male with a small lateral white spot;
forewing of female clouded apically, crossveins of areolet almost parallel to each other
and faint similar to 2nd recurrent vein (Fig. 1060); host - C. suppressalis . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrothereutes lanceolatus (Walker)
574' Mesoscutum mat, with indistinct punctures; hind tibia not white basally . . . . . . . 575
575(574') Scutellum mat, with dense punctures; tergite II and III of females mat and very densely
punctured; tergites usually black (Fig. 1061); hindlegs black, tarsus of male white
medially; female flagella black with median white rings (Fig. 1062); host - C. suppres-
salis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gambrus wadai (Uchida)
575' Scutellum polished or subpolished with few indistinct punctures; tergites II and III of
female polished to subpolished with weak sparse punctures, II-III or I-IV red; hindlegs,
tibiae, and tarsi slightly paler with apices of femora and tibiae black, and apices of tarsal
segments slightly darkened; flagella of female antennae tricolored, red basally, white
medially, and blackish apically (Fig. 1063); host - C. suppressalis . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Gambrus ruficoxatus (Sonan)
576(573') Frons without a semicircular carina above each antennal socket; mandibles broad
apically with teeth subequal in length; ovipositor thicker and tip usually blunt; upper
margin of pronotum moderately swollen; propodeal apophyses or sublateral crests
present as teeth in females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 577
576' Frons with a semicircular carina above each antennal socket . . . . . . . . . . . . . . . . . 580
577(576) Mesoscutum dull, striate or aciculate along margins and grooves; scutellum flat to
weakly arched, dull or subpolished, distinctly and closely punctate; lateral carina of
scutellum extending to beyond its basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 578
577' Mesoscutum smooth or subpolished; scutellum smooth and convex; lateral carina not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579
578(577) Subtegular ridge and tegulae always entirely black; scutellum red or black; pronotum
coarsely trans-striate medially and irregularly wrinkled above; mesoscutum finely acicu-
late along margins of each lobe, median of midlobe rugulose and lateral lobes sub-
polished; anterior half of thorax black and posterior half light colored (Fig. 1064);
forewing with a fuscous band extending from base of stigma to lower margin; middle,
hind trochanters, coxa, and femur never black, usually reddish (Fig. 1065); female
abdomen (Fig. 1066) with apices of tergites I and II, entire VII, and narrow apex of VII
whitish yellow, hosts - S. nivella, C. sacchariphagus, Bactra venosata . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goryphus basilaris Holmgren
578' Subtegular ridge, tegulae, and scutellum whitish yellow; female forewing without a
brown tinge below stigma (Fig, 1067); face finely longitudinally striate; upper margin of
pronotum with fine oblique striae; mat mesoscutum finely punctured; tergites I and II
with white apices, VII and VIII broadly yellowish white, II to V alternately blackish to
reddish brown (Fig. 1068); host — C. suppressalis . . . . . Goryphus apicalis Holmgren
579(577') Forewings (Fig. 1069) with a small brown spot below stigma; scutellum yellow; first
tergite slender, apical part slightly widened; areolet pentagonal with side parallel; apices
of tergite I, II and VII broadly white, rest of I reddish brown like propodeum (Fig. 1070);
antennae dark brown without white marks of flagellar segments V-IX; forelegs pale
yellow, middle leg pale brown with apical tarsal segments dark brown; hindlegs reddish
brown except trochanter, 1st, 2nd and 5th hind tarsal segments dark brown and rest of
hind tarsus white (Fig. 1071); hosts — Oulema melanopa, O. oryzae in Taiwan . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goryphus lemae (Sonan)
579' Forewings with 2 brownish bands; hindwing apices dark brown (Fig. 1072); scutellum
reddish brown along apical margins; first tergite not slender and dark orange; apices of
tergites I to IV and VI1 yellow similar to fore, middle coxae, and trochanters;
propodeum and rest of tergite I dark orange; legs largely dark orange, all tarsal
segments, hind trochanter, apex of femur, base and apex of tibia blackish (Fig. 1073);
antenna black with white marks above each of flagellar segments V-IX; host — Scir-
pophaga incertulas . . . . . . . . . . . . . . . . . . Goryphus mesoxanthus mesoxanthus (Brulle)
580(576') Forewing with a broad band below stigma covering distal one-third of discocubital cell,
inner basal part of radial cell, almost entire areolet, distal two-third of 2nd discoidal cell
and inner apical part of 2nd brachial cell; wing apices along distal ends of 3rd cubital,
3rd discoidal and 2nd brachial cells with brown tinge (Fig. 1074); antennae dark brown
and thin mesonotum with fine punctations; palpi white and tarsi yellowish; hosts —
Chilo infuscatellus, Bactra venosata in East Asia . . . . . . . . . Isotima chilonis (Uchida)
580' Forewing tinge different from above; tergite I black or red; apical transverse carina of
propodeum absent or weakly indicated laterally; mesoscutum black; apical margin of
tergite II of female and apical margins of I-III of male white; forecoxa and trochanters
of female fuscous; hind trochanters, tibiae, and tarsi black; hosts — Scirpophaga in-
notata, s. incertulas, Sesamia inferens in South and Southeast Asia . . . . . . . . . . . . 581
581(580') Tergite I black (Fig. 1075); hosts — S. innotata, S. incertulas in Java (Indonesia) and
Philippines, and Sesamia inferens in Malaysia . . . . . Isotima dammermani (Rohwer)
581' Tergite I red (Fig. 1076); host — S. incertulas in India and Indonesia . . . . . . . . . . . . . . .
.................................................... Isotima javensis (Rohwer)
582(557') Discocubital cell of forewing (Fig. 1077) with a conspicuous hairless area bearing two
scleromes (a sclerotized streak and spot); mesoscutum light brown; apical tergites light
brown, not marked with black; only one intercubital vein present and second recurrent
vein conspicuously basad of the intercubital vein; host – S. inferens, Taiwanese and
Japanese species . . . . . . . . . . . . . . . . . Enicospilus sakaguchii (Matsumura & Uchida)
582' Forewings without such structure or appearance . . . . . . . . . . . . . . . . . . . . . . . . . . . 583
583(582') Areolet present on forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 584
583' Areolet absent on forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 587

584(583) Eyes rather strongly emarginate; temple narrow; tegula black; head subquadrate; inter-
ocellar distance 1.1-1.2X the ocello-ocular distance; thorax longer with speculum shiny;
lower outer angle of 2nd discoidal cell acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 585
584' Eyes weakly or not all emarginate; temple moderately wide; clypeal margin present or
indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 586
585(584) Scape and pedicel yellow (Fig. 1078); mesopleuron below subtegular ridge finely
transtriate; foreleg entirely yellowish brown with black coxa; III brownish black with
black coxae (Fig. 1079); propodeum flattened and abruptly narrowed apically, median
propodeal carinae indistinct; host – Pamara bada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Casinaria ajanta Gupta & Maheshwary
585' Scape and pedicel black (Fig. 1080); mesopleuron below subtegular ridge irregularly
and coarsely wrinkled; median carina wavy and present; fore trochanters and base of
forefemur blackish brown usually up to middle, tarsal segments largely black; midtibia
blackish apically and tarsi wholly black; leg III uniformly black (Fig. 1081); propodeum
large and elongate; tergite I long and straight; postpetiole bulbous; hosts – Pelopidas
mathias, P. cinnara . . . . . . . . . . . . . . . . Casinaria pedunculata pedunculata (Szepligeti)
586(584') Tergite I of abdomen without baso-lateral pit or groove in front of spiracle; propodeum
extending backward over middle of hindscoxa; male clasper with a preapical dorsal
notch; tergites almost entirely black; mandibles, scape and pedicel yellow; midcoxa and
hindcoxa black; hindfemur, tibia and tarsus blackish brown; trochanter yellow; face
black, clypeus confluent with the face (Fig. 1082); host – C. suppressalis . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Venturia sp.
586' Tergite I of abdomen with a baso-lateral pit or groove in front of spiracle; tergite II-V
red with bases of II and III blackish; mid- and hindcoxae, hindfemur, and hindtibia red,
tibia slightly darkened at apex; trochanters light brown; hindtarsus reddish brown, each
segment pale basally; areolet of forewing present (2 intercubital veins present) (Fig.
1083); host – S. incertulas . . . . . . . . . . . . . . . . . . . . . . . . . Diadegma akoensis (Shiraki)
587(583') Clypeus separated from face by a groove . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 588
587' Clypeus confluent with face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 591
588(587) Ventral margins of tergite I subparallel to each other (Fig. 1084); occipital carina almost
complete above; lateral ocellus distance from eye about 1.0 (in female) or 0.5 (in male)
its diameter; mesoscutum light brown and yellow, most often with 3 blackish stripes
(sometimes with one or with no black stripes ); scutellum yellow; hindtibia darkened on
both ends; hosts C. suppressalis, Scirpophaga spp., C. medinalis, Marasmia spp. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trathala flavoorbitalis (Cameron)
588' Ventral margins of tergite I bowed inward to touch or almost touch each other; occipital
carina broadly incomplete above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 589
589(588') Second discoidal cell of forewing pointed at base (Fig. 1085); hindfemur about 3.1 times
as long as deep; male clasper with an apical small peculiar projection; mesoscutum and
scutellum entirely black; tergites III- VI alternately black and reddish brown (Fig. 1086);
hindtibia blackish brown; hosts - Scirpophaga spp., C. medinalis, Marasmia spp. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temelucha stangli (Ashmead)
589' Second discoidal cell of forewing truncate at base (Fig. 1087); hindfemur about 4.3-5.2
times as long as deep; male clasper without an apical projection . . . . . . . . . . . . . . 590
590(589') Lateral ocellus very close to eye, separated by about 0.5 (in female) or 0.1-0.2 (in males)
its diameter (Fig. 1088); mesoscutum light brown and yellow, usually with three dark
brown stripes; scutellum yellow; hindtibia scarcely darkened at apex; hosts - C. sup-
pressalis, Naranga diffusa, Bradina admixtalis, C. medinalis . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temelucha biguttula (Munakata)
590' Lateral ocellus rather far from eye, separated by about 1.5 (in female) or 1.0-1.1 (in
male) its diameter (Fig. 1089); mesoscutum, scutellum, and hindtibia colored as in T.
biguttula; general color yellowish orange; hosts - C. suppressalis, C. medinalis, Maras-
mia spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temelucha philippinensis (Ashmead)
591(587') Eyes with conspicuous long hairs, strongly emarginate and, distinctly convergent
ventrally; wings without discoidella; mesopleuron without smooth shiny area, with
horizontal striae; face yellow including temple, but with frons and ocellar areablack; top
of propleuron, subtegular area, scutellum, coxae I-III and legs (except leg IV brownish
black with apical two-thirds of tibiae, entire trochanter and basal one-third of femora
blackish brown) all yellow; scutellum rather convex with two longitudinal carinae
towards posterior half; tergite I without lateral groove; tergite II without a thyridium;
ovipositor about 2-3.8 as long as apical length of abdomen (Fig. 1090); hosts
Cnaphalocrocis medinalis, Marasmia spp., Ostrinia jumacalis . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichomma (Trichomella) cnaphalocrosis Uchida
591' Eyes bare, weakly or strongly emarginate and not convergent ventrally; lateral groove in
tergite I and thyridium in tergite II present; face, subtegular area, and scutellum black
....................................................................... 592
592(591') Eyes strongly emarginate; area of vertex behind eyes narrow; mesopleuron without
smooth area, coarsely punctate; scutellum flat, somewhat deep medially; lower outer
angle of second discoidal cell forms a right angle; tip of mediellan cell well rounded with
a short protruberance apically; radiella concave or rounded (as seen on top) . . . 593
592' Eyes weakly emarginate or not at all; area of vertex behind eyes moderately wide;
scutellum convex; tip of mediellan cell not well rounded, radiella straight and cubitella
deeply concave before apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 595
593(592) Median propodeal carinae distinct and usually very strong; notaular region similarly
sculptured as the rest of mesoscutum; mesopleuron finely and closely rugose; tegulae
black; face rugulose; male genital claspers broad and flat apically (Fig. 1091); subgenital
plate (Fig. 1092) of male oblo-elongate, more or less uniformly colored; host – Achaea
janata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charops ganges Cushman
593' Median propodeal carinae absent or weakly present in the apical region of the
propodeum (Fig. 1093); middle femora wholly yellow . . . . . . . . . . . . . . . . . . . . . . . 594
594(593') Leg III black except yellow trochanter, base of tibia, and ends of femur (Fig. 1094);
tarsal claw with 5 teeth (Fig. 1095); black mesopleuron reticulate rugose; forecoxa and
tegulae yellow; abdomen largely reddish brown; male genital clasper with a slender
apical process (Fig. 1096); subgenital plate (Fig. 1097) of male broadly subglobular to
ovoid with a tapering point tubercle-like, plate dark colored except lateral periphery
and tubercle light colored; hosts – S. innotata, C. medinalis, Marasmia spp. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charops brachypterum (Cameron)
594' Hindfemur yellowish brown; base of tarsus I yellow (Fig. 1098); petiole wholly reddish;
mesopleuron closely rugose; male genital clasper with a moderately slender apical
process, midlength of process a little broader (Fig. 1099); subgenital plate (Fig. 1100)
almost wholly dark colored except light colored margins; subtriangular with slightly
convex top; hosts – S. incertulas, S. innotata, Naranga aenescens, Spodoptera mauritia,
Mythimna loreyi, Pelopidas mathias . . . . . . . . . . . . . . . . . . . Charops bicolor (Szepligeti)
595(592') Abdomen reddish brown except black tergites I & II (although apices are reddish brown
usually); trochanter III yellow; apical half of basal tarsal segment of leg III dark; thorax
and head black; wings with 2nd recurrent vein erased at midhalf (Fig. 1101) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriborus vulgaris (Morley)
595 ' Entire abdomen black; tarsal claws with small pectines basally; hind tarsal segment III
about as long as segment V (Fig. 1102); scape black with brownish yellow tinge beneath;
mid- and hind coxa black; hind femur red; hind tibia and tarsi reddish brown to
yellowish brown, tibiae blackish at apex and tarsal segments I-IV apically and entire of
segment V slightly darkened; trochanters yellowish brown to reddish brown with hind
trochanter blackish basally; hosts – C. suppressalis, Sesamia inferens . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriborus sinicus (Holmgren)
595" Abdomen uniformly black dorsally; tarsal claws entirely pectined from base to apex;
hindtarsal segment III slightly longer than segment V (Fig. 1103); coloration as in E.
sinicus; hosts – S. inferens, S. incertulas . . . . . . . . . . Eriborus terebrans (Gravenhorst)
596(520') Forewings with 2-3 closed cubital cells and a single recurrent vein attached to the base
of second cubital cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 597
Figs. 1089-1098
596' Forewings with two closed cubital cells and two recurrent (1st and 2nd) veins attached
to the base of the second cubital cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 598
597(596) Scutellum with a squarish yellowish orange band; metascutellum with a transverse
rectangular yellowish orange mark; head, thorax, legs and abdomen clothed with black
hairs; clypeus punctated but light medially, convex, pointed apically and without
transverse ridges; mandibles without grooves and serrated with 2 inner teeth; antennal
base very close to eyes with a distance of 0.25 diameter of antennal base (Fig. 1104);
wings with bluish lustre basally and reddish brown towards apex; l st recurrent vein in
the basal one-fourth of 2nd cubital cell (Fig. 1105) 36 mm long . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scolia nr. scutellaris Gribodo
597' Scutellum and metascutellum black; head with yellowish white hairs arising from each
punctate; clypeus not convex, mostly smooth without punctures except laterals below
antennal base; anterior margin with transverse ridges and light longitudinal carinae
medially above it; mandibles without serrated teeth; antennal base distant from eyes;
female tergites I-IV (Fig. 1106) with a thin yellow band and a row of yellow hairs at
apices, rest black; median yellow band narrow but broadens laterally in tergite I, II with
an additional pair of subglobular yellow spots toward margins; apices of sternites with
yellowish white hairs and yellow patch along apico-laterals of II and III; male tergites
I-V with yellow narrow apical bands; bands in I-III widen laterally and become spot-like;
apices of sternites II-III with yellow bands, erased medially in IV and reduced to spot
laterally in V (Fig. 1107); male 100 mm long and female 14 mm long . . . . . Scolia sp.
598(596') Abdomen uniformly black, apices of tergites with black hairs; head with black or whitish
and reddish orange hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 599
598' Abdomen black with tergite apices lined with yellowish hairs and bands behind it or
white hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 600
599(598) Hairs on head and pronotum black; vertex punctated above lateral ocelli; eye margins
and entire clypeus well punctated; with deep pits in the entire area between eye
emargination (Fig. 1108); distance between first and second recurrent vein 1.3 times that
space between first and second transverse cubital; 2nd transverse cubital oblique in the
apical four-fifths and vertical in the basal one-fifth; bulge of cell R in line with 2nd
transverse cubital; extension of Cu as long as the distance between two recurrent veins;
(Fig. 1109); radiella far away from cubitella and their tips diverge apically, venation
reaches midhalf of hindwing (Fig. 1110); 27 mm long . . . . . . . . . . . Campsomeris sp. A
599' Hairs on back of head and entire pronotum reddish orange; vertex smooth; shiny,
impunctate including ocellar area, seven small transverse pits with white hairs above
lateral ocelli (Fig. 1111); eye margins hardly punctate; midclypeus smooth and im-
punctate; distance between two recurrent veins 0.7 of distance between transverse
cubital 1 & 2; midlength of transverse cubital slightly pointed outward; bulge of cell R
beyond level of 2nd transverse cubital; extension of Cu long, two-fifths longer than
distance between two recurrent veins; radiella and cubitella parallel-sided and parallel
to each other; 17 mm long . . . . . . . . . . Campsomeris collaris quadrifasciata Fabricius
600(598') Apices of abdominal tergites II to IV with white hairs; tibial spur of leg III white . . . . .
....................................................................... 601
600' Apices of abdominal tergite I-V with yellow bands and hairs; tibial spurs white to
blackish brown ........................................................ 602
601(600) Apical distance between first and second recurrent veins longer than the apical distance
between transverse cubital 1 and 2; first recurrent vein with an angled corner; subdis-
coideus straight; vein R rounded at its bulging point with its tip directed slightly
downwards below level of wing apex; tip of Cu beyond second cubital cell short, barely
one-fifth the apical distance of two recurrent veins (Fig. 1112); radiella and cubitella of
hindwings not traceable (Fig. 1113); mandibles narrow apically with short inner basal
hairs; apex of hindfemur with a reddish brown enlargement in the inner side, central
plate lined with subglobular marks (Fig. 1114); basal tarsal segment of leg III with 2 long
white subapical ventral setae (Fig. 1115) . . . . . . . . . . . . . . . . . . . . . Campsomeris sp. B
601' Apical distance between 2 recurrent veins as long as apical distance between transverse
cubital 1 & 2; first recurrent vein with a rounded corner; subdiscoideus downwardly
curved apically; bulging point of vein R well beyond slightly S-shaped 2nd transverse
cubital and tip directed towards wing apex; extension of Cu beyond 2nd cubital cell,
one-fifth longer than the distance between two recurrent veins (Fig. 1116); radiella and
cubitella prominent (Fig. 1117); mandibles broad and tip moderately acute; abdomen
entirely black similar to Campsomeris sp. B, lined along apices of tergites with white
hairs (Fig. 1118); brown plate at inner-apical part of hind femur (Fig. 1119) without
globular marks; basal tarsal segment of leg III with five long white setae . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsomeris sp. C
602(600') Hairs on head, thorax, legs and apices of tergites yellow to dark yellow or brownish
yellow; clypeus, scutellum, and metascutellum without yellow bands; mandibles
prominently open, tips touching each other; yellow band on apex of tergite II straight
basally, without U- or V-shaped pattern; tergite III with a hat-like yellow mark apically,
elevated medially and low laterally (viewed posteriorly) (Fig. 1120); apical distance
between 2 recurrent veins as long as apical distance between 2 transverse cubitals (Fig.
1121) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsomeris sp. D
602' Hairs on body and legs white to whitish yellow; clypeus and scutellum with yellow areas;
mandibles on top of each other and closed; yellow bands on tergite II-V all U-shaped
..................................................................... 603
603(602') Tergite II with a wide V-shape mark; III concave medially, yellow band much wider than
black band (Fig. 1122); scutellum with a pair of yellow subrectangular spots; metascutel-
lum without yellow marks; clypeus with small black median spot; distance between
antennal base and eyes 0.36 diameter of antennal base . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsomeris nr. aurulenta (Smith)
603' Tergite II with a wide U-shape band; I with an additional black band on top of yellow
band; III widely concave medially, black band much wider than yellow band (Fig. 1123);
scutellum and metascutellum with a broad transverse yellow band; clypeus with a large
median black spot; distance between antennal base and eyes 0.2 diameter of antennal
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsomeris annulata (Fabricius)
604(523') All tibiae with 2 spurs, reduced to a calcar and a simple spur in the fore tibia; wing
simple with well sclerotized long marginal vein extended to wing base; stigmal vein
moderately long and concave; antennae 11-12 segmented [Ceraphronidae] . . . . . .605
604' Mid and hind tibia with only 1 spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607
605(604) Forewing with 2 broad brown bands, dark at midhalf and light at apex (Fig. 1124);
thorax, legs and scape yellow; head brownish yellow; abdomen brownish black similar
to funicular segments VI-VIII (Fig. 1125); 0.9-1 mm long . . . . . . . . . .Ceraphron sp. A
605' Forewing without brown bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 606
606(605') Brown except black head, propodeum, flagella of antenna, pedicel and apical half of
scape (Fig. 1126), and apical one-half of abdomen; all legs brownish yellow; forewing a
little longer than abdomen (6057) and with 2 light yellow brown bands running parallel
with the marginal vein, in between with a row of setae (Fig. 1127); funicle segment I (Fig.
1126) as long as combined length of II and III; 2-3 mm long . . . . . . . . Ceraphron sp. B
606' Entirely black except abdomen with brownish tinge; antennal scape yellow, pedicel and
funicles I-IV brownish yellow, funicle I shorter than combined length of II and III, rest
of segments blackish brown (Fig. 1128); legs I-III yellow; forewing without yellow-brown
band parallel with the marginal vein and one-fourth longer than abdomen, stigmal vein
faint and short (Fig. 1129); 1.2 mm long . . . . . . . . . . . Aphanogmus fijiensis (Ferriere)
606' Similar to A. fijiensis; 1st funicle nearly as long as combined length of II and III, terminal
4 segments blackish brown (Fig. 1130); stigmal vein moderately long and with few hairs
near wing base (Fig. 1131) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphanogmus sp.
607(604') Venation absent or reduced to submarginal vein with a slightly knobbed apex; egg-larval
parasites of Orseolia oryzae [Platygasteridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 608
607' Forewing with distinct, though sometimes short marginal and stigmal veins
[Scelionidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 609
608(607) Antennal scape and legs light orange-yellow; apical four antennal segments pale brown,
broad, and subquadrate, basal four smaller in size (length and width) than apical
segments and light orange-yellow; coxae yellow except brown coxa I; lateral posterior
part of thorax with dense greyish pubescence (Fig. 1132); 0.9-1.0 mm long . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platygaster oryzae Cameron
608' Antennal scape and legs yellow; funicular segments brownish black, long and slender;
slightly more than twice as long as broad; coxae brownish black except yellow coxa III;
posterior part of thorax with greyish white pubescence; body length more than 1 mm
(Fig. 1133) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platygaster foersteri (Gahan)
609(607') Gaster with lateral margins rounded; wide latero-tergites of metasoma loosely attached
to sternite (not incised) forming no depressed submarginal ridge; 2nd tergite (T2) of
metasoma the longest; antenna 12-segmented in male and 10-11 in the female
[Telenominae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 610
609' Gaster with lateral margins carinated; narrow laterotergites incised to form impressed
submarginal ridge; metasoma usually subequally segmented with 3rd segment (T3) often
the longest; lateral ocelli usually nearer to the inner orbits than to median cells
[Scelioninae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 633
610(609) Frons predominantly sculptured particularly in the lower half; eyes bare and usually
glabrous; parapsidal furrows often present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 611
610' Frons smooth and shining, lack sculpturing in the lower half of frons; eyes usually hairy;
parapsidal furrows absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 616
611(610) Fan-like carina prominent in the head emanating near mandibular base, extended
dorsad to the frons and cheeks, and branches ventrad on either sides of insertions;
central keel, submedian and orbital carinae developed (Fig. 1134); lateral side of thorax
strongly pitted (Fig. 1135); acetabular field and anteroventral portion of mesepisternum
setose; lateral sulci on 2nd sternite (S2) broadly separated posteriorly but more nearly
straight and setigerous punctures widely separated; antennae with yellow scape (similar
to coxae), black radicle; hosts – Scotinophara latiuscula, S. coarctata . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psix lacunatus Johnson & Masner
611' Fan-like carina absent; cheeks and frons without ridges; postmarginalis of wings usually
longer than stigmalis; metasoma uniformly black; scutellum of females with semi-
decumbent hairs not interfering the sculpture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 612
612(611') Tergite I of abdomen without any sublateral setae; notauli present . . . . . . . . . . . . . 613
612' Tergite I of abdomen with sublateral setae; notauli absent . . . . . . . . . . . . . . . . . . . . 614
613(612) Scutellum coarsely sculptured; slightly more roughly punctured than mesoscutum and
scrobiculate towards edges; broad, well convex mesoscutum strongly reticulate – sculp-
tured and clothed with short hairs; notauli sharply defined posteriorly; gaster longer
than broad, 1st tergum with deep longitudinal costate on basal half, 2nd tergum 1.4 times
wider than long and midbasal two-thirds longitudinally striated, the rest of tergites
smooth and shining (Fig. 1136); female antenna (Fig. 1137) reddish yellow, 5-segmented
club broad and long with 2nd and 3rd club segments transverse, and conical apical
segment; scape subequal to next three segments; length 1.3-1.5 mm; egg parasites of
Nezara viridula L., N. antennata and Piezodorus rubrofasciatus in E. Asia . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trissolcus mitsukurii (Ashmead)
613' Scutellum smooth without microsculpture; mesoscutum covered with leather-tough
microsculpture; notauli (Fig. 1138) narrow but visible; dorsellum with dorsal transverse
row of deep pits along scutellar-metanotal suture and venter with longitudinal carinae;
episternal foveae well-developed with a line of 4-5 dorsoventrally elongate deep pits
running from dorsal tip of acetabular carina toward mesopleural pit; mesopleural carina
absent ventrally (Fig. 1138); tergum II with weak wrinkles reaching well beyond basal
costae and short irregular subapical line of setae; antenna dark brown to black with light
yellow-brown radicle and scape; malar region narrow; length 0.9-1.2 mm; egg parasite
of Nezara viridula in N. America . . . . . . . . . . . . . . . . . . . . Trissolcus thyantae Ashmead
614(612') Second tergite with prominent rugulae beyond basal costae; scutellum with distinct
tough leather-like microsculpture; bases of setae with pustules or blisters; head in lateral
view, with wide gena highly bulging posteriorly; vertex broadly rounded; episternal
foveae only a pair of circular pits dorsad of acetabular carina apex (Fig. 1139); 2 nd
tergum highly rugulose eyes hairy and diverging posteriorly (Fig. 1140); hosts – Nezara
viridula, Piezodorus guildinii in Americas . . . . . . . . . . Trissolcus basalis (Wollaston)
614' Second tergite smooth; bases of setae not strongly raised . . . . . . . . . . . . . . . . . . . . 615
615(614') Vertex abruptly rounded up to occiput; orbital furrow dorsally and ventrally con-
stricted; hyperoccipital carina absent; frons with rough thick leather-like microsculp-
ture (Fig. 1141); anteroventral portion of mesepisternum coriaceous, median
longitudinal carina well developed; scutellum smooth (Fig. 1142); episternal foveae 2-3
poorly delimited depressions; host – Nezara viridula in Americas . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trissolcus hullensis (Harrington)
615' Vertex sharply angled up to occiput; orbital furrow narrow and fading ventrally; hyper-
occipital carina present; frons with granulose microsculpture (Fig. 1143); scutellum
coarsely rugose-areolate, antero-ventral mesoepisternum rugulose; episternal foveae
only 2 shallow pits (Fig. 1144); egg parasite of Nezara in N. America . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trissolcus solocis Johnson
616(610') Hosts – stem borer eggs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 617
616' Hosts-eggs of pentatomid bugs in the genera Nezara, Scotinophara, Eysarcoris)
Oebalus and eggs of lepidopterans, Melanitis, Spodoptera and Mythimna . . . . . . . 624
617(616) Small telenomines, body length 0.5-1.1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 618
617' Moderately large telenomines, body length 1.5-1.8 mm . . . . . . . . . . . . . . . . . . . . . . 620
618(617) Tergite I uniformly smooth, unsculptured, its costae absent or reduced to an impercep-
tible row of minute foveae basally; tergite II (Fig. 1145) with emarginate apex; lower
midhalf of abdomen evenly tapering towards apex and somewhat flattened; metasoma
long and slender, clearly concave in dried specimens; antenna of males yellow except
apical 3-4 segments, funicles 4 to 9 subglobular and bead-like, funicles 1 to 3 about twice
as wide as the rest (Fig. 1146); female scape 4.9 times as long as broad, 1st funicle 1.6
times as long as broad, 2nd and 3rd subequal and subquadrate, club 4-segmented with
each segment a little longer than broad and conical-ovate apical segment (Fig. 1147);
penis sheath not narrowed before the apex (Fig. 1148); female 0.8-0.85 mm long and
male small, 0.5-0.6 mm long (Fig. 1149); hosts — Scirpophaga incertulas, S. innotata . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus rowani (Gahan)
618' Tergite I strongly and evenly costate at apical margin; thorax hardly longer than its
greatest width; head markedly transverse; shortest distance between the eyes clearly
greater than their width as seen from above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 619
619(618') Abdomen nearly three times as long as its greatest width (Fig. 1150); hind tibia slightly
less than twice as long as its metatarsus; forecoxa of male prominently darkened,
without conspicuous darkening along inner border of apical appendages; basal segment
of hind tarsus long and slender, equally thin as the rest, and almost as long as the apical
three segments; penis sheath narrowed before apex (Fig. 1151); male 0.9 mm long and
female 1-1.1 mm long; hosts — S. incertulas, S. auriflua and Chilo sacchariphagus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus dignoides Nixon
619' Abdomen about twice as long as wide, narrowed to a sharp point at apex; tergite I with
sharply defined longitudinal costae right across; tergites 3 and 4 strongly transverse
(Fig. 1152); pedicel longer than combined length of funicular segments I and II (Fig.
1153); hind tibia slightly more than twice as long as its metatarsus; legs bright yellow;
genitalia distinct on account of thickening which appears as a dark band along inner side
of apical appendage (Fig. 1154); 0.7-0.8 mm long; hosts — Scirpophaga innotata, S.
incertulas, Chilo suppressalis, C. polychysus, C. auricilius . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus dignus (Gahan)
620(617') Eyes bare; transverse, head, and body dark brown with yellow legs except brown coxa I
and apical tarsomeres; antennal segment II >III, IV to VI weakly rounded and
transverse, VII trapezoidal, VIII-X transverse, XI longer than wide and apically globose
(Fig. 1155); wings narrow without a basal vein but with long marginal setae, stigmal vein
shorter than postmarginal vein; legs short and stout, 5th tarsomeres of all legs dilated,
5th middle tarsomere with four strong ventral spines (Fig. 1156); mesoscutum weakly
and evenly convex; scutellum bulging, smooth with long sparse setae; submarginal
foveae absent; tergite I of abdomen with 2 pairs of lateral setae, 1 pair of sublateral with
basal margins finely punctured; tergite 2 without longitudinal costae but with transverse
striations basally (Fig. 1157); 1.7 mm long; host — S. innotata in S. E. Asia . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus transversus Bin & Johnson
620' Eyes setose; combination of characters not as above . . . . . . . . . . . . . . . . . . . . . . . . 621
621(620') Mesoscutum flat or strongly depressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 622
621' Mesoscutum evenly convex; head more transverse-ovate (Fig. 1158); female antennal
segment II much longer than any of III-XI, VI strongly transverse and subglobular,
VII < VIII, VIII-X transverse (Fig. 1159); in male, II much longer than any of III-XI, IV
and V slightly longer and with basiconic sensillum and wider than III, VI-X transverse;
5th tarsomere of legs without strong setae ventrally; abdomen elongated, tergite II much
longer than wide and tip pointed; tergite I with 3 pairs of lateral setae, 2 pairs of
sublateral setae and costae reduced to a median row of foveae similar to tergite II (Fig.
1158); genitalia with a narrow median plate and small digital teeth (3/digitus), penis
valves strongly sclerotized and sheath tapers apically (Fig. 1160); 1.7 mm long; host –
S. melanoclysta in Africa . . . . . . . . . . . . . . . . . . . . Telenomus versicolor Bin & Johnson
622(621) Head strongly reflexed; lateral angles of clypeal margins sharply pointed; labral margins
excised medially; temples narrow, weakly bulged and with a narrow band of coriaceous
sculpture along posterior orbits reaching dorsal occipital carina (Fig. 1161); mesos-
cutum strongly depressed; scutellum strongly transverse; episternal foveae present but
very shallow; acetabular field large entering mesopleural furrow but not reaching
intercoxal space; antennae of female (Fig. 1162) with segment II and III equal in length,
IV (longest) > V, VI globular to quadrate, VIII > VII, VII to X transverse; segment V
with a prominent tyloid-sensillum and VIII-XI each with a long seta in the expanded
ventro-apical angle in the male (Fig. 1163); tergite I with 2-3 pairs of lateral setae,
wrinkles of tergite II fan-like medially reaching one-third of its length (T2); digital teeth
small with 3 teeth/digitus; penis sheath strongly rounded at apex (Fig. 1164); 1.8 mm
long; hosts – Diatraea rufescens and D. saccharalis in South America . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus alecto (Crawford)
622' Head not strongly reflexed; occipital carina complete with or without median interrup-
tions; temples with or without a band of coriaceous sculpture . . . . . . . . . . . . . . . . . 623
623(622') Temples with a band of coriaceous sculpture along posterior orbit reaching half the
distance from occipital carina; head moderately transverse (Fig. 1165); occipital carina
complete without median interruptions; acetabular field reaching mesopleural furrow;
female (Fig. 1166) antennal segment V and VI globular, male (Fig. 1167) with segment
III > II, IV = V, V with a tyloid sensillum ventrally, VI-XI clearly transverse; tergite I
with 2 pairs of lateral setae and costae in basal half, tergite II nearly as broad as long
(Fig. 1165); genitalia with small digital teeth, median plate forms two indistinct converg-
ing rods, penis sheath narrow and pointed apically (Fig. 1168); 1.5 mm long; host –
Chilo sacchariphagus in S. Asia . . . . . . . . . . . . . . . Telenomus globosus Bin & Johnson
623' Temples without such band; head subglobular, occipital carina indistinct medially (Fig.
1169); mesoscutum strongly depressed (Fig. 1170); acetabular field not reaching
mesopleural furrow; antenna in female with segment, VI strongly transverse (Fig. 1171);
male (Fig. 1172) with segment II=III, IV< V, V with a distinct basiconic sensillum,
VI-XI globose; abdomen subelongate, broadens below lower end of broad tergite II,
tergite I with 2 pairs of lateral setae and costate in entire length of laterals and basal half
of middle area (Fig. 1170); genitalia with large digital teeth; median plate forms an
elongate trapezoidal plate; penis sheath short and strongly long rounded to moderately
tapered apically (Fig. 1173); 1.6 mm long; hosts — Maliarpha separatella and Eldana
saccharina in Africa . . . . . . . . . . . . . . . . . . . . . . . Telenomus applanatus Bin & Johnson
624(616') Egg parasitoids of noctuids (Mythimna and Spodoptera) and satyrid (Melanitis); first
funicular segment as long as or slightly longer than pedicel . . . . . . . . . . . . . . . . . . 625
624' Egg parasitoids of pentatomid bugs (Nezara, Piezodorus, Scotinophara, Eysarcoris,
Oebalus); first funicular variable, in some, as long as pedicel . . . . . . . . . . . . . . . . . 627
625(624) Head and pronotum of male reddish yellow, antenna deep brown except yellowish
brown scape, leg yellow except fusco-testaceous femur and tibia of leg III and apical
tarsal segments yellow or black; female, black with chestnut brown antenna and fuscous
legs; head transverse with well convex frontal region, broad and flat vertex; male
antennal segment I 3.2 times as long as II, II a little longer than III, III = IV; VI-VI11
subquadrate, IX = X, XI < X, XII conically elongate; female club 4-segmented;
segment VI very short and strongly transverse, proximal three segments slightly
transverse (Fig. 1174); forewings rather broad and hyaline, ciliated, stigmalis one-fifth
of postmarginalis; tergite I of abdomen transverse with longitudinal costae all over; II
nearly as long as broad with short striae basally; 0.6-0.7 mm long; egg parasitoid of
armyworm Mythimna unipuncta in E. Asia . . . . . . . . . . . . . Telenomus cirphivorus Liu
625' Not as above, egg parasitoids of different hosts . . . . . . . . . . . . . . . . . . . . . . . . . . . . 626
626(625') Large species with short and stout body, at least 0.9 mm long; head transverse, vertex
convex, finely and evenly scaly-reticulate with sparse short hairs; preoccipital carina
present and distinct even in the median area; antenna honey yellow except dark brown
club, pedicel as long as first funicular segment; VI subglobular, VII and proximal three
segments of club transverse (Fig. 1175); forewing pale, venation yellowish and not
sharply outstanding; stigmalis rather long 0.6 times of postmarginalis; fringes of hind
wing a little more than half maximum width of wing (Fig. 1176); hosts — unknown
noctuid moth and Melanitis leda in Asia . . . . . . . . . . . . . . . . Telenomus Iucullus Nixon
626' Very small species, length 0.5 mm; head strongly transverse; vertex without a posterior
declivous surface; preoccipital carina not distinct (Fig. 1177); antenna uniformly black-
ish, funicles I-IV very small, bead-like and much narrower than pedicel and VI-VI11
similar in size, female segment III distinctly longer than IV, VI strongly transverse (Fig.
1178); male with segment III-V of same length (Fig. 1179); forewing greyish; stigmalis
short; fringes of hindwing longer than or fully as long as maximum wing width; genitalia
slightly narrow medially, with stout digiti and digital teeth, 3/digitus, median plate very
wide basally and strongly concave at apex near digiti, penis sheath moderately long,
subrounded at apex; genitalia in general short and broad (Fig. 1180); hosts — Spodop-
tera litura and S. mauritia acronyctoides in Asia . . . . . . . . . . . . Telenomus remus Nixon
627(624') Oriental species of Telenomus parasitizing Nezara or Scotinophara . . . . . . . . . . . 628
627' Non-oriental (Palearctic, Nearctic and Neo-tropical) species of Telenomus attacking a

wider genera of pentatomid bugs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629
628(627) Small species, 0.8 mm long; postscutellum with a semi-triangular strongly rugose median
dilation; ocellar region without punctures (Fig. 1181); 1st funicular segment shorter than
pedicel, 3rd at least not elongate; club clearly 5-segmented (Fig. 1182); stigmalis rather
short; fringe about two-thirds maximum wing width; abdomen longer than wide (11:7)
with tergite I sculptured to its extreme apex; striations on tergite II reach basal; genitalia
feebly sclerotized, narrowed medially above digiti and below basal ring, digiti stout with
3 small digital teeth each, median short and broad with nearly straight apex, penis sheath
acutely rounded (Fig. 1183); general coloration black except yellow legs and scape, and
uniformly brown funicles; hosts – Scotinophara lunda, S. coarctata and S. latiuscula in
S. and S.E. Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus triptus Nixon
628' Moderately large species, 1.05 mm; postscutellum with a median subglobular strongly
rugose swelling; ocellar region with 3-4 punctures; antenna slender without a clearly
differentiated club; 1st funicular segment as long as pedicel, II and III elongate (Fig.
1184); fringe about half maximum wing width; stigmalis of forewings long; abdomen 1.5
times as long as wide with tergite I evenly striate (Fig. 1185); black with unicolorous dark
brown antennae, yellow legs except blackened coxae I-III and infuscated femur; host -
Nezara vindula and S. latiuscula in Asia . . . . . . . . . . . . . . . . . . Telenomus cyrus Nixon
629(627') Ocellar setae 2-3 pairs either widely separated or weakly developed; shallow postocellar
grooves behind lateral ocelli; hyperoccipital carina absent; temples drop-off sharply to
occipital carina, coriaceous sculpture along posterior orbits extending 0.33 distance to
occipital carina (Fig. 1186); metapleural carina well developed in the posterior half of
sclerite; antenna 11-segmented in the female, pedicel < 1st funicle, 4th funicle quad-
rate; V < VI; VII-VIII transverse; clava 5-segments; head and body black with all coxae
black to dark brown; body length 1.2-1.3 mm; host – Nezara viridula in Asia and Europe
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus chloropus Thomson
629' Ocellar setae absent, if present postocellar grooves wanting . . . . . . . . . . . . . . . . . . 630
630(629') Hyperoccipital carina present; frontal depression well developed; frons slightly bulging
between antennal insertions and inner orbits; 1st funicle distinctly longer than pedicel in
the female (Fig. 1187); funicular segments I to III much longer than wide and VI-IX
slightly longer than wide in the male; scutellum with submarginal foveae smaller than
dorsellar punctures; head and body black; yellow forecoxa with blackish base, mid-and
hind coxa dark brown concolorous to antennal segments II-XI; scape brownish yellow;
genitalia (Fig. 1188) with moderately broad digiti, 3 teeth/digitus; penis sheath rounded;
body length 1.10 mm in male and 1.1-1.4 mm in the female; host – Nezara viridula in
America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus cristatus Johnson
630' Hyperoccipital carina absent; frontal depression usually weakly developed . . . . . 631
631(630') Polyphagous egg parasitoids in American region; head and body dark brown to black;
vertex smoothly rounded, angled or rounded behind lateral ocelli; mesoscutum strongly
convex,- coriaceous-pustulate without longitudinal sculpture (Fig. 1189); setal bases
pustulate in the smooth scutellum; submarginal foveae subequal in size to dorsellar
punctures; dorsellum broadly triangular overlapping propodeum; abdominal tergite I
with 2-3 pairs of sublateral setae (Fig. 1190); antenna of female 11-segmented, segment
II<III; VI (4th funicle) transverse, VII < VIII; wing clear; genitalia slender, digiti and
teeth (3/digitus) large; median plate with parallel sclerotized rods near digiti; penis
sheath short and almost broadly rounded (Fig. 1191); body length 0.9-1.4 mm in female,
0.9-1.6 mm in male; hosts — N. viridula, Oebalus pugnax, Mormoidea poecilia . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus podisi Ashmead
631' Not as above .......................................................... 632
632(631') Egg parasitoids of Nezara antennata in E. Asia; vertex rather narrow and well convex;
antennae and coxae nearly black throughout, pedicel slightly longer than broad, shorter
than 1st funicular segment, 2nd funicular segment about as long and broad as first; 3rd
1.7 times as long as broad and 0.9 times as long as 2nd; 4th - 9th slightly longer than broad
(Fig. 1192); abdominal tergite II about as long as broad and weakly longitudinally
striated basally; body length of male 0.9 mm and 1.2 mm in the female . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus nakagawaii Watanabe
632 ' Egg parasitoids of Scotinophura, Piezodorus and Eysarcoris in E. Asia; vertex rather flat
and broad; postero-lateral part behind eye angulate and very narrow (Fig. 1193);
pedicel 2.2 times as long as the 1st funicular segment; 3rd funicular oblong, 4th globular
and club with 5 segments in female (Fig. 1194); male with funicular segment I to III
much longer than IV to IX; black except for dark brown antenna with yellowish brown
to yellow scape and apex of pedicel; legs including coxae uniformly yellow; penis sheath
short and apically rounded (Fig. 1195); body length 0.9 mm in male and 1.1 mm in
female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telenomus gifuensis Ashmead
633(609') Small scelioninae, body length barely 1.6 mm; egg parasitoids of rice bug Leptocorisa
oratorius; head with reticulate sculpture all throughout except transverse intercon-
nected rectangular sculptures in frontal area above antennal bases (Fig. 1196); deep
ridge at the back of head and strongly rounded at front; mesothorax and scutellum with
rough peg-like structures (Fig. 1197); otherwise finely punctated, parapsidal furrows
absent; antenna brownish yellow with yellowish scape and distal end of pedicel; pedicel
inverted, cone-shaped and much longer than 3rd; 4th - 6th quadrate; 7th - 12th form the
club, also quadrate except cone-shaped apical segment (Fig. 1198); wing hyaline with
small hairs; veins brown; abdomen broadly ovoid; barely longer than wide, l st tergite 4
times broader than long; black with yellow legs . . . . . . . . . . . . . . Gryon nixoni Masner
633' Relatively large species, body length 3-4 mm; egg parasitoids of grasshopper ... 634
634(633') Submarginal veins in hindwing incomplete, short, usually stump-like; wing smoky,
forewing with a longitudinal hyaline streak just below middle and veins dark brown;
lower part of frons between eye and antennal insertion with some short carinae converg-
ing towards mouth; hosts - Oxya velox and O. japonica . . . . . . . . . . . . . . . . . . . . . . 635
634' Submarginal veins complete, reaching fore margin at frenal hooks; hosts — grasshop-
pers or lepidopteran eggs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 636
635(634) First tergite of abdomen subquadrate, a little longer than broad at base (Fig. 1199);
scutellum not armed; legs reddish yellow; veins dark brown; stigmal vein slender, finely
defined and not apically rounded (Fig. 1200); 2nd tergite hardly impressed on basal half;
in male the 1st joint of flagellum thickened, being twice as long as thick; 3rd the longest
but not conspicuously enlarged; basal half of scape reddish yellow in male with the rest
fuscous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scelio muraii Watanabe
635' First tergite of abdomen transverse, being distinctly shorter than broad at base (Fig.
1201); scutellum armed; legs yellow; veins darker, almost black; stigmal vein stout,
sharply defined and apically rounded (Fig. 1202); 2nd tergite clearly impressed on basal
half; in male first flagellum slender, not distinctly thickened; 3rd joint conspicuously
enlarged; female scape reddish yellow and only fuscous at extreme apex . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scelio tsuruokensis Watanabe
635'' First tergite of abdomen transverse similar to S. tsuruokensis but short and truncate
anteriorly (Fig. 1203); 2nd broader and a little longer than 1st, 3rd twice as long as 2nd;
each segment longitudinally striate from base to end with 11 striae on first and 15 finer
ones of second; legs red except black coxae and brown midposterior femur of female;
wing slightly infuscated, covered with brownish cilia except hairless base; hindwing large
and quite hyaline; scutellum unarmed; metanotum with a row of large grooves separated
by transverse carinae; propodeum hollowed posteriorly and with a large tooth on each
side; antenna of female scape 4 times as long as pedicel and pedicel nearly as long as
first flagellum; male’s 3rd flagellum globular (Fig. 1204); host — Locusta migratoria
migratorioides in Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scelio zolotarevskyi Ferriere
636(634') Antennal scape very long, dilated dorso-ventrally and triangular in the females (Fig.
1205); body black, flat and highly depressed dorso-ventrally; antennae pale orange
concolorus to legs except blackened clubs; abdomen with punctures and striations, tip
end not strongly pointed (Fig. 1206); larval parasite of lepidopteran leaffeeders; body
length 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platyscelio abnormis Crawford
636' Antennal scape slender but not dilated as above (Fig. 1207); genae strongly developed
behind mid-inner portion of eye, making eyes rather oblique in lateral view; body
uniformly black except brownish legs; hindcoxae and antennae excluding darkened club
brownish black; entire body clothed with short white hairs; abdomen very long and
slender, 5.5 times longer than thorax and strongly pointed at tip (Fig. 1208); egg
parasitoids of Conocephalus longipennis; body length 4 mm . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macroteleia crawfordi Keiffer
637(519') Tarsi 3-segmented [Trichogrammatidae] (Fig. 1209) . . . . . . . . . . . . . . . . . . . . . . . . . 638
637' Tarsi 4- or 5-segmented (Fig. 1210) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 662
638(637) Antennal club of female 1-segmented; funicle 2-segmented; marginal vein curved distal-
ly away from anterior border of front wings to extend up to stigmal vein; setae of disc
usually in vein tracks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 639
638' Antennal club of female with more than one segment . . . . . . . . . . . . . . . . . . . . . . . . 654
639(638) Antennal club of male 3-segmented bearing 2 funicular segments; vein track of radial
sector (RS1) absent; forewing with long marginal fringe; stigmal vein elongate
(Trichogrammatoidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 640
639' Antennal club of male 1-segmented with unsegmented flagellum but with 2 slight
constrictions below in the region of funicular segments; forewing with short marginal
fringe; vein track RS1 present (Trichogramma) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 644
640(639) Chelate structure of male genitalia large and protruding extended beyond apical ends
of gonoforceps; median ventral projection (MVP) indistinct; lateral tubercles
prominent in between of ventral ridges highly chitinized (Fig. 1211); forewing with
rounded outer margin, long fringe setae about width of remigium (Fig. 1212); ovipositor
of female slightly longer than hind tibiae (egg parasite of Chilo sp. and Diopsis
thuracica) (Ghana) . . . . . . . . . . . . . . . . . . . . Trichogrammatoidea simmondsi Nagaraja
640' Chelate structure small or large, far below or at most near gonoforceps . . . . . . . . . 641
641(640') Chelate structures distinctly far below gonoforceps, median ventral projection distinct
although short, lateral tubercles not distinct; aedeagus apodeme nearly 0.8 times length
of tibia III (Fig. 1213); ovipositor as long as tibia 111, width of plate about half the length;
basal infuscation in line with stigma, remigium with sparse and long setae, fringe setae
long, more than one-third width of remigium (Fig. 1214); flagella of male antenna with
22-26 long hairs; orange color except bright red eyes; light brown thorax and abdomen;
S. and S.E. Asia . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogrammatoidea nana (Zehntner)
641' Chelate structures rather near gonoforceps; other characters not as above . . . . . . 642
642(641') Median ventral projection, lateral tubercles, and central ridges indistinct; aedeagus with
apodemes subequal in length, together 0.6 length of tibia III (Fig. 1215); ovipositor
slightly longer than tibia III, width of plate less than half the length; antennal flagella
nearly 1.5 times scape, with moderately long 25-32 hairs, the longest more than 2 times
width of flagellum; basal infuscation light, in line with stigma (Fig. 1216); ochreous with
bright red eyes, yellowish brown thorax and dark reddish brown abdomen; egg parasite
of Heliothis armigera . . . . . . . . . . . . . . . . . . . . . Trichogrammatoidea armigera Nagaraja
642' Median ventral projection present; lateral tubercles distinct or indistinct . . . . . . 643
643(642') Median ventral projection distinct with distinct lateral tubercles (Fig. 1217); basal
infuscation not reaching stigma; fringe setae short (Fig. 1218); flagellar hairs of anten-
nae about 30, all long . . . . . . . . . . . . . . . . . . . . . . . . . .Trichogrammatoidea lutea Girault
643' Median ventral projection not conspicuous with lateral tubercles distinct (Fig. 1219);
basal infuscation in line with stigma; fringe setae long (Fig. 1220); antennal flagella with
22-26 long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogrammatoidea bactrae Nagaraja
644(639') Median ventral projection absent or inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . 645
644' Median ventral projection present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
645(644) Dorsal expansion of gonobase (DEG) with a broadly rounded posterior extremity,
extruding slightly beyond level of gonoforceps and base constricted; median ventral
process not conspicuous; aedeagus slightly longer than apodeme; chelate structure near
tips of gonoforceps (Fig. 1221); ovipositor longer than tibia III; antennal flagella with
about 35 nearly blunt hairs of moderate length, the longest 2.5 times width of flagella;
light yellow except blackish cheeks, sides of thorax, scutellum, legs and abdomen; (egg
parasites of hesperiids) . . . . . . . . . . . . . . . . . . . . . . . Trichogramma hesperidis Nagaraja
645' Dorsal expansion of gonobase horseshoe-shaped basally; fringe of forewings one-fifth
width of wings; aedeagus distinctly longer than apodeme, both together as long as or
longer than tibia III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
646(645') Ovipositor markedly extruded beyond tip of abdomen; female with white thoracic
ventral region; antennal hairs of male long, the longest 4 times maximum width of
flagella; male genitalia elongate, chelate structure far below tip of gonoforceps,
aedeagus about 3 times longer than apodeme (Fig. 1222) . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogramma pallidiventris Nagaraja
646' Ovipositor slightly exserted, longer than tibia III which is as long as aedeagus with short
arm (Fig. 1223); male genitalia elongate ovate, chelate structure in line with MVP (Fig.
1224); thoracic venter of female never white; antennal hairs of male long, the longest 3.5
times maximum width of flagella . . . . . . . . . . . . . . Trichogramma japonicum Ashmead
647(644') Antennal hairs very long, longest hair 3.5-4 times maximum width of flagellum . . . 648
647' Antennal hairs moderately long, longest hair 2 to 2.5 times maximum width of flagellum
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 649
647'' Antennal hairs very short, the longest as long as maximum width of antenna; flagella
unsegmented (Fig. 1225); longest marginal setae of forewing one-quarter wing width
(Fig. 1226); dorsal expansion of gonobase triangular tapering at tip and extended to
chelate structure (Fig. 1227); aedeagus longer than apodeme (Fig. 1228); ovipositor
longer than hind tibia (egg parasite of Chilo diffusilineus) . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogramma mwanzai Schulten & Feijen
648(647) Dorsal expansion of gonobase basally narrow, highly chitinized, triangular and con-
stricted basally (Fig. 1229); longest antennal hair 3.5 times maximum width of flagellum;
aedeagus (Fig. 1230) slightly longer than apodeme, both together slightly shorter than
tibia III, ovipositor longer than tibia III (Fig. 1231); fringe on tornus nearly one-eighth
of wing width; brownish yellow with black thorax and abdomen (Americas) . . . . . . . . .
.......................................... Trichogramma semifunatum (Perkins)
648' Dorsal expansion of gonobase broadly triangular at base with conspicuously rounded
sides, median ventral process (MVP) below chelate structure (Fig. 1232); longest hair
of antenna 4 times maximum width of flagellum; forewings much longer than body;
aedeagus and apodeme equally long (Fig. 1233); together shorter than tibia which is as
long as ovipositor (Fig. 1234); dull yellow body with brownish head, pronotum and
abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogramma evanescens Westwood
649(647') Longest antennal hair 2.5 times maximum width of flagellum . . . . . . . . . . . . . . . . . . 650
649' Longest antennal hairs not more than 2 times maximum width of flagellum . . . . . . 651
650(649) Antennal flagella of male with 50 or more hairs; dorsal expansion of gonobase with
broad lateral lobes basally; median ventral projection very large, slightly below chelate
structure (Fig. 1235); apodeme longer than aedeagus (Fig. 1236) (E. Asia) . . . . . . . . . .
.......................................... Trichogramma dendrolimi Matsumura
650' Antennal flagella of male with less than 35-40 hairs; dorsal expansion of gonobase with
narrow lateral lobes basally; small median ventral projection sharp, ovate to long ovate
(Fig. 1237); apodeme as long as aedeagus (Fig. 1238), together shorter than tibia III;
ovipositor longer than tibia III (Fig. 1239) . . . . . . . . . . . . . Trichogramma chilonis Ishii
651(649') Aedeagus as long as apodeme, both together equal to 1.2 length of gonobase (Fig. 1240);
median ventral projection sharp and triangular (Fig. 1241), ovipositor slightly longer
than tibia III; fringe on tornus of forewing about one-sixth of wing width (Fig. 1242);
yellow with dark brown abdomen; terminal segment of male antenna narrowed apically
(Fig. 1243); egg parasite of Mythimna separata) (China) . . . . . . . . . . . . . . . . . . . . . . . . . .
......................................... Trichogramma leucaniae Pang & Chen
651' Aedeagus longer than apodeme, other characters not as above ................ 652
652(651') Aedeagus much longer than apodeme (Fig. 1244); dorsal expansion of gonobase highly
sclerotized, with large lateral lobes, sharp, and triangular central lobe; median ventral
projection almost in line with chelate structure (Fig. 1245); wing infuscation darker than
T. leucaniae (Fig. 1246); male antennal hairs long, apical half of terminal segment large
(Fig. 1247); egg parasite of Mythimna separata in China . . . . . . . . . . . . . . . . . . . . . . . . . .
............................................ Trichogramma ivelae Pang & Chen
652' Aedeagus just longer than apodeme; dorsal expansion of gonobase moderate to highly
chitinized (America) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653
653(652') Chelate structures nearer to gonoforceps; median ventral projection below level of
chelate structures (Fig. 1248); aedeagus longer than apodeme (Fig. 1249), both shorter
than tibia III which is a little shorter than ovipositor (Fig. 1250); adult yellow except
anterior terga of abdomen dark pigmented in female and entire abdomen in male . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogramma perkinsi Girault
653' Chelate structures far below level of gonoforceps; median ventral projection nearly in
line with chelate structures (Fig. 1251); apodeme as long as aedeagus (Fig. 1252),
combined length shorter than tibia III which in turn is shorter than ovipositor (Fig.
1253); adult yellowish brown with fuscous head, thorax, and abdomen . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichogramma frasciatum (Perkins)
654(638') Marginal fringe of forewing at least half maximum wing width; discal ciliation usually in
rows but sparse; female antenna 2-segmented (ring and funicle) (Oligosita) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 655
654' Forewing with a narrow smoky or brownish tinge across wing beneath stigmal vein (Fig.
1254); scape as long as combined lengths of pedicel and funicle (Fig. 1255); gaster pale
yellow except tergites latero-dorsally dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracentrobia yasumatsui Subba Rao
654" Forewing with a rounded brown tinge below stigmal vein (Fig. 1256); scape a little
longer than combined length of pedicel and funicle (Fig. 1257); tergites I-IV of ab-
domen uniformly brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracentrobia andoi Ishii
655(654) Marginal fringe of forewings one-third or one-half the maximum width of wings . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 656
655' Marginal fringe of forewings slightly shorter or longer than maximum width of wings . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
656(655) Discal cilia short, fine, and very sparse, at least 10 rows visible; short hairs absent along
margins; infuscation distinct, a little beyond stigmal vein at middle, 15 small setae
arranged in 3 rows at middle of infuscation below marginal vein; marginal vein with 3
long setae, stigmal vein knob-like with subtruncate apex and 4 distal sensilla and a long
seta above midlength of stigma; forewing broad and rounded; fringe about one-third
maximum wing width; pedicel about half of scape (Fig. 1258); femur III with 13
transverse striations and 3 ventral setae (Fig. 1259); orange-bodied species with light
yellow-brown legs, antennae and mesoscutum . . . . . . Oligosita consanguinea Girault
656' Discal cilia long and coarse; marginal fringe of forewings about half maximum width of
wings, always greater than one-third (Fig. 1260); pedicel 3 times as long as funicle
segment, the latter prominently wider than long; scape broadened basally, funicle and
basal 2 clubs bisected medially by a transverse row of long setae (Fig. 1261); host
unknown but collected from ricefield . . . . . . . . . . . . . . . . . . Oligosita brevicilia Girault
657(655') Pedicel as long as the short and clavate scape (Fig. 1262); funicular segment longer than
broad; forewings with distinct and large substigmal cloud; yellow except fuscous tips of
coxae and midfemora, and dark brown basal half of abdomen . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligosita manii Viggiani
657' Pedicel usually much shorter than scape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 658
658(6571) Pedicel almost as long as the narrow funicle; club as long as the combined length of
pedicel, ring, and funicle segments; cilia on discal area very sparse, a single complete
row lining the apical end of wing and 10 more cilia irregularly scattered at the distal end
(Fig. 1263) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligosita shibuyae Ishii
658' Pedicel at least twice as long as funicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 659
659(658') Marginal fringe of forewing as long as or longer than the maximum width of wings; discal
cilia moderately sparse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 660
659' Marginal fringe of forewing more than 0.7 maximum width of wing but not as long as
wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 661
660(659) Marginal fringe of forewing distinctly longer than maximum wing width (Fig. 1264);
pedicel more than twice the length of funicle, the latter slightly wider than pedicel (Fig.
1265); sheath of ovipositor not exserted . . . . . . . . . . . . . . . . . . . Oligosita naias Girault
660' Marginal fringe and maximum width of forewing equally long (Fig. 1266); funicle half
the length of pedicel (Fig. 1267); sheath of ovipositor clearly exserted . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligosita yasumatsui Viggiani & Subba Rao
661(659') Marginal fringe of forewing only slightly shorter than maximum wing width; discal
ciliation rather thick opposite subtriangular stigmal vein, moderately scattered towards
distal end (Fig. 1268); pedicel more than 2 times longer than funicle (Fig. 1269) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligosita aesopi Girault
661' Marginal fringe almost as in O. aesopi; funicle subglobular and less than half of pedicel
(Fig. 1270); discal ciliation more evenly scattered towards distal end, without dense
ciliation opposite the knob-like stigmal vein (Fig. 1271) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oligosita nephotettica Mani
662(637') Basal part of hindwing reduced to thread-like stalk composed of the sub-marginal vein;
wing disc ribbon-like; forewing with a long marginal fringe; antennal sockets wide apart
[Mymaridae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 663
662' Hindwing never stalk-like basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 682
663(662) Gaster with more or less distinct petiole, convexly rounded or subglobular basally;
mesopostphragma not projecting into the gaster; tarsi 4- or 5-segmented . . . . . . . 664
663' Gaster broadly connected to the propodeum (hindpart of thorax); mesopostphragma

plainly projecting into gaster; a pair of distinctly separated plates ( = postscutellum)
behind scutellum; antennae 9-segmented in the females (6 funicular segments, solid
club) and 13-segmented in the males; tarsi Csegmented (Anagrus) . . . . . . . . . . . .674
664(663) Tarsi 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 665

664' Tarsi 4-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .672
665(664) Petiole long and slender (Fig. 1272); propodeum prominently carinated; antenna II-
segmented with 8 funicular segments and undivided club (Fig. 1273) . . .Ooctonus sp.
665' Petiole short, wider than long; antennae of male 13-segmented (Fig. 1274) and female
11-segmented, with 8 funicular segments (Fig. 1275); marginal vein not elongated,
venation not reaching basal one-third of wing; gaster subsessile (Gonatocerus spp.). . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 666
666(665') Anal plate with 4 long hairs (Fig. 1276) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667
666' Anal plate with more than 4 long hairs (Fig. 1277) . . . . . . . . . . . . . . . . . . . . . . . . . . .669
667(666) Apical 4 funicular segments distinctly longer than the basal four . . . . . . . . . . . . . . .668
667’ Apical 4 funicular segments as long as or slightly shorter than the basal 4, 4th to 6th
segments cylindrical equally long and without sensory ridge; segments VII and VIII with
a pair of sensory ridges each (Fig. 1278); median of mesoscutum with a subglobular
brown band in apical half (Fig. 1279); forewing (Fig. 1280) 4.5 times as long as wide; anal
plate large, tapers apically and pygostyle indented subbasally inside (Fig. 1231) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatocerus uttardecanus Mani & Saraswat
668(667) Funicular segments V-VIII longer than basal 4 segments, cylindrical to subcylindrical,
equally long and with a pair each of sensory ridges (Fig. 1282); entire median area of
mesoscutum occupied by a brown band except lateral margin near parapsidal furrow
(Fig. 1283); forewing 3 times longer than wide (Fig. 1284); anal plate short hairs in 2
rows, apex of plate pointed; pygostyle without indention (Fig. 1285); postphragma very
flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatocerus narayani (Subba Rao & Kaur)
668' Funicular segments V to VIII globular, each with sensory ridges except segment VI, and
segment VIII with a cavity (Fig. 1286); mesoscutum moderately produced and rounded
apicomedially with a brown band above tip of parapsidal furrow (Fig. 1287); forewing
rather concave along anterior margin, about 4 times as long as wide; marginal vein 7.4
times as long as wide; proximal sensillum midway of distal sensilla and distal macro-
chaeta (Fig. 1288); 4 anal plate hairs in a transverse row in apical one-third (Fig. 1289)
and cleft at apex . . . . . . . . . . . . . . . . . . . . . . Gonatocerus devitatacus Mani & Saraswat
669(666') Anal plate triangular with 7 long hairs mostly along margins (Fig. 1290); apicomedian
band of mesoscutum reaching only pair of setae (Fig. 1291); forewing slightly con-
stricted near stigmal vein, nearly 4 times as long as wide, marginal vein 14 times as long
as wide; stigmal vein bears 3 distal sensilla (Fig. 1292); pedicel as long as funicular
segment III, F6 = F7, and club nearly as long as radicula and scape (Fig. 1293); abdomen
light yellow except blackish brown tergites V and VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatocerus munarus Mani & Saraswat
669' Anal plate suboval or ovoid; funicular segments of various lengths . . . . . . . . . . . . 670
670(669') Anal plate ovoid with 5 hairs in 3 transverse rows (Fig. 1294); basal funicular segment of
antennae shortest in both sexes; 0.7 as long as 2nd, 2nd and 3rd subequal, 4th to 8th
equally long, a pair of sensory ridges only in segments V to VIII in the female (Fig. 1295)
and all segments in male except scape and pedicel (Fig. 1296); lateral sides of mesos-
cutum behind parasidal furrow entirely yellowish brown to brown (Fig. 1297); forewing
4 times as long as wide; proximal sensillium just below distal macrochaeta (Fig. 1298);
ovipositor highly exserted . . . . . . . . . . . . . . . . . . . . . Gonatocerus cicadellae Nikolskaya
670' Anal plate subovoid; basal 3 funicular segments equally long . . . . . . . . . . . . . . . . . 671
671(670') Anal plate bluntly conical with 5 long hairs in 2 transverse rows medially (Fig. 1299);
female antenna with 4th and 6th funicular segments globular and equal in length, 6th the
shortest, paired sensory ridges in segments V, VII, and VIII (Fig. 1300); shortest
funicular segment I (F1) as long as scape, club = F3 = F4 = F8 = F10 in male (Fig.
1301); discal hairs a little beyond base of marginal vein in origin (Fig. 1302); abdomen
with alternating yellow and brown transverse bands (Fig. 1303) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . Gonatocerus cincticipitis Sahad
671' Anal plate narrowly conical with a subacute apex (Fig. 1304); female funicular segments
IV, V, and VI globular and equally long, paired sensory ridges only in VII and VIII
segments (Fig. 1305); funicular segment X (F10) as long as club, and longer than F6 to
F8 in the male (Fig. 1306); discal hairs emanate from base of marginal vein and form a
prominent oblique line to posterior margin; abdomen with four brown interrupted
transverse bands (Fig. 1307) . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatocerus miurai Sahad
672(664') Hindwing with few marginal cilia, abbreviated, filiform and highly reduced; forewing
(Fig. 1308) strongly petiolated and oar-shaped, distal half of broad part dark brown with
one clear row of setae traversing middle from shaded to light area and 2-3 short rows of
setae dorsad it in shaded portion; marginal fringe 4 times maximum width of wing; scape
medially thin, enlarged on both ends, 1st funicle as long as pedicel, II the longest,
segments III to VI short and together with undivided club slightly longer than the very
long 2nd segment (Fig. 1309); basal tarsal segment 2 times length of 2nd . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mymar taprobanicum Ward
672' Not with the combination of above characters; forewing hyaline with normal discal cilia
....................................................................... 673
673(672') Scape without scale-like structures; marginal vein thickened (Fig. 1310); prothoracic
spiracles in normal position . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynema sp.
673' Scape with prominent scale-like structures, pedicel broad, funicles I to III long and in
decreasing length, and in increasing diameter towards club (Fig. 1311); marginal vein
thin (Fig. 1312); prothoracic spiracles positioned towards middle of the body, on the
line between pronotum and mesoscutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stephanodes imbricatus (Narayanan & Subba Rao)
674(663') Funicular segment I long, usually 2 times as long as wide; scape smooth without
transverse ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 675
674' Funicular segment I very short, subglobular; scape transversely carinated . . . . . . . 677
675(674) Funicular segment I slender and longer than pedicel (Fig. 1313); antennae long and
narrow; ovipositor hardly exserted (Fig. 1314), ratio of ovipositor and exserted part 20:1;
disc of forewing narrow, nearly parallel sided, with 1 row in basal two-thirds and 2 rows
in the apical one-third (Fig. 1315) . . . . . . . . . . . . . . . . . . . . Anagrus optabilis (Perkins)
675' Funicular segment I shorter or as long as pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . 676
676(675') First funicular (Fig. 1316) segment as long as pedicel; ovipositor highly exserted; apical
2 segments of tarsi III as long as the exserted part of ovipositor; disc (Fig. 1317) of
forewing slightly dilated in the apical one-third, ciliation in an irregular row towards
basal half and in 2-3 irregular rows in apical one-third . . . Anagrus perforator Perkins
676' First funicular segment distinctly shorter than pedicel (Fig. 1318); ovipositor just
moderately exserted, length of exserted part as long as apical segment of tarsus III; ratio
of ovipositor and exserted part 5.4:1; proximal two-thirds of forewing parallel after
marginal vein, distal one-third gradually expanded and curved with long marginal cilia,
a long midlongitudinal line of discal hairs from distal end of marginal vein to wing apex
present, accompanied by another irregular short line of 6-8 hairs dorsad of long line
(Fig. 1319) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagrus panicicolae Sahad
677(674') Distal end of forewing disc dilated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 678
677' Distal end of forewing disc not dilated; 2nd funicular (E) segment longest . . . . . . 680
678(677) Marginal vein with 2 long setae; forewings with 3-4 rows of discal hairs with distinct
small bare space at the widest part (Fig. 1320); gaster conical; ovipositor moderately
exserted; 1st funicle very small and subglobular, F2 longer than F3 and the longest (Fig.
1321) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagrus flaveolus Waterhouse
678' Marginal vein with only 1 long seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 679
679(678') Forewing with 8-9 irregular rows of discal hairs and without bare or hyaline space at its
widest part (Fig. 1322); 3rd funicular segment with only 1 sensory ridge (Fig. 1323);
ovipositor moderately exserted, ratio of ovipositor and its exserted part 9.2:1 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagrus incantatus Haliday
679' Forewing with 2 regular rows (rarely 3) of discal hairs and with hyaline space in its
widest part, ciliation moderately more towards proximal end (compared with the distal
end) (Fig. 1324); ovipositor clearly exserted beyond tip of abdomen; club with 1 sensoria
(Fig. 1325) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagrus sp. A
680(677') Funicular segment II the longest; segments IV to VI subequal in length and diameter
(Fig. 1326); discal ciliation 2 lines, forms a long clear hyaline area distally; marginal
fringe almost 3 times the maximum width of wing (Fig. 1327); ovipositor short to slightly
exserted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anagrus frequens Perkins
680' Funicular segment II as long as segment IV; may or may not be as long as segment VI;
ovipositor short or long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 681
681(680') Ovipositor long, prominently exserted beyond tip of gaster; ovipositor length a little over
twice the length of apical segment of ovipositor sheath; 2nd funicular segment as long as
4th but shorter and much more slender than the broad 6th segment (Fig. 1328); 1 row of
discal ciliation visible along clear area (Fig. 1329) . . . . . . . . . . . . . . . . . Anagrus sp. B
681' Ovipositor short, slightly exserted beyond tip of gaster; funicular segments II, IV, and
VI subequal; distant end of forewing with a hyaline area; marginal fringe a little longer
than the maximum wing width (Fig. 1330) . . . . . . . . . . . Anagrus armatus (Ashmead)
682(662') Tarsi 4-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 683
682' Tarsi 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 700
683(682) Clava enlarged, entire with apex strongly and obliquely truncate; antenna 8-segmented
(Fig. 1331); sensillae of forewings at apex of stigmal vein arranged symmetrically, basal
one-third of front wings with few hairs, tip of marginal vein weakly hyaline prior to
stigmal vein (Fig. 1332); this grey to black polyembryonic parasite oviposits in eggs and
emerges during the host larval stage (Encyrtidae) — host C. medinalis, Marasmia spp.,
Hedylepta indicata . . . . . . . . . . . . . . . . . . . . . . . . . . . Copidosomopsis nacoleiae (Eady)
683' Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 684
684(683') Hind coxa enlarged laterally compressed, and disc-like; long bristles on hind tibia form
diamond-like configurations; antennae with three funicles and a conspicuous anellus,
usually ramose in males; marginal vein very long; body compressed laterally [Elasmidae]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 685
684’ Hind coxa not greatly enlarged, if so, not compressed or disc-like and with different
pattern; hind tibia without diamond-like arrangement of bristles; gaster distinctly con-
stricted at junction with the propodeum [Eulophidae] . . . . . . . . . . . . . . . . . . . . . . . 687
685(684) Head, thorax, and abdomen yellow except blackened ocellar triangle, dorsal part of
coxae, mid and lateral edges of scutellum, lateral edge of propodeum, abdominal
tergites II and III, middle of V-VI and edges of IV to VI in the female (Fig. 1333) and
entire tergites II to IV and apical half of male . . . . . . . . Elasmus albopictus Crawford
685' Color pattern not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 686
686(685') Uniformly dark colored (Fig. 1334); forewing long and very narrow and evenly rounded
distally (Fig. 1335) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elasmus corbetti Ferriere
686' Not entirely black; venter, mid-and lateral sides of abdomen orange (Fig. 1336); antenna
yellowish brown with very short funicular joints; hind coxae black with tip yellow, femur
III with blackened dorsum and venter (Fig. 1337); body length 1-1.3 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elasmus philippinensis Ashmead
686'' Blackish green on head, mesonotum, and propodeum; postscutellum yellow; antenna
brown except yellow scape; abdomen reddish yellow along the sides; legs yellow except
greenish black coxae II and III and black striped upper part of hind femora (Fig, 1338);
forewing slightly widened distally, wing tip with its apical half rounded (Fig. 1339) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elasmus claripennis (Cameron)
687(684') Forewing with interruption between submarginal vein and parastigma; tarsus usually
4-segmented [Eulophidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 688
687' Forewing with submarginal vein smoothly joining parastigma . . . . . . . . . . . . . . . . 697
688(687) Notaular depression shallow and not defined, hind midlobe broad and weakly emar-
ginate with deep impressed line; pronotum strongly arched with smooth and laterally
obtuse-angular collar; scutellum less convex, reticulation on basal half more elongate
and weak anteriorly; dorsellum large and impressed laterally; forewing with bare costal
cell, pilose basal vein, small speculum, densely hirsute wing disc, very small stigma,
postmarginal vein more than twice as long as stigma vein; petiole slightly transverse,
reticulate with an elevated median carina in anterior half; head transverse; temples
converging behind eyes; eyes moderately hairy; antennal scape slightly swollen in apical
one-third and 5 times as long as broad; funicles longer than wide; gaster much shorter
than thorax; 1.2 to 1.4 times as long as broad, broadest subposteriorly; 1st tergite very
large about 0.80 of gaster (Fig. 1340); black with dark faint greenish tinge along sides of
thorax, propodeum and anterior part of tergite I (pupal parasite of Parnara guttata) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pediobius mitsukurii (Ashmead)
688' Similar to P. mitsukurii; pronotum without dorsally prominent extension towards
anteriorly rounded head; petiole not wider than long; antennae with slender scape
viewed dorsally and subglobular funicles; abdomen noticeably ovoid; forewing with at
least 2 transverse rows of hairs in the anal region, hindwing with few hairs toward basal
area (Fig. 1341) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pediobius atamiensis (Ashmead)
688'' Notaular line complete; axillae angularly produced; scutellum with two longitudinal
submedian grooves; pronotum without cross-carina; middle lobe of mesoscutum with
hairs, median area never bare, usually with a median groove; malar sulcus present;
stigmal vein long; funicle 3-segmented in female . . . . . . . . . . . . . . . . . . . . . . . . . . . 689
689(688'') Oriental species; body shining green with bluish lustre to uniformly-brownish black or
pale yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690
689' African species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692

690(689) Antenna with 4 ring segments; body pale yellow except patches of metallic green with
brassy and golden reflections in the thorax and sides of abdomen (Fig. 1342); scape of
male antenna enormously expanded beneath (Fig. 1343), innerly convex and outerly
concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ooletrastichus formosanus Timberlake
690' Antenna with one ring segment; blackish brown with greenish reflections or metallic
blue with lustre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691
691(690') Body uniformly brownish black with greenish reflections in the head and thorax; legs
whitish yellow except brownish coxa I and femur I; antenna blackish brown except
yellow-brown scape, funicular segments not slender (Fig. 1344); club of male antenna
black, broad and much wider than any antennal segment (Fig. 1345) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus howardi (Olliff)
691' Body metallic blue or shining green with bluish lustre; antenna brown except yellow
scape with brown sensoriae in male; legs yellow except greenish coxae I and IV; ocelli
in a low triangle, front ocellus never reached by scape; thorax smooth and shining,
abdomen elongate, pointed behind with a slightly exserted ovipositor; female antenna
(Fig. 1346) 3 equally long funicular segments 3 times as long as broad; male; 1st funicular
segment smaller than the rest, funicle and club clothed with short hairs, sensorial organ
of scape narrow and elongate and nearly as long as scape (Fig. 1347); abdomen ovoid,
slightly longer than thorax . . . . . . . . . . . . . . . . . . . . . . Tetrastichus schoenobii Ferriere
691'' Very similar to T. schoenobii; shiny green with bluish lustre; legs uniformly whitish
yellow with dark brown last tarsal segments (Fig. 1348); 3 equally long funicular seg-
ments only twice longer than broad; abdomen narrow and pointed behind, ovipositor
more exserted than in T. schoenobii (Fig. 1349) (pupal parasite of Hydrellia philippina)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus sp.
691''' Body dark brown or black with slight blue-green lustre; scape, trochanters, apices of
femora, entire tibia and tarsus pale yellow; head transverse wider than mesonotum;
clypeus emarginate medially and mandibles sharply bidentate (Fig. 1350); mesoscutum
with 3 pairs of bristles in inner basal half of parapsidal furrow (Fig. 1351); forewing
hyaline and broader apically; submarginal vein with 2 setae, stigma vein about one-fifth
of marginal vein (Fig. 1352); antenna with segment I > II = III (Fig. 1353); sensoria
about two-thirds of scape (Fig. 1354) host : S. inferens . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus inferens Yoshimoto
692(689') Head, thorax, and abdomen shiny and very finely reticulated; costal cell of forewing
12-13 times as long as broad, ventral area limed with many setae; submarginal vein with
7 large bristles dorsally (Fig. 1355); marginal vein as long as stigmal vein; propodeum
smooth with median carina; 1st funicular segment the longest, nearly 3 times as long as
broad, slightly shorter than club; male antenna with 5 funicular segments and clava with
a basal whorl of long setae (Fig. 1356); sensorial organ near apex of scape, less than half
length of scape; color pattern of pronotum, mesonotum, and scutellum light to dark
brown; larval-pupal parasitoid of African gall midge, Orseolia oryzivora . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus pachydiplosisae Risbec
692' Larval-pupal parasite of diopsid rice stemborers . . . . . . . . . . . . . . . . . . . . . . . . . . . 693
693(692') Head, thorax, and abdomen irregularly reticulated; large part of coxae reticulated or
unreticulated; ovipositor slight to clearly exserted . . . . . . . . . . . . . . . . . . . . . . . . . . 694
693' Head, thorax, and abdomen regularly reticulated; large part of coxae usually reticu-
lated; ovipositor moderate to fully exserted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
694(693) Yellow-bodied species with dark brown pronotum, anterior midlobe of mesoscutum,
propodeum, and stylus; ocellar triangle brown concolorous to scape and pedicel; ab-
domen with irregular brown areas along margins and posterior tergites; malar space 0.8
times length of eyes; groove present between ocellar triangle (lateral ocellus) and eye
margins; hindcoxae not reticulated; forewings 2.8 times as long as broad, not reaching
tip of gaster, costal cell without bristles at middle (Fig. 1357), dorsum of submarginal
vein with 4 bristles; gaster oblong-lanceolate; funicular segments without dense row of
marginal hairs (Fig. 1358); ovipositor clearly exserted . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus flavobruneus Schulten & Feijen
694' Blue-green with glossy green abdomen; malar space less than 0.8 times eye length; most
of hind coxa reticulated; forewing reaches tip of gaster, nearly 3X as long as broad,
costal area with few evenly dispersed hairs; dorsum of submarginal vein with 5 bristles
(Fig. 1359); gaster elongate; ovipositor slightly exserted; male antenna with 6 funicular
segments, rimmed basally by a whorl of 8-11 long bristles (Fig. 1360) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus confusus Schulten & Feijen
695(693') Malar space very wide, as long as eye length; forewing as long as broad, extended to tip
of gaster, dorsum of submarginal vein with four bristles costal area with about 12 setae,
sparsely set distally (Fig. 1361); ocellar triangle elevated, without groove between lateral
ocellus and eye margin; base of antenna rimmed dorsally with 4-5 long hairs, funicle I a
little over half of 2nd funicle (Fig. 1362); head and thorax bluish green, abdomen reddish
brown with blue-green lustre; ovipositor slightly exserted . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus risbeci Schulten & Feijen
695' Malar space less than eye length; other characters not as above . . . . . . . . . . . . . . 696
696(695') Male with 6 funicular segments, each with a basal incomplete row or whorl of 8-11
bristles; sensorial process 0.8 of scape length (Fig. 1363); malar space one-half eye
length; postspiracular carina well developed; body blue-green except yellow parts below
and between antennal sockets, genae; scape yellowish white with brown upperside
distally, pedicel brown to light brown, flagella uniformly brown; costal area with 6-7
setae evenly spaced towards distal end, submarginal with 6 setae; basal area of forewing
below submarginal without hairs (Fig. 1364); ovipositor clearly exserted; relatively large
species, male 1.6 mm and female 2.9-3.6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus variabilis Schulten & Feijen
696' Male with only 5 funicular segments, each lined at basal one-third or one-fourth with an
incomplete ring of 6-8 long bristles (Fig. 1365); sensorial process a little over half of
scape; scape as long as eye length; malar space 0.7 of eye length; basal area of forewing
with setae, 4-5 in the submarginal and 6 in the base of costal area (Fig. 1366); body
generally metallic green with green to dark green abdomen . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetrastichus diopsisi (Risbec)
697(687') Mesoscutum with complete notaular lines; male antennal funicle not ramose . . . . 698
697' Mesoscutum with notaular lines incomplete or absent; male antennal funicle with rami
or lateral branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 699
698(697) Abdomen subsessile; hind tibia with small spurs, shorter than basitarsus; head broader
than mesoscutum; eyes finely pubescent, large in facial view; face triangular in frontal
view (Fig. 1367); mandibles hexadentate with 2 large teeth (Fig. 1368); antenna 11-seg-
mented with 2 rings and 3-segmented club; scape beyond vertex; propodeum highly
carinated, not oblique, with 4 setae in the hind lateral margin of propodeum; scutellum
flat and carinated distally; axillae well developed (Fig. 1369); yellow with blackish
antennae, grey eyes, brown patches on scutellum and abdominal tergites, midtergites of
IV and V usually with broad brown spots (Fig. 1370) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenomesius japonicus (Ashmead)
698' Abdomen subpetiolate; hind tibia with long spurs, longer one exceeds basitarsus;
propodeum oblique; scutellum without longitudinal furrows; body black except yellow
area at base of gaster; abdomen broadest submedially (Fig. 1371) . . . . . Euplectrus sp.
698' Abdomen ovoid, basal half yellow with 2 pairs of black spots laterally, latero-median
spots one-third - half that of subbasal lateral spots (Fig. 1372) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euplectrus chapadae (Ashmead)
699(697') Hind tibia with 2 apical spurs; toruli of antennae below middle of face; flagella some-
what flattened; costal cell wide, postmarginal vein more than 2X as long as stigmal vein;
propodeum weakly sculptured, convex and plicae present but without median carina
(Fig. 1373); abdomen black with or without subbasal white band (Fig. 1374) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sympiesis spp.
699' Hind tibia with 1 apical spur; toruli near middle of face; costa cell narrow; scape well
beyond front ocellus; male (Fig. 1375) antenna with 3 long rami; female with 2 anelli
(rings), 4 funicular segments with 2nd segment the longest and nearly 3X length of
pedicel, club 3-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiptarsenus sp.
700(682') Hind femur strongly enlarged with teeth in the ventral edge, thoracic dorsum rough,
with dense piliferous, umbilicate punctations; prepectus small; funicle 7-segmented plus
a ring segment; forewings not folding; ovipositor not upwardly curved over dorsum
[Chalcididae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 701
700' Hind femur more or less normal, ventral edge without serrated tooth . . . . . . . . . . 705
701(700) Coxae without an inner ventral tooth in either sex . . . . . . . . . . . . . . . . . . . . . . . . . . 702
701' Coxae without an inner ventral tooth in the females . . . . . . . . . . . . . . . . . . . . . . . . . 704
702(701) Preorbital carina of frons distinct, widely abbreviated near malar ridge; postorbital
carina indistinct; scrobe area polished, moderately deep reaching front ocellus (Fig.
1376); genae flat below eyes; right mandibles with 3 apical teeth; mesoscutellum
rounded at apex; scutellum gently declined posteriorly with apical part narrowly ex-
planate and reflexed, apical margin rounded (Fig. 1377); hind tibia mostly black, slightly
reddish basally with yellow spot subbasally and apically; femur III (Fig. 1378) black
except apical yellow spot, outer ventral margin with 10-12 teeth; dorsum of tergite 2 with
dense minute punctures . . . . . . . . . . . . . . . . . . . . . . . . . Brachymeria excarinata Gahan
702' Frons without preorbital carina but with or without postorbital carina . . . . . . . . . 703
703(702') Head 3 times wider than thorax; lateral ridges of scrobe produced in front of antennal
toruli; vertex distinctly deeply pitted; postorbital carina present reaching genotemporal
margin; genal angle acute in front and nearly rectangular behind; scutellum cleft medial-
ly in dorsal view; hind femur and tibia III yellow with a black broad patch at middle (Fig.
1379); abdomen a little shorter than pronotum; tergite VI with 6-7 transverse rows of
distinct and deep pits . . . . . . . . . . . . . . . . . . . . . . . . . Brachymeria megaspila Cameron
703' Head width nearly as long as thorax; lateral ridges of scrobe not produced in front of
antennal toruli; postorbital carina not well developed to almost absent; front genal angle
slightly acute and obtusely rounded off behind (Fig. 1380); scutellum in dorsal view not
divided medially; hindcoxa black with brownish red distal part, femur reddish brown
except yellow apex with a pale black patch on outer side, tibia yellow with black ventral
carina and brown base; abdomen a little shorter than 1.5 times length of pronotum,
mesoscutum, and scutellum combined; tergite VI with 5-6 transverse rows of indistinct
punctures . . . . . . . . . . . . . . . . . . . . . . . Brachymeria jayaraji Joseph, Narendran & Joy
704(701') Lateral ridges of scrobe produced in front of antennal toruli; smooth median area
prominent below scrobe, the latter reaching front ocellus; postorbital carina distinct,
extended to posterior margin of genotemporal part; front genal angle acute to slightly
rounded, hind angle rectangular but broadly rounded (Fig. 1381); scutellum weakly
bilobed medially; femur III (Fig. 1382) shiny black with clear yellow apical part, tibia III
creamy yellow with entire inner half reddish black; abdomen shortly ovoid with smooth
first tergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachymeria lasus (Walker)
704' Lateral ridges of scrobe not produced in front of antennal toruli; area below scrobe with
punctures and without a smooth median raised portion; scrobe not reaching front
ocellus (Fig. 1383); scutellum parallel-sided towards base and rounded apically; hind
femur (Fig. 1384) black with yellow distal ends; tibia III yellow with reddish black base;
abdomen black with first tergite faintly rough . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachymeria albotibialis (Ashmead)
705(700') Mesopleura large, convex and without any vertical grooves; midtibia with a strongly
produced straight apical spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 706
705' Mesopleura usually grooved and at least partly impressed . . . . . . . . . . . . . . . . . . . 708
706(705) Midcoxa closer to forecoxa; mesoscutum flat and convex dorsally, without impressions;
thorax finely reticulate, mesonotum more glossy than scutellum and wider than long;
scutellum rounded; antenna (Fig. 1385) yellow, inserted at base of face near clypeus,
scape short, pedicel elongate 0.4X length of scape and more than twice as long as broad,
funicle 1-3 small, subquadrate, 4-6 longer than broad and each longer than basal 3,
club-3-segmented and a little longer than 4-6; head transverse, frontally rounded with
narrow vertex; abdomen triangular, shorter than thorax; hypopygium reaching more
than 0.7 along gaster; body black with metallic luster on head and thorax [Encyrtidae]
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ooencyrtus malayensis Ferriere
706' Midcoxa closer to hind coxa; mesoscutum impressed posteriorly, longer than broad
without distinct shoulders; tibial spur of midleg very long, as long as combined length of
basal 3 segments and strongly hirsute; forewing with an oblique bare linea-clava; mar-
ginal vein longer than submarginal; antenna with 5 funicles and an anellus, clava with
obsolete second suture [Eupelmidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707
707(706') Brown infuscation of forewing originates at apex of speculum; mesoscutum white
medially; head and pronotum black, triangular area of pronotum sometimes whitish
(Fig. 1386) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neanastatus cinctiventris Girault
707 ' Infuscation starts halfway between oblique speculum and stigma, stigmal area to wing
tip infuscated; mesoscutum black with 2 whitish bands (Fig. 1387); vertex of head and
triangular area of pronotum yellow . . . . . . . . . . . . . . . . . . . Neanastatus oryzae Ferriere
708(705') Pronotum large and long, nearly quadrate and as broad as mesoscutum; notaular
grooves complete; propodeum sloping, median longitudinal groove present; marginal
vein normal, without infumation below; female (Fig. 1388) with 5-segmented funicle;
funicular segments each with a ring of bristles near the base and apex in male (Fig.
1389), club with many bristles; gaster with 4th tergite (Fig. 1390) the longest; body
non-metallic [Eurytomidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eurytoma spp.
708' Pronotum short, if long then with a punctated thorax, usually with metallic lustre; male
antenna with 5-7 funicular segments; forewing bare in basal one-third or with 13-seg-
mented antenna and pilose forewing; notauli often complete; hind coxa hardly larger
than forecoxa [Pteromalidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 709
709(708') Hind tibia with two spurs; fore femur golden yellow and unpigmented; head same color
as thorax, black with green metallic luster; abdomen pale to dark yellow with somewhat
broken median and clearly broken brown marginal stripes (Fig. 1391); antenna with 2
anelli in both sexes, female antenna with very long funicle segments and long erect hairs,
scape to anelli yellow and funicle to club dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Propicroscytus mirificus (Girault)
709' Hind tibia with only one spur; head and thorax reticulately punctated; scutellum without
frenum; occipital ridge near foramen far behind and below lateral ocelli; forewing not
very hairy, with fringe, postmarginal vein longer than stigmal vein . . . . . . . . . . . . . 710
710(709') Striae one clypeus confined to clypeal region; temples about one-fourth as long as eyes;
head in dorsal view not thick about one-fifth times as broad as long . . . . . . . . . . . 711
710’ Striae on clypeus reach lower margin of eyes and malar sulcus; temples nearly half as
long as eyes; dorsal view of head thick, less than twice as broad as long . . . . . . . . 712
711(710) Body black to bluish black; breadth of head slightly wider than combined length of
pedicel and flagellum in females; occipital carina weak; collar of pronotum weak but
sharply marginal except at side; malar space slightly shorter than 0.5 time height of eye;
anterior margin of clypeus slightly emarginate medially; mandible with 4 teeth; scape of
female antenna 0.8-0.9X eye height, flagella not slender, some subquadrate, 6th segment
quadrate to slightly transverse; club twice as long as broad; male with slender flagella,
scape above level of vertex, club 3 times as long as broad (Fig. 1392); subglobular gaster
with petiole as long as nucha; parasite of Oulema oryzae, Agromyza oryzae, Hydrellia
griseola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichomalopsis oryzae Kamijo & Grissell
711' Body green to bluish green; combined length of pedicel and flagellum 0.9-1.0X breadth
of head; occipital carina rather strong; collar of pronotum weak, irregularly marginal or
not marginal; malar space slightly longer than 0.5 times eye height; lower margin of
clypeus weakly emarginate medially; mandible with unequal teeth, left side with 3 and
right side with 4; scape of female antenna 0.9 to 1.0 times eye height; flagella rather
slender and weakly clavate distally, 1st and 2nd segment equally long, 6th subquadrate
to transverse and shorter than the first; club 2.2 times as long as broad; male with
antenna inserted higher, toruli nearer to anterior margin of median ocellus than to lower
margin of clypeus, combined length of pedicel and flagellum a little longer than head
breadth, 6th funicle longer than broad (Fig. 1393); hosts same as T. oryzae . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichomalopsis shiraki Crawford
712(710') Lower margin of clypeus cleft medially (Fig. 1394); lower margins of face not projecting
below level of lower margins of clypeus; occipital carina strongly curved medially;
dorsum of thorax moderately curved . . . . . Trichomalopsis apanteloctena (Crawford)
712' Lower margin of clypeus weakly emarginate; lower margins of face curved and
projected below level of lower margins of clypeus (Fig. 1395); occipital carina evenly
curved medially; thoracic dorsum evenly to weakly curved . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichomalopsis deplanata Kamijo & Grissell
713(2') Sedentary parasites, attached mainly to the abdomen of host leafhoppers and planthop-
pers; legless; female twisted-winged fly ( STREPSIPTERA ) . . . . . . . . . . . . . . . . . . .714
713' Mobile insects; legged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 716
714(713) Broad-passage opening circular-like, thorax reduced and ring-like behind broad-pas-
sage (Fig. 1396); parasites of delphacids and fulgorids . . . . . . . . . . . . . . Elenchus spp.
714' Not as above; parasite of cicadellids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715
715(714') Basal collar with 2 dark brown spots at base (Fig. 1397); parasite of Nephotettix, Recilia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halictophagus bipunctatus Yang
715' Basal collar without dark spots but with a pale median band (Fig. 1398); parasite of
Cofana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halictophagus spectrus Yang
716(713') Node absent; fontanelle present (ISOPTERA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 717
716' Node present; fontanelle absent (HYMENOPTERA) . . . . . . . . . . . . . . . . . . . . . . . 719

717(716) Pronotum flat without anterior lobes; head capsule oval, fontanelle lying at the middle
of the anterior margin; antenna 16-segmented; labrum subtriangular with a semihyaline
tip; mandible thin, moderately long and slightly sabre-shaped (Fig. 1399) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coptotermes fonnosanus Shiraki
717' Pronotum saddle-shaped with anterior lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 718
718(717') Labrum with hyaline tip; head subovate; antenna 16-17 segmented, segment III longer
than II or IV; left mandible with subbasal serrations, tip of mandible not strongly curved
inwards; pronotum with slightly pointed sides (Fig. 1400); S. Asian species . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrotermes gilvus Hagen
718' Labrum with a subhyaline tip; head much longer than wide and less ovate; antenna 18-19
segmented; left mandible without subbasal serrations, tips more inwardly curved;
pronotum with a transverse mark in anterior one-third, sides more rounded (Fig. 1401);
African species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrotermes bellicosus Smeathman
718" Labrum without a hyaline tip; head ovate; antenna 13-segmented, segment II almost
twice as long as III; mandible without serrations, tips strongly cross each other but
moderately curved inwards; pronotum with elevated anterior lobes, sides hump-like
towards lateral (Fig, 1402); African species . . . . . . . . . . . . . . . . . . . . . . Microtermes sp.
719(716') Two nodes ( = petiole) present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 720

719' One node present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 724
720(719) Propodeum with spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 721

720' Propodeum spineless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 723
721(720) Antenna 11-segmented with and 2-segmented club; palpal formula 2,2; workers strong-
ly polymorphic with small eyes and subquadrate head (Fig. 1403); propodeal spine
slightly curved downward towards the posterior (Fig. 1404); reddish brown ants . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pheidologeton sp.
721' Antenna 12-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 722
722(721') Antennal club 4-segmented (Fig. 1405); thoracic dorsum with a deep furrow in the
mesoepinotal joint; head subglobular and convex dorsally, vertex in frontal view notched
medially; hind leg without pectinate calcaria; propodea, spines small but nearly vertical
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pheidole sp.
722' Antennal club 3-segmented; thoracic dorsum entire and flat to convex anteriorly; head
subrectangular, width more than 0.7 mm, vertex not notched medially as seen in front,
enlarged part of first node subquadrate in lateral view, anterior face and dorsum
approaching each other through a curve, posterior face longer than anterior,
posterodorsal angle higher than anterodorsal; propodeal spine long and acute, up-
curved along its length (Fig. 1406) . . . . . . . . . . . . . . . . . . Tetramorium pacificum Mayr
723(720') Antenna (Fig. 1407) 12-segmented, club with 3 segments; eyes about 0.06 mm in
greatest diameter; head width less than 0.6 mm; body rough and opaque; basal half of
first node as high as the second (Fig. 1408) . . . . . Monomorium pharaonis (Linnaeus)
723' Antenna 10-segmented; club with 2 segments, nearly half of flagellum (Fig. 1409); eyes
relatively larger than M. pharaonis with more than 30 ommatidia; head width not less
than 0.5 mm in the smallest form; second petiole almost ovate; propodeal spiracle
almost ovate; propodeal spiracle almost at midlength (Fig. 1410) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenopsis geminata (Fabricius)
724(719') Petiole armed with 2 posterolaterally projected horn-like spines similar to that in the
propodeum (Figs. 931-933); pronotum with 2 similar spines projected forward; body
usually black or abdomen with silvery tinge . . . . . . . . . . . . . . . . . . . . . . . Polyrachis spp.
724' Petiole not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 725
725(724') Very small ants, less than 2 mm long; antenna 12-segmented, segment XII as long as X
+ XI; head blackish brown with wide V-shaped frons; antenna and gaster yellowish
brown; petiolar node rudimentary (Fig. 1411); scape surpassing occipital corner by
more than length of first funicular segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tapinoma melanocephalum (Fabricius)
725' Large ants, usually more than 10 mm long; node very prominent . . . . . . . . . . . . . . 726
726(725') Head, thorax and petiole including postpetiole strongly striated or grooved; body dull
black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 727
726' Head, thorax and petiole not striated; if so, striations very fine; mandibles parallel to
each other with 3 strong subapical teeth; area between base of antenna and eyes with a
deep groove, body glossy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 728
727(726) Pronotum with a lateral spine, transversely striated similar to the rest of thoracic dorsum
(Fig. 929); head striations widely diverge posteriorly and vertex moderately raised (Fig.
1412); scape hardly beyond back of head; petiole triangular in lateral view, slightly
concave in top view and with 3-4 long hairs; postpetiole smooth with long brown-yellow
setae and short white hairs; black with reddish brown legs, antennae and mandibles . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontoponera transversa (F. Smith)
727' Pronotum spineless, striations circular and transversely ovate at middle (Fig. 1413);
meso- and metastriations longitudinal and rather long; propodeal spiracle almost at
midlength and between striae 8 and 10 from bottom; petiole robust, striated transversely
with a moderate concavity between short erect spines (Fig. 1414); striae on top of head
above antennal base not expanded and long scape high above the back of head (Fig.
1415); femur and tibia black . . . . . . . . . . . . . . . . . . . . . . . Diacamma vagans (F. Smith)
727" Pronotum as in D. vagans but striations more transversely ovate (Fig. 1416); top meso-
and metastriations more oblique; propodeal spiracles on striae 11 and 13 from top;
petiole moderately stout, spines long and V-shaped in between spines (Fig. 1417); striae
on top of head expanded at middle; scape as above (Fig. 1418); all legs uniformly
reddish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diacamma sp.
728(726') Sting well developed (Fig. 1419); head with a medium longitudinal groove posteriorly;
mandibles long and parallel to each other, serrated with 3 strong teeth apically (Fig.
1420); petiole produced into a sharply pointed spine apically (Fig. 1421); head grooved
between antennal base and eyes (Fig. 1420) . . . . . . . . . . . . . . . . . . . Odontomachus spp.
728' Sting not developed (Fig. 1422); head without groove posteriorly and between eyes and
antennal base; mandibles short but robust, uniformly serrated with teeth of equal sizes
(Fig. 1423); petiole thinly narrowed apically, enlarged at middle (Fig. 1424) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camponotus sp.
729(1') Tarsi 2-clawed with or without claw tufts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 730
729' Tarsi 3-clawed, claw tuft absent though spurious claws present . . . . . . . . . . . . . . . . 769
730(729) Eyes arranged in 3 distinct rows, front row with 4 and highly enlarged anterior median,
second row with 2 small eyes and a third row with 2 medium sized eyes placed more
posteriorly on top of carapace (Fig. 1425); jumping spiders [Salticidae] . . . . . . . . . 731
730' Eyes arranged in 2 rows (Fig. 1426) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 739
731(730) Femur and tibia of leg I usually noticeably enlarged; tibia I armed with 3 pairs of ventral
stout spines sometimes with leaf-like setae extended up to femur and patella in leg I
only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 732
731' Femur and tibia of leg I not exaggerated in size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 734
732(731) Leg I with leaf-like setae (Fig. 1427) except in metatarsus and tarsus; tibia I 3-4 times
wider than metatarsus I, dorsally and ventrally lined with leaf-like setae; in male
metatarsus I much longer and lightly colored than tibia I; posterior lateral eyes slightly
outside carapacial margin (Fig. 1428); epigynum with a dome-shaped structure at

middle and a cavity on both sides (Fig. 1429); palpal organ brown with a shorter and
thinner embolius, tibial apophysis acute (Fig. 1430); abdomen a little longer than
cephalothorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harmochirus bruchiatus (Thorell)
732' Leg I without leaf-like setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 733
733(732') Posterior lateral eyes (PLE) almost at midlength of frontally flat carapace with a white
spot below each eye and a pair of subovate to subtriangular white spots at middle of
PLEs (Fig. 1431); median anterior eyes very large, mandibles very short and clypeus
relatively high (Fig. 1432); male abdomen with 3 pairs of white spots (Fig. 1433), female
pattern irregular; epigynum with a hat-like median structure (Fig. 1434); palpal organ
with a bent sharp-ended apophysis (Fig. 1435) . . . . . . . . Bianor hotingchiehi Schenkel
733' Posterior lateral eyes above midlength of carapace; cephalothorax flattened dor-
soventrally and short; abdomen ovoid with 2 pairs of apodemes and 3 white transverse
median, submedian, and subposterior bands (Fig. 1436), anterior half of abdomen white
along margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhene sp.
734(731') Sternum narrowed apically, narrower than base of labium; female abdomen with 2
longitudinal bands dorsally; chelicera with 2 promarginal and 1 retromarginal teeth; first
pair of legs darker and longer than the rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 735
734' Sternum normally wider than base of labium; cheliceral teeth variable . . . . . . . . . 736
735(734) Eyefield black to blackish brown; male: cephalothorax brown; abdomen light brown,
grey laterally with big white-yellow irregular spots and rows of small yellow spots; venter
greyish yellow with 3 grey longitudinal streaks; palpal organ orange-brown with a broad
tibial apophysis pointed outward (Fig. 1437); female: cephalothorax orange; abdomen
with straight longitudinal bands and 2 pairs of rectangular spots posteriorly (Fig. 1438);
tibia I with 4 pairs of ventral spines; epigynum with 2 oval depressions divided by a
wedge-shaped furrow above epigastric fold and coiled canals above (Fig. 1439) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maripissa magister (Karsch)
735' Entire female cephalothorax including eyefield reddish brown with black shades, ab-
dominal longitudinal band undulated in the inside, and tibia I with 3 pairs of ventral
spines; epigynum with half-starlike mark above 2 oppositely C-shaped depressions (Fig.
1440); male with lightly colored areas inside the eyefield and behind posterior lateral
eyes; abdomen with 3 light colored transverse bands; palpal organ with an acute tibial
apophysis, tapering tegulum and apically rounded cymbium (Fig. 1441) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maripissa elongata (Karsch)
736(734') Tibia + patella of leg III as long as or longer than tibia + patella of leg IV; first 2 pairs
of legs similar; tibia I with 3 pairs of ventral spines; cephalothorax high and convex, sides
parallel in the cephalic and rounded in the thoracic; ocular quadrangle occupies about
one-third carapace length, narrower in front than behind; retromargin with a single
tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 737
736' Tibia + patella of leg III shorter than tibia + patella of leg IV . . . . . . . . . . . . . . . .738
737(736) Posterior part of abdomen with 2 or 4 symmetric white-yellow spots (usually 2 large
yellow spots in female) (Fig. 1442); female eyefield black; median band broadly yellow
and black banded sublaterally, sublateral black bands run up to cephalothorax in male;
epigynum large with slit-like copulatory organs, U-shaped anterior part and oval sper-
matheca (Fig. 1443); palpal organ thick with a slightly hooked tegulum, nearly rectan-
gular bulbus and embolus in its upper part; tibial apophysis short, laterally hooked and
reaches one-fourth of cymbium; top of cymbium broadly rounded ventrally and subtrun-
cated laterally (Fig. 1444) . . . . . . . . . . . . . . . . Plexippus paykulli (Savigny & Audouin)
737' Posterior part of abdomen laterally with a mosaic of yeIlow-orange dots on a grey
background, median part of entire abdomen with an orange longitudinal band; eyefield
light brown to brown, male with dark eye margins pointed towards each other in
between posterior lateral eyes (Fig. 1445); epigynum with a dome-shaped bridge be-
tween copulatory slit-like organs (Fig. 1446); palpal organ with a rounded tegulum,
narrowly rounded top of cymbium both in ventral and lateral view . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plexippus setipes Karsch
738(736') Retromarginal tooth bifid; eyefield black; cephalothorax with a white band running
from behind of left anterior lateral eye to the sides of posterior eye transversing a little
below midcephalothorax and ends near right anterior lateral eye; patella + tibia of leg
III as long as patella + tibia of leg IV; abdomen greyish to slightly mottled with short
and brown hairs in female and black with a wide transverse crescentic band of white
across front (Fig. 1447, left) and 2 small white spots towards rear in male (Fig. 1447,
right); male without eyebrows in the eye region; epigynum very small with apically
converging oblong-shaped structures (Fig. 1448); palpal organ with very small apophysis
and embolus, tibia clearly longer than tarsus (Fig. 1449) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hasarius adansoni (Audouin)
738' Retromargin with a single tooth, cephalothorax without such band running from right to
left; anterior lateral eyes passing back of posterior lateral eyes; tibia + patella of leg III
shorter than patella + tibia of leg IV; abdomen usually with a light transverse basal
band; posterior median eyes twice as far from posterior lateral eyes as from the anterior
laterals (Fig. 1450); male with eyebrows in the eye region (Fig. 1451); epigynum
moderately large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phidippus sp.
739(730') Legs I and II usually larger and longer than III and IV, powerful and laterigrade; body
crab-like in shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740
739' All legs of the usual prograde type; body shape not crab-like . . . . . . . . . . . . . . . . . 755
740(739) Diminutive spinneret (colulus) absent; inner margin of chelicera armed with teeth; apex
of metatarsus with a soft trilobate membrane allowing hyperextension of tarsus;
posterior eye row recurved with prominent laterals; maxilla not crested; male carapace
marked with a large black nearly V-shaped band (Fig. 1452); tibial apophysis short but
with 2 teeth (Fig. 1453); female with a yellow-black bordered stripe on the posterior
slope of carapace, epigynum with lateral lobes in contact (Fig. 1454); legs very long and
spotted black [Sparassidae] . . . . . . . . . . . . . . . . . . . . Heteropoda venatoria (Linnaeus)
740' Diminutive spinneret (colulus) present; inner margin of chelicera smooth, the outer
alone with teeth; apex of metatarsus sclerotized not allowing hyperextension
[Thomisidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .741
741(740') Legs about equal in length and with true claw tufts; integument with plumose or
squamose hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .742
741' Legs I and II longer than III and IV, claw tuft simple when present; integument with
simple hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .745
742(741) Anterior lateral eyes (ALE) nearer to the anterior median eyes than to the posterior
median eyes; anterior eyes smaller and shorter in length than the posterior eyes;
posterior eyes more recurved than anterior eyes, medians much farther apart from each
other than from the lateral eyes; abdomen with a light broad median band and banded
black subdorsally, broadest in apical one-third and moderately rounded towards the tip;
legs mottled usually subbasally and subapically in all tibiae (Fig. 1455); epigynum
ovoid-shaped with a pair of elongate canals at median (Fig. 1456) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Philodromus spinitarsis Simon
742' Anterior lateral eyes closer to the posterior median eyes than to the anterior median
eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .743
743(742') Posterior eyes not strongly recurved and nearly equidistant from each other; anterior
eyes closer to the anterior median than to the posterior eyes; cephalothorax hardly
longer than wide; abdomen usually oval with a well marked cardiac area (Fig. 1457) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thanatus sp.
743' Posterior eyes strongly recurved; abdomen long, cigar-shaped and with a median lon-
gitudinal band extended to cephalothorax up to anterior median eyes; posterior part of
abdomen with or without paired spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .744
744(743') Posterior part of abdomen without paired black spots (Fig. 1458) . . . . . . . Tibellus sp.
744' Posterior part of abdomen with a pair of black spots; cardiac area with small paired
black spots within the band (Fig. 1459); ejaculatory duct of palpal organ in male with a
single slightly oblique loop at median, embolus small and acutely pointed (Fig. 1460);
epigynum with a pair of small globular rings touching epigastric furrow and separated
by a concave posterior piece (Fig. 1461) . . . . . . . . . . Tibellus oblongus (Walckenaer)
745(741') Eyes situated on distinct protuberances . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .746
745' Eye tubercles indistinct, only slightly protuberant in appearance . . . . . . . . . . . . . . .749
746(745) Abdomen widened and truncated posteriorly; eyes borne on the tubercles . . . . . .747
746' Abdomen long and laterally corrugated; eyes borne on a carina . . . . . . . . . . . . . . .748
747(746) Carapace with a median white T-shaped band and a broad longitudinal deep brown
band on each side; abdomen longer than wide and subelliptical; dorsum with 4-5
pointed black sigilla, lateral sides of abdomen with 3-4 deep brown longitudinal thin
lines (Fig. 1462); epigynum half moon-like facing each other (Fig. 1463) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thomisus cherapunjeus Tikader
747' Carapace without such pattern but with a transverse white band along posterior eyes in
the female; abdomen of male reddish brown and whitish in female, with 5 rounded
dorsal sigilla in both sexes; wider than long and semitruncated posteriorly (Fig. 1464);
epigynum inverted J-shape opposite each other (Fig. 1465); palpal organ with a laterally
projected tibial apophysis and an upwardly directed one reaching almost half of cym-
bium (Fig. 1466) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thomisus nr. labefactus Karsch
747'' Carapace without white T-band, posterolateral angle of cephalothorax rounded;
anterior and posterior portion of abdomen strongly rounded with humps laterally (Fig.
1467); leg I with a median or submedian spot in femur, tibia, and metatarsus; epigynum
simple with a globular band above epigastric furrow and transverse ovate slit at middle
(Fig. 1468) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thomisus okirawensis Strand
748(746') Carapace with a prominent, thinly white T-shaped band at middle, sides with a parallel
brown band of uniform diameter in its entire length; abdomen nearly parallel sided with
slightly rounded posterior end, median area with a pair of small longitudinal brown
bands and lateral sides with about 4 longitudinal corrugations (Fig. 1469); epigynum
with a dome-shaped median structure, basal ends loop inwards close to the epigastric
furrow (Fig. 1470) . . . . . . . . . . . . . . . . . . . . Runcinia albostriata Boesenberg & Strand
748' Carapacial T-shaped white band and broad lateral dark brown bands both prominent
but not reaching posterior margin; abdomen with 3 prominent longitudinal corruga-
tions, venter with oblique dark brown marks (Fig. 1471); epigynum marked with 2
subovate structures bridged together anteriorly by a narrow dome-shaped process (Fig.
1472) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Runcinia sp.
748'' Carapace with an indistinct T-band, lateral sides with a posteriorly widened dark brown
band; abdomen tapers posteriorly, median area with a long broad pigmented discon-
tinuous line from the base to the tip, lateral sides with conspicuous muscular corruga-
tions and elevated ridges each provided with a row of spines (Fig. 1473); epigynum with
a transversely C-shaped structure at middle (Fig. 1474) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Runcinia roortwali Tikader
749(745') Cephalic region as wide as the thoracic region; squarish cephalothorax red, high and
clothed with black hairs; anterior and lateral parts deep brown to black, median area
also deep brown but not in the male; globular abdomen with a longitudinal, chalk-white
anchor-like band at middle and 2 transverse white bands medio-laterally (Fig. 1475) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camaricus formosus Thorell
749' Cephalic region narrower than the thoracic region . . . . . . . . . . . . . . . . . . . . . . . . . 750
Figs. 1445-1461
750(749') Inner half of femur I slightly swollen and narrowed in the basal half, 2-3 spines present
near the swollen area; tibia with 5 pairs of ventral spines; posterior one-fourth of
abdomen transversely lined with clavate setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 751
750' Femur I normally shaped, posterior one-fourth of abdomen without clavate setae . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 752
751(750) Abdomen parallel-sided except in the posterior one-fourth, median area with a lon-
gitudinal broad dark band; posterolateral corner of cephalothorax with clavate setae
(Fig. 1476); tip of tibial apophysis subtruncate (Fig. 1477); chelicera with 4 outer and 3
inner teeth and 4 minute ones in between the 2 rows (Fig. 1478) . . . . . . Regillus sp. A
751' Abdomen not parallel-sided and median area without a broad dark band; posterolateral
corner of cephalothorax without clavate setae (Fig. 1479); tip of tibial apophysis
rounded (Fig. 1480); chelicera with 4 teeth each in the outer and inner row (Fig. 1481)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Regillus sp. B
752(750') Tubercles of lateral eyes broadly separated, anterior eyes slightly recurved and
posterior row strongly recurved; carapace flat above with 2 subparallel, broad dark
bands running from posterior lateral eyes to the basal one-eighth of cephalothorax in
female, U-shaped in the male; abdomen with 2 parallel dark brown bands in female (Fig.
1482) and 5-6 transverse posterior bands in male (Fig. 1483) . . . . . . . . . . . Xysticus sp.
752' Tubercles of lateral eyes confluent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 753
753(752') Anterior eyes similar in size and nearly equidistant; carapace unmarked; abdomen with
moderate longitudinal striations laterally and slightly pointed posteriorly (Fig. 1484) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Misumena sp.
753' Anterior lateral eyes larger than the anterior medians . . . . . . . . . . . . . . . . . . . . . . . 754
754(753') Abdomen with an inverted V-shaped reddish brown band anteriorly and 3-4 thin
transverse reddish brown stripes posteriorly, cephalothorax green similar to femur I-IV
and almost entire leg III and IV in female (Fig. 1485); epigynum subovate with a
transverse C-shaped median mark (Fig. 1486); male cephalothorax reddish brown with
a pair of longitudinally broad dark reddish brown bands, abdomen with an ovoid green
band at center and a brown spot each opposite its broadest part (Fig. 1487); tibia of palp
with a bifurcated retrolateral and inwardly curved ventral apophysis (Fig. 1488) ......
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Misumenops tricuspidatus (Fabricius)
754' Abdomen devoid of an inverted V-shaped band but with 6-8 globular black spots
anteriorly and 2 oblique dark bands posteriorly; cephalothorax with dark patches
laterally and posteriorly (Fig. 1489) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Misumenops sp.
755(739') Anterior spinnerets cylindrical and separated by a distance about equal to the diameter
of one; maxilla with oblique depression; eyes heterogenous, anterior medians dark and
normal in position; posterior medians usually oblique [Gnaphosidae] . . . . . . . . . 756
755' Anterior spinnerets conical and contiguous; maxilla without oblique or transverse
depression; eyes homogeneous [Clubionidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 758
756(755) Tibia IVwith 2 or more median dorsal spines; eye rows well separated, posterior median
eyes nearer each other than to the laterals and oblique; abdomen ovoid (Fig. 1490);
sternum posteriorly pointed with labium and maxilla rounded apically (Fig. 1491) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drassodes sp.
756' Tibia IV with 0-1 median dorsal spine; eyes close together with normal posterior
medians and recurved posterior eye row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 757
757(756') Abdomen with a white narrow transverse band below the submedian pair of brown spots
and about 4 thin concave dark stripes posteriorly (Fig. 1492); ventro-posterior part with
a transverse subundulate white stripe above spinnerets (Fig. 1493); epigynum triangular
with a pair of elongated, unevenly shaped structures (Fig. 1494) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilochroa sp. A
757' Abdomen with a broad transverse white band dorsosubmedially below the second pair
of spots and posteroventrally above spinnerets (Fig. 1495); epigynum with a pair of small
ovoid and diverging structures at middle close to epigastric furrow (Fig. 1496) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilochroa sp. B
758(755') Thoracic fovea absent, leg I the longest; posterior spinnerets longer than the anterior .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 759
758' Thoracic fovea present; leg IV the longest; anterior spinnerets almost as long as the
posterior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763
759(758) Abdomen naturally ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 760
759' Abdomen elongate-ovate or slightly narrowed in apical one-fourth with sides of
cephalic region produced to a small point; cardiac area very distinct . . . . . . . . . . 761
760(759) Apices of palp, tarsus and metatarsus blackish (Fig. 1497); epigynum with a knob-like
cavity at center emanating from epigastric furrow (Fig. 1498); palpal organ with a
slender and apically rounded cymbium, tarsal apophysis long with the inwardly angled-
curve part sharply pointed, tibial apophysis single but bifurcated at tip (Fig. 1499);
female produces a regularly looped shelter in a rice leaf (Fig. 1500) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiracanthium japonicum Boesenberg & Strand
760' Apices of leg segments unmarked (Fig. 1501); epigynum with a pair of moderately
robust C-shaped structures facing each other above epigastric furrow, dorsal portion
clearly separated (Fig. 1502) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiracanthium sp. A
761(759') Cephalic region moderately developed along lateral sides (raised as viewed dorsally);
abdomen rounded towards the posterior but with subparallel lateral margins in the
basal one-fourth (Fig. 1503); tarsal apophysis sickle-shaped, long and extended beyond
diameter of tibia, tibial apophysis single but minutely bifurcated at tip (Fig. 1504); inner
margin of chelicera with 5 teeth (6 in some spiders and the most basal), apical one the
largest (Fig. 1505) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiracanthium sp. B
761' Cephalic region not elevated laterally, protrusion absent . . . . . . . . . . . . . . . . . . . . . . . . 762
762(761') Posterior spinnerets noticeably long; lateral sides of cephalothorax with some reddish
brown patches (Fig. 1506); chelicera without subapical lateral tooth; abdomen elongate
ovate; epigynum lip-like with black bands laterally (Fig. 1507); tarsal apophysis short
and sharply pointed downwards, tibial apophysis single, reddish dark brown and
rounded at tip (Fig. 1508) . . . . . . . . . Chiracanthium nr. unicum Boesenberg & Strand
762' Posterior spinnerets relatively long; cephalothorax unmarked; abdomen widest a little
above midhalf and equally rounded anteriorly and posteriorly (Fig. 1509); chelicera
with a subapical lateral tooth (Fig. 1510); tarsal apophysis rather broad and pointed at
tip; apex of tibia with a small ventral apophysis and a slightly inwardly curved tip of
retrolateral apophysis (Fig. 1511) . . . . . . . . . . . . . . . . . . . . . . . . . . Chiracanthium sp. C
762" Posterior spinnerets moderately exserted; cephalothorax uniformly brownish, darker in
the cephalic region and lightly colored in the thoracic part (Fig. 1512); a female spider
rests inside a folded rice leaf looped 3 times in a diagonal pattern (see arrow); epigynum
with a transverse elongate mark adjacent to the epigastric furrow, and with 4 additional
pale marks, 2 globular dorsally and 2 ovoid laterally (Fig. 1513); tarsal apophysis of male
sharply pointed inwards but without angled corner in its entire length (Fig. 1514) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiracanthium lascivum Karsch
763(758') Abdominal dorsum mottled with greyish to black spots forming a transverse line; legs
with greyish to brown bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 764
763' Abdomen and legs unmarked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 765
764(763) Carapace with 2 long lateral longitudinal bands and 3 short bands in the cephalic area
almost converging along fovea; abdomen with at least 5 concave interrupted bands (Fig.
1515); legs with dark bands near apex of femur, both ends of tibia, and apex of
metatarsus (Fig. 1516) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clubiona sp. A
764' Carapace without lateral longitudinal bands; abdomen with 7 convex interrupted bands
(Fig. 1517); legs without bands; epigynum with 2 semi S-shaped structures facing each
other, very broad to subglobular basally towards the epigastric furrow and diverging,
normally shaped towards apex and converging, and in contact at middle (Fig. 1518);
palpal organ with an outwardly curved embolus (Fig. 1519) . . . Clubiona vigil Karsch
765(763') Chelicera blackish brown to dark reddish brown with 2 promarginal and 4-5 retromar-
ginal teeth; abdomen of female usually oblong with largely rounded anterior and
posterior ends (Fig. 1520); epigynum with a pair of spherical structures anteriorly and
paired elongate process posteriorly (Fig. 1521); palpal organ with a sickle-shaped
embolus, cymbium laterally rounded at apex; and a bifurcated tibial apophysis
(Fig. 1522) . . . . . . . . . . . . . . . . . . . . . . . . . . . Clubiona japonicola Boesenberg & Strand
765' Chelicera usually brown to light brown; female abdomen usually smaller than above;
epigynum and palpal organ differently structured . . . . . . . . . . . . . . . . . . . . . . . . . . . 766
766(765') Abdomen shortly ovoid; E. Asian species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 767
746' Abdomen moderately long to very long; S. E. Asian species . . . . . . . . . . . . . . . . . . . 768
767(766) Carapace reddish brown; abdomen brown with silky white hairs; femora of legs with 2
dorsal setae (Fig. 1523); epigynum with a pair of circular structures at middle and
half-moon like black patches on each side, posterior margin double-notched (Fig. 1524);
palpal organ with a short concealed embolus, cymbium slightly pointed at tip and with a
depression below it viewed laterally, ventral tibial apophyses much longer than the
retrolateral (Fig. 1525) . . . . . . . . . . . . . . . . . . . . . Clubiona lena Boesenberg & Strand
767' Carapace light brown; abdomen yellowish brown; femur of legs with 3 dorsal setae (Fig.
1526); epigynum simple, midposterior area closely bilobed with a narrow cleft in be-
tween (Fig. 1527); apex of tibia of palpal organ with 2 apophyses nearly converging,
ventral apophysis long and bifurcated reaching midhalf of cymbium (Fig. 1528) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clubiona kurilensis Boesenberg & Strand
768(766') Lateral side of abdomen with a small tubercle subapically and with some moderately
long setae dorsomedially; abdomen 1.4 times longer than cephalothorax (Fig. 1529);
chelicera with 6 promarginal and 4 retromarginal teeth (Fig. 1530) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clubiona sp. B
768' Abdomen long, more than 1.6 times longer than cephalothorax and without lateral
swellings subapically (Fig. 1531); epigynum broadly U-shaped with slightly converging
arms well rounded apically, median with a pair of J-like processes opposite each other
(Fig. 1532); tibia of palpal organ with a short and broad retrolateral apophysis and 2
long setae at median length (Fig. 1533) . . . . . . . . . . . . . . . . . . . . . . . . . . Clubiona sp. C
769(729') All legs conspicuously spinose; eye group in a hexagonal pattern, anterior median eyes
very small; clypeus usually high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 770
769' Legs not conspicuously spinose; eyes never arranged hexagonally . . . . . . . . . . . . . . 773
770(769) Abdomen with 2 blackish red sublateral ( = subdorsal) longitudinal bands similar to the
carapace; hexagonal eyes very close to each other (Fig. 1534); epigynum with a dome-
shaped median process (Fig. 1535); palpal organ with a long cymbium, apophysis on
tibia blunt and hardly protruded outside (Fig. 1536) . . . Oxyopes macilentus L. Koch
770' Abdomen with oblique sublateral bands; hexagonal eyes not very close to each other .
........................................................................ 771
771(770') Tibial apophysis of palpal organ very long; cymbium moderately long; embolus curved
downward (Fig. 1537); epigynum with a moderately broad median protruding process
posteriorly, arms subparallel anteriorly (Fig. 1538) . . . . . . . . Oxyopes javanus Thorell
Figs 1507-1519
771' Tibial apophysis of palpal organ without a long process; epigynum with different
structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 772
772(771') Epigynum widely cleft posteromedially with a pair of small cone-like structures
anteriorly (Fig. 1539); embolus short and hooked moderately downwards; cymbium
slightly short at apex; apex of tibia with 2 small and short apophyses (Fig. 1540) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxyopes sertatus L. Koch
772' Epigynum strongly concave with a pair of ball-like structures anteriorly (Fig. 1541);
embolar area with 2 slender and a large apically truncate processes; cymbium with a
slightly slender and acute apex, base opposite apex of tibia cleft producing 2 tubercles;
apex of tibia with a blunt and semi-rounded lateral process (Fig. 1542) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxyopes Iinealipes (C. L. Koch)
773(769') Six spinnerets in a transverse row (Fig. 1543) and tracheal spiracles closer to the
spinnerets than to the epigastric fold; abdomen with 3 black longitudinal bands inter-
connected on both ends and 3-4 transverse narrow bands posteriorly (Fig. 1544);
anterior median eyes smaller than anterior laterals; posterior eye row strongly
procurved (Fig. 1545); chelicerae with 2 promarginal and retromarginal teeth each (Fig.
1546)[Hahniidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hahnia sp.
773' Spinnerets not in a transverse line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 774
774(773') Tarsus IV with a comb of serrated setae (Fig. 1547) [Theridiidae] . . . . . . . . . . . . . 775
774 ' Tarsus IV without a serrated comb of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 782
775(774) Cephalothorax with a transverse groove in the middle; abdomen usually prolonged
either above (with triangular appearance) or elongated behind the spinnerets; cephalic
area of male usually raised . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 776
775 ' Cephalothorax without transverse groove; abdomen subglobular or slightly prolonged
....................................................... ................ 780
776(775) Abdomen very long and slender, much prolonged behind the spinnerets and tapers
posteriorly; carapace with longitudinal brown bands on each side of the elevated foveal
area (Fig. 1548); projection in the male head moderately knob-like and without eyes
viewed laterally (Fig. 1549); legs long and slender with alternate transparent (colorless)
and yellow band (Fig. 1548) . . . . . . . . . . . . . . . Argyrodes saganus (Doenitz & Strand)
776' Abdomen more or less subtriangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777
777(776') Abdomen orange-red with a black spot posteroventrally around spinnerets and on top
posterodorsally, 4 silvery lateral spots present just below dorsal black spot; margin
between 2 black spots almost straight (Fig. 1550); tibia I a little shorter than metatarsus
I; median part of epigynum with a pair of diverging horn-like processes directed and
overhanging expanded ear-like structures in the lateral sides (Fig. 1551); laterally, male
horn on head with 2 eyes, anterior eye slightly bigger than the posterior and deeply
concave to U-shaped from base of cephalic region (Fig. 1552); parategular apophysis
shortly curved and with a middle tooth (Fig. 1553) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrodes miniaceus (Doleschall)
777' Abdomen mostly silvery, subtriangular, and pointed posterodorsally . . . . . . . . . . 778
778(777') Venter and basolateral area of abdomen yellowish brown; in lateral view, posterodorsal
tip of abdomen in an oblique position from the spinnerets, the latter almost at midhalf
from pedicel to posterodorsal end (Fig. 1554); tibia I much longer than metatarsus I;
epigynum truncated anteriorly and concaved posteriorly with a bilobed structure at
middle (Fig. 1555); laterally, male horn V-shaped from base of cephalic region, with 2
eyes of equal diameter (Fig. 1556); paracymbium narrow medially and slightly swollen
apically, median apophysis triangular (Fig. 1557) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrodes fissifrons O.P. -Cambridge
778' Venter and basolateral part of abdomen black, rest uniformly silvery . . . . . . . . . . 779
774(778') Abdomen tapers and pointed upward towards the posterior (Fig. 1558); cephalic area
of male deeply cleft or V-shaped in lateral view, horn rounded anteriorly with 2 eyes of
different diameter, anterior eye larger than the posterior (Fig. 1559); tibia I shorter than
metatarsus I; epigynum half circular anteriorly with a subquadrate plate posteriorly
containing a pair of globular structures inside (Fig. 1560); palpal organ with a large
apophysis anteriorly and transverse tibia (Fig. 1561) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrodes bonadea (Karsch)
779' Abdomen more or less rounded (half ovate) posterodorsally in lateral view; tibia I
longer than metatarsus I (Fig. 1562); epigynum with a tongue-like median structure
(Fig. 1563) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrodes sp.
780(775') Cephalothorax, legs and abdomen light green; abdomen with 4 pairs of black spots
dorsally in the female (Fig. 1564) and 3 pairs in the male (Fig. 1565); epigynum a simple
plate with rounded anterior margin and truncate posterior margin, middle with a double
hook-like process (Fig. 1566); tibia of palpal organ without a neck basally, tegular
apophysis slender and pointed at tip, cymbium rounded on top (Fig. 1567) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Theridion octomaculatum (Boesenberg & Strand)
780' Body and leg coloration yellowish brownish to black or reddish brown; abdomen
without paired black spots, usually rounded or with a small hump or protrusion
posterodorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 781
781(780') Carapace with a broad grey-brown longitudinal band trifurcated towards lateral eyes;
abdomen subglobular, dorsally blackish brown interrupted medially with silvery marks
along cardiac area and laterally with 2 indentions (Fig. 1568); chelicera with an upward-
ly and laterally projected spine (Fig. 1569); epigynum with a transverse lace-like median
structure (Fig. 1570); cymbium bifurcated towards apex, tibia almost 2 times longer than
wide (Fig. 1571) . . . . . . . . . . . . . . . . . . Enoplognatha japonica (Boesenberg & Strand)
781' Carapace without any band; abdomen dorsally ovoid to moderately elongate-ovate,
posterodorsally with a protrusion in female and a small hump in male viewed laterally
(Fig. 1572); anterior part of male abdomen cleft; epigynum with a pair of subparallel
black bands opposite the indention of the posterior margin, median piece Y-shaped
(Fig. 1573); cymbium rounded apically, tibia as long as wide apically, embolus long,
curved, and with a broad base (Fig. 1574) . . . Coleosoma blandum O. P. -Cambridge
782(774') Tarsus IV with very fine hairs (trichobothria) arising at a 90° angle from leg (Fig. 1575);
trochanters with or without notch; lip not rebordered . . . . . . . . . . . . . . . . . . . . . . . 783
782' Tarsus IV without trichobothria; trochanters without notch; lip usually rebordered . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .809
783(782) Trochanters without notch; tarsus with a single row of trichobothria and usually 3-
clawed; terminal segment of posterior spinnerets visible above; longer than basal seg-
ment; abdomen with irregularly straight transverse light bands interrupted in cardiac
area; carapace with 2 reddish brown bands, broad in the thoracic area and narrowed to
a point towards posterior lateral eyes (Fig. 1576); epigynum with a transverse orifice
anteriorly divided medially by a longitudinal line (Fig. 1577); tibia of palpal organ with
a moderately acute apical apophysis; cymbium swollen subbasally and slender apically
(Fig. 1578) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agelena labyrinthica (Clerck)
783' Trochanters with a notch; tarsus with numerous trichobothria . . . . . . . . . . . . . . . . .784
784(783') Median claw of tarsus with 2-3 teeth; lower margin of cheliceral furrow with 3-4 teeth;
posterior eye row not forming 2 distinct rows and not strongly recurved; anterior piece
of lorum of pedicel with a notch into which the posterior part fits [Pisauridae) . . .785
784' Median claw smooth or with a single tooth; posterior eye row strongly recurved forming
2 rows; anterior piece of lorum rounded behind and fits into a notch of the posterior part
[Lycosidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 787
785(784) Lower margin of cheliceral furrow with 3 teeth; carapace with a relatively broad median
longitudinal white band and 2 dark brown to reddish brown longitudinal bands laterally
all extended up to the entire length of abdomen (Fig. 1579); sternum yellow media-lon-
gitudinally and with 6 ovoid yellow spots along sides (Fig. 1580); epigynum anchor-like
with rounded tip of lateral arms and broadened body base (Fig. 1581); palpal organ with
a knife-like retrolateral apophysis hairy on the back side (Fig. 1582) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pisauru lama Boesenberg & Strand
785' Lower margin of cheliceral furrow with 4 teeth; middorsum of carapace and abdomen
with a broad brown band followed by a narrow white band on each side (Fig. 1583) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 786
786(785') Epigynum bulb-like, broad below midhalf but almost truncated in the apical margin
provided with elongated patches off each side, posterior epigynal margin not cleft
medially (Fig. 1584) . . . . . . . . . . . . . . . . . . . . . . . . . . Dolomedes angustivirgatus Kishida
786' Epigynum tapers to a rounded anterior margin, posterior epigynal margin with 2 clefts
producing a small tongue-like process at center (Fig. 1585); apophysis of tibia bluntly
truncate in the retrolateral and small but pointed in the ventrolateral, cavity of cymbium
rounded apically below embolus (Fig. 1586) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dolomedes sulfureus Boesenberg & Strand
787(784') Posterior spinnerets distinctly longer than the anterior, apical segment conical and as
long as basal; anterior eye row a little longer than the posterior row; sternum with a
median longitudinal black band (Fig. 1587) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 788
787' Posterior spinnerets slightly longer than the anterior and apical segment hemispherical
and very short; anterior eye shorter than the posterior row; sternum without a median
longitudinal black band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 789
788(787) Small spiders, total length 9 mm; epigynum bifurcated posteriorly, showing a W-shaped
appearance (Fig. 1588); anterior eye row straight; labium longer than wide; distal end of
maxilla with distinct scopulae; cephalothorax clothed with line hairs and spine-like hairs
in the lateral margins; abdomen pale brown with transverse patches dorsomedially,
subtriangular light band opposite tip of anterior median longitudinal brown band
(Fig. 1589); palpal organ with a slender apical half of cymbium (Fig. 1590) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hippasa greenalliae (Blackwall)
788' Total length less than 9 mm; epigynum with a subtruncate tongue-like structure along
the posterior margin (Fig. 1591); anterior eye row slightly procurved; labium wider than
long; distal end of maxilla without distinct scopulae; cephalothorax with pubescence,
without spine-like hairs in the lateral margins; abdomen strongly chequered, anterior
median longitudinally reddish brown followed by pairs of large and less regular lateral
grey brown spots (Fig. 1592); palpal organ with a rather short apical part of the cymbium
(Fig. 1593) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hippasa holmerae Thorell
789(787') Metatarsus IV longer than or as long as tibia + patella together; labium wider than long;
clypeus vertical viewed laterally; cephalic region not much elevated from thoracic
region; superior claw entirely toothed; carapace usually with median band . . . . . 790
789' Metatarsus IV shorter than tibia + patella combined; clypeus may or may not be
slanting viewed frontally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 799
790(789) East Asian species of Pardosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 791
790' South and Southeast Asian species of Pardosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 794
791(790) Anterior part of the median fight band as broad as the distance between the posterior
lateral eyes, tapers posteriorly and constricted a little above fovea; head with a truncated
area below anterior eye row; chelicera with a thin longitudinal band basally (Fig. 1594);
abdomen more lightly colored than carapace with semirectangular to hat-like marks
dorsally towards the posterior (Fig. 1595); epigynum cleft posteromedially between 2
lateral openings and bears 2 oblique structures at middle (Fig. 1596); cymbium with 2
teeth at apex; median apophysis broad but short and rounded apically ventrally and
truncate laterally; terminal apophysis slightly hooked (Fig. 1597) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa laura (Karsch)
791' Median light band of carapace widens towards posterior lateral eye (PLE) row, as wide
as or wider than the distance from PLE-PLE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 792
792(791') Median band narrow and slightly trifurcated cephalad, outer arms curved to the lateral
sides of each posterior lateral eye and the median almost traversing mid-PLE distance,
submargins with narrower bands; abdomen with transverse elongate marks below the
tower-like light band in the cardiac area (Fig. 1598); front of head strongly recurved
below anterior eye row, each chelicera with a short basolateral longitudinal thin band
(Fig. 1599); epigynum with a very broad median piece posteriorly and constricted
before anterior margin (Fig. 1600); median apophysis longer than wide, narrowed
before a rounded apex, terminal one bifurcate at apex (Fig. 1601) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa palustris (Linnaeus)
792' Median band not trifid apically behind posterior eye row; abdominal dorsum with light
colored rectangular or paired to transverse elongate bands below cardiac area . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793
793(792') Dorsum of abdomen with about 3-4 transversely elongated light bands posteriorly and a
pair of ovoid light bands on the side of the tapering posterior end of the cardiac area;
median light band of the carapace slightly T-shaped, broad below posterior eyes and
narrowed posteriorly (Fig. 1602); clypeus high, more than 1 anterior median eye
diameter, and concave medially, chelicera without distinct longitudinal bands frontally
(Fig. 1603); epigynum with a narrow truncated structure at midposterior margin, widens
towards the inside orifice becoming dome-shaped subanteriorly (Fig. 1604); median
apophysis sickle-shaped with a rounded tip, tip of cymbium without spine (Fig. 1605) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa astrigera L. Koch
793' Dorsum of abdomen with 3-4 transversely rectangular light bands opposite posterior
end of cardiac area: median light band of carapace not T-shaped (Fig. 1606); clypeus
high, almost 2 anterior median eye diameters and recurved, chelicera with 2 parallel
longitudinal bands dorsomedially (Fig. 1607); epigynum with a truncated plate slightly
beyond posterior epigynal margin and a pair of oblique orifices towards the anterior
margin (Fig. 1608); tip of cymbium with 3 teeth, median apophysis broadly rounded
apically with a small hook at lower midhalf, terminal apophysis small and lanceolate-like
(Fig. 1609) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa isago Tanaka
794(790') Median light band of carapace prominently forked or Y-shaped anteriorly, sublateral
margins with a longitudinal white band; abdomen with 4-5 transverse light bands
dorsally (Fig. 1610) in the male and 3 elongate-ovate light bands and a pair of globular
light spots dorsally in the female (Fig. 1611); epigynal lateral margin indented inside
before posterior end, guide slender and enlarged apically toward posterior epigynal
margin (Fig. 1612); tip of cymbium without teeth, terminal apophysis downwardly
sickle-shaped, embolus noticeable (Fig. 1613) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa pseudoannulata (Boesenberg & Strand)
794' Dorsum of carapace without a Y-shaped or forked anterior median band . . . . . . . 795
795(794') Median and submarginal bands indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 796
795' Median and submarginal bands distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 797
796(795) Ocular area covered by posterior eye blackish brown made to an inverted U-shaped
light band between posterior lateral eyes; anterior eyes short and straight; sides of
cephalic area lightly colored (Fig. 1614); abdomen clothed with fine hairs, dorsum
pigmented brown with pale dots; tibia and metatarsus of legs I and II with 4 and 3 pairs
of ventral spines; epigynum concave anteriorly with a pair of sclerotized oblique C-
shaped cavities at center (Fig. 1615); palpal tibia long about 0.5 times of cymbium;
terminal apophysis sharply pointed (Fig. 1616) . . . . . . . Pardosa sutherlandi (Gravely)
796' Ocular area uniformly dark without an inverted U-shaped mark; sides of cephalic area
blackish brown; anterior eyes slightly procurved; abdomen with 3 pairs of subglobular
yellow spots and 3-4 transversely irregular light bands (Fig. 1617); epigynum with a
rounded to angulate-truncate anterior margin, guide body very narrow only about 0.5
times diameter of guide pocket (Fig. 1618); palp with a white tibia and black tarsus in
male (Fig. 1619) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa birmanica Simon
797(795') Abdomen clothed with minute light spots all over dorsally; ocular area black and hairy;
light median band on carapace tapers posteriorly, constricted above fovea and bifur-
cated in between posterior lateral eyes (Fig. 1620); submarginal band not very wide;
epigynum strongly rounded in the anterior margin, indented posteriorly and bears an
expanded tongue-like median plate (Fig. 1621); maxilla with inconspicuous scopulae .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa annandalei (Gravely)
797' Abdomen with paired lens-like spots; median light band extended inside the ocular area
forming an inverted U or partly V-shaped band in the cephalic area; submarginal band
prominently wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 798
798(797') Cephalothorax brown; anterior end of light median band of carapace with a pair of small
ovate light spots between the posterior lateral eyes (PLE) (Fig. 1622), or apex becoming
pointed in between PLE to PME and between posterior median eyes (PME) (Fig.
1623); abdomen dark brown with transverse paired subglobular spots dorsally; apex of
the median piece (guide) inverted T-shaped with arms hooked inward-upward (Fig.
1624); maxilla with conspicuous scopulae . . . . . . . . . . . . Pardosa sumatrana (Thorell)
798' Cephalothorax yellowish brown, median light band widens above fovea and below
posterior lateral eyes, black bases of lateral eye darker than submedian bands (Fig.
1625); abdomen yellowish brown, mid-anterodorsal area with a lens-like marking fol-
lowed behind by a pale band, sides with 2 longitudinal rows of brownish black bands
from base to end; epigynum with a pair of subparallel sclerotized structures anteriorly
and a pair of bibulbous structures along posterior epigynal margin (Fig. 1626) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pardosa oakleyi Gravely
799(789') Carapace without a V-shaped marking above fovea; labium longer than wide . . . 800
799' Carapace with a prominent V-shaped marking above fovea; labium as long as wide to
longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 804
800(799) Tibia IV with 2 equally stout dorsal spines (Fig. 1627); anterior eye row straight to
slightly procurved, anterior median eyes (AME) never smaller than anterior lateral eyes
(ALE) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 801
800' Tibia IV with the dorsal proximal spine thinner or more drawn out than the distal one
and sometimes reduced to a bristle (Fig. 1628) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 802
801(800) Median light band of carapace broad extended up to straight anterior median eyes,
submedian dark bands similarly broad running from posterior lateral eyes and almost
converged at base of carapace; abdomen longer than carapace, provided with reddish
brown spots and transverse light brown patches posterodorsally (Fig. 1629); epigynum
strongly rounded along anterior margin, posteriorly with 3 indentions, midcleft deeply
Y-shaped (Fig. 1630); median apophysis almost rounded at tip, apex of cymbium slightly
tapering (Fig. 1631); chelicera of male without dorsal tooth in the fang . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lycosa chaperi Simon
801' Median band of carapace reaching the area in between posterior median eyes (Fig.
1632); anterior eye row slightly recurved; submedian dark band open basally; abdomen
shorter than carapace provided with irregular light and dark colored patches; fang of
chelicera with a dorsal tooth almost at midlength (Fig. 1633) . . . . . . . . . . . . Lycosa sp.
802(800') Carapace glabrous without definite longitudinal light band; cephalothorax reddish
brown with head region not so distinct; anterior eye row recurved, medians twice as
large as the laterals; sternum orange-brown; abdomen black with 3 pairs of brown dots
dorsally and light gray ventrally, and shorter than carapace; tarsus I with dorsobasal
bristle drawn out thin and fine at the end; legs with transverse bands (Fig. 1634);
epigynum broadly subtriangular apically and slightly procurved posteriorly; median with
a pair of oppositely faced sclerotized areas (Fig. 1635) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctosa janetscheki Buchar
802' Carapace hairy with a median light band; tarsus I without a long and thin dorsobasal
bristle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 803
803(802') Abdomen with a pair of light yellow longitudinal bands in the anterior half; cephalic
area more lightly colored than the thoracic region; eye area without black bands
(Fig. 1636); maxilla yellowish towards scopulae and brownish basally (Fig. 1637);
epigynum semitriangular apically with a pair of light spots and subparallel trunk of
median guide (Fig. 1638) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trochosa sp. A
803' Abdomen with transverse light bands usually with a pair of hairs in each cell; submedian
carapacid band prominently dark colored similar to the U-shaped band of posterior
eyes (Fig. 1639); maxilla uniformly light colored (Fig. 1640); epigynum with an anchor-
like median guide producing 2 distinct clefts in the posterior margin, lateral margins
each with an inwardly directed sclerotized hook-like process (Fig. 1641) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trochosa sp. B
804(799') Anterior median eyes (AME) larger than anterior lateral eyes (ALE) . . . . . . . . . 805
804' AME smaller than or equal to ALE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 808
805(804) Anterior eye row as long as posterior median row; carapacial light lateral bands not
separated from margins by dark brown bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 806
805' Anterior eye row shorter than posterior median row; carapace with a narrow dark
brown line along margin; dark brown hands separate light lateral bands from margins in
the carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 807
806(805) Height of clypeus less than 0.5 diameter of AME; AME diameter about twice as large
as anterior lateral eyes (Fig. 1642); V-shaped mark of carapace greyish brown; lateral
sides of abdomen yellowish brown with blackish grey markings (Fig. 1643); legs grey-
brown with indistinct annulations in all femora, metatarsus I with 2 ventral apical spines;
epigynum with paired spatulate processes posteriorly and indistinct stalked sper-
mathecae in apical margin (Fig. 1644); median apophysis with a long and slender margin
and a small process at apex (Fig. 1645) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirata subpiraticus (Boesenberg & Strand)
806' Height of clypeus a little shorter than diameter of AME; AME about 1.5 times as large
as ALE; V-shaped mark reddish brown; lateral sides of abdomen reddish brown with
white pubescenee (Fig. 1646); legs dark yellowish brown, metatarsus I with 3 ventral
apical spines, epigynum with trilobe-like structures at lateral sides, paired spermathecae
spatulate and bottle-shaped (Fig. 1647); median apophysis obtuse with a pointed
process (Fig. 1648) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirata piraticus (Clerck)
807(805') Height of clypeus as wide as AME diameter; AME moderately larger than ALE (Fig.
1649); V-Shaped band blackish brown; abdomen blackish grey with 3 pairs of white
spots, a triangular one prominent medially (Fig. 1650); epigynum with a pair of oval
processes creating a deep median cleft along posterior margin (Fig. 1651); median
apophysis robust with a small dorsal projection (Fig. 1652) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirata meridionalis Tanaka
807' Height of clypeus less than half of AME diameter; AME twice as large as ALE (Fig.
1653); V-shaped band brown; abdomen yellowish brown with blackish grey markings,
light transverse bands with acute and raised medians prominent dorsally (Fig. 1654);
epigynum with paired posterolaterally inverted V-shaped structures (Fig. 1655), sper-
mathecae long and erect (Fig. 1656), slightly diverging apically; robust median
apophysis with long apical projection (Fig. 1657), retrolaterally it is wide at the upper
part with one projection tapered at end on ventral margin (Fig. 1658) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirata piratoides (Boesenberg & Strand)
808(804') Anterior eye row strongly procurved; height of clypeus equal to 1 AME diameter;
ME-AME and AME-ALE separation as long as 1 AME diameter (Fig. 1659);
carapace with a very narrow dark brown line along margin; abdomen blackish grey
dorsally with light transverse bands radiating from median longitudinal area (Fig. 1660);
epigynum with a pair of slightly oblique button-like processes provided with 3 pairs of
indistinct round structures, anterior epigynal margin strongly rounded and medially
cleft along posterior margin (Fig. 1661) . . . . Pirata procurvus (Boesenberg & Strand)
808' Anterior eye row weakly procurved; height of clypeus 1.5 times AME diameter; AME
closer to each other than to ALE (Fig. 1662); carapace without lines marginally;
abdomen blackish grey with broad yellow-brown bands medio-dorsally (Fig. 1663);
epigynum with a pair of prominent brown globular structures, median posterior
epigynal margin deeply cleft, anterior margin rounded (Fig. 1664); robust median
apophysis with a long and slender projection upward (Fig. 1665); retrolaterally sharp at
apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirata clercki (Boesenberg & Strand)
809(782') Chelicera with stridulating ridges (Fig. 1666); paracymbium of male a separate sclerite
(Fig. 1667) [Linyphiidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 810
809' Chelicera without stridulating ridges; paracymbium fused with cymbium (Fig. 1668); not
a separate sclerite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 814
810(809) Lateral margins of carapace serrated (Fig. 1669), anterior part of chelicera, femur of
palpal organ maxilla with teeth; carapace glossy and dark brown or red-brown with
darker cervical groove and radial striae; head raised without definite lobe; chelicera
armed with a large pointed boss anteriorly and with a row of 6-7 warty teeth antero-
laterally (Fig. 1670); sternum black; tibia I-III with 2 dorsal spines and IV with one;
position of trichobothrium in metatarus IV 0.42-0.5; metatarsus longer than tarsi, IV
without trichobothrium; epigynum prominently lobe-like and armed with a small
chitinized brown tooth in the midposterior epigynal margin (Fig. 1671); femur of palpal
organ nearly as long as femur II and curved mesally; patella half of femur, distally
thickened with a long tapered apophysis ventrally; tibia slender basally with a long
ventral projection armed with a small tooth ventrally; paracymbium strongly widened
distally (Fig. 1672) . . . . . . . . . . . . . . . . . . . . Erigone prominens (Boesenberg & Strand)
810' Lateral margins of carapace not serrated; metatarsus IV with a trichobothrium; supe-
rior tarsal claws with a few minute teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 811
811(810') Position of trichobothrium in metatarsus IV 0.81; chelicera laterally convex, without
stridulating organ; cephalic lobe of male distinct with a pit opened within an ocular sulci
midway between posterior median eye to lateral (Fig. 1673); female without a cephalic
lobe; abdomen with 4 pairs of grey spots in 2 longitudinal rows (Fig. 1674); epigynum
with 2 clefts along posterior margin, each leading to an ovate diverging receptacle (Fig.
1675); palpal femur 2 times longer than patella; tibia with 3 sharp apical apophyses;
embolic division with a broad triangular scaphium; paracymbium with a small basal part,
slender but short median part and a large triangular end-piece (Fig. 1676) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atypena ( = Callitrichia) formosana (Oi)
811' Position of trichobothrium in metatarsus IV only 0.42-0.66; head may or may not be
elevated in both sexes; stridulating organ usually present; chelicera with warts or
antero-lateral granulations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 812
812(811') Head not elevated into a lobe in both sexes; eyes rather small, anterior row nearly
straight, and posterior row moderately procurved; carapace glossy and nut brown, oval
in outline dorsally and evenly rounded on sides; clypeus straight, subvertical and as wide
as ocular area; abdomen yellowish dark brown (sometimes black with 4 impressed dots
and several faint pale chevrons dorsally (Fig. 1677); position of trichobothrium in
metatarsus IV 0.6 epigynum with a large transversely elongate oval opening (Fig. 1678);
short patella thickened distally, curved ventrally and with a small ventral tooth; tibia with
a broadly truncate lateral tooth (Fig. 1679); genital bulb thin; paracymbium widened
into 2 small distal lobes (Fig. 1680) . . . . . . . . . . . Erigonidium graminicola (Sundevall)
812' Head of male moderately elevated into a lobe; eyes fairly large, sometimes closely
grouped; position of trichobothrium in metatarsus IV 0.6-0.66 . . . . . . . . . . . . . . . . . 813
813(812') Abdomen ovoid greyish yellow to black with a pale longitudinal median line dorsally;
carapace yellowish brown to dark brown (Fig. 1681); male head elevated behind eyes
with a deep transverse suture at midelevation part, female head considerably gibbous
above but without transverse suture; chelicera swollen at base with 5 promarginal teeth,
armed with a large boss in front and clothed with antero-lateral granulations in male
(Fig. 1682); boss absent in the female; position of trichobothrium in metatarsus IV
0.56-0.62; epigynum dark brown with a small orifice divided by a triangular scape (Fig.
1683); tibia as long as patella drawn distally, dorsal margin with a broad apophysis
indentated distally; embolus long and coiled at the balbal tip (Fig. 1684) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . Ummeliata ( = Oedothorax) insecticeps (Boesenberg & Strand)
813' Abdomen elongate-ovate, dull yellow with longitudinal closed black stripes on each side
of the dorsum (Fig. 1685) and on each side of the venter; carapace brown-orange with
blackish ocular area and a large hump behind eyes in the male (hump absent in the
female); hump broadly rounded in front, moderately deep transverse groove between
ocular area and hump (Fig. 1686); chelicera with a large pointed anterior boss curved
mesially and warty granulation antero-laterally; position of trichobothrium in metatar-
sus IV 0.7; epigynum with a large orifice in the lip of scape with clear spermathecae (Fig.
1687); patella twice as long as patella I with a small process on the distal end; tibia
funnel-shaped distally, dorsolaterally provided with a black long sickle-like apophysis;
bulb slender and curved embolus along the membranous semicircular pars pendula
(Fig. 1688) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gnathonarium gibberumn Oi
814(809') Tiny spiders barely 3 mm long; sternum (Fig. 1689) broadly truncate anterior part with
indentations; abdominal pattern without folium; female palp clawless
[Theridiosomatidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 815
814' Small to large spiders usually more than 5 mm long; sternum cordate; abdominal folium
prominent; female palp with a claw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 816
815(814) Scape prominently exserted beneath entire epigynal posterior margin (Fig. 1690); palpal
organ with squarish median apophysis, mesal bristle in the embolic division protruding
from beneath the conductor (Fig. 1691); carapace orange-yellow with greyish abdomen;
posterior median eye separation at least 1 PME diameter; tibial trichobothria in 2-3
irregular rows, usually moderately long . . . . . . . . . . . . . . . . . . . . . . . . . Wendilgarda sp.
815' Scape not protruding beneath epigynal posterior margin; palpal organ with the embolic
division a short, tubular embolus with apophysis consisting of separated bristle-like
parts; median apophysis slightly grooved and gradually tapered apically; carapace dark
brown to light tan; eyes nearly of similar sizes, posterior median eye separation 0.5 eye
diameter (Fig. 1692) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Theridiosoma sp.
816(814') Abdomen long and elongate (except in Dyschiriognatha with rounded abdomen);
chelicera long, usually projected forward and with various teeth arrangements; female
genital aperture not sclerotized [Tetragnathidae] . . . . . . . . . . . . . . . . . . . . . . . . . . . 817
816 Abdomen seldom elongated; chelicera short and often vertical; female genital aperture
sclerotized or partially sclerotized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 831
817(816) Abdomen globular or oval with silvery appearance . . . . . . . . . . . . . . . . . . . . . . . . . 818
817' Abdomen prominently elongated and less silvery . . . . . . . . . . . . . . . . . . . . . . . . . . . 819
818(817) Abdomen globular with 4 black spherical spots dorsally; palpal organ without long
processes above tibia (Fig. 1693); chelicera and carapace red (Fig. 1694) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Dyschiriognatha quadrimaculata Boesenberg & Strand
818' Abdomen globular with longitudinal inwardly hook; dark streaks (sometimes greyish to
black) dorsally; chelicera and carapace reddish black (Fig. 1695); palpal organ with 2
long processes above tibia (Fig. 1696) . . . . . . . . . . . . Dyschiriognatha tenera (Karsch)
818" Abdomen ovoid with a moderately broad median and 2 thin sublateral longitudinal
white bands and a broad reddish to dark brown band between the whitish median and
sublateral bands; carapace with 3 longitudinal brown stripes (Fig. 1697); sternum
punctured; labium not markedly swollen distally; proximal end of femora I and II with a
group of 2-3 trichobothria; paracymbium with a pointed lateral apophysis, rather short
not extended beyond bulb, rounded towards apex (Fig. 1698); epigynal fold simple with
transverse narrow orifice (Fig. 1699) . . . . . . . . . . . . . . . Pachygnatha clercki Sundevall
819(817') Spinnerets almost midventrally located; abdomen pointed posteriorly (Fig. 1700);
anterior lateral eyes on the margin of carapace (Fig. 1701); guide tooth far from first
large tooth in the female (Fig. 1702); apophysis in male bifurcated (Fig. 1703) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha javana (Thorell)
819' Spinnerets on the abdominal tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 820
820(819') Posterior eye row strongly recurved; posterior lateral eye far from anterior lateral eye
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 821
820' Posterior eye row gently recurved to straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824
821(820) Tarsus of palp enlarged; male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 822
821' Tarsus of palp normal, female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 823
822(821) Fang of chelicera with a subbasal outer excrescence; auxillary fang guide (Ax) bluntly
rounded at tip and slightly away from guide tooth (G); upper teeth below guide tooth of
equal size and spacing (Fig. 1704); ventrally auxillary fang guide large and close to large
guide tooth with a tubercle at middle (Fig. 1705); paracymbium bluntly rounded at tip
and conductor hook-like apically; anterior lateral eyes distinctly very small compared to
the medians (Fig. 1706) and closer to PLE than to AME . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha squamata Karsch
822' Fang of chelicera with 2 inner cusps (median and subbasal); apophysis (a) short, blunt
at tip and slightly curved inward, slanting tooth (SI) on top middle of auxillary fang guide
and first large tooth (T) (Fig. 1707); ventrally G closer to T than to Ax, paracymbium
slender towards apically rounded tip with a lateral tubercle at midlength (Fig. 1708);
distal portion of embolus and conductor truncated laterally after a small cleft (Fig.
1709); ALE smaller than medians, closer to AME than to PLE (Fig. 1710) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha virescens Okuma
823(821') Chelicera with 7 promarginal and 5 retromarginal teeth; G closer to T than to Ax
dorsally (Fig. 1711); genital fold almost parallel-sided and subtruncate apically (Fig.
1712) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha squamata Karsch
823' Chelicera with 6 promarginal and 5 retromarginal teeth; G almost midway Ax and T
dorsally (Fig. 1713); genital fold with lateral indentions before narrowly rounded apex
(Fig. 1714) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha virescens Okuma
824(820') Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 828
824' Female ............................................................... 825
825(824') Chelicera with 3 strong apical teeth, apophysis (a) with a median tubercle; retromargin
with about 12 teeth (Fig. 1715); paracymbium with an apical cleft (Fig. 1716); ALE
closer to PLE than AME, ocular quad narrower in front than behind (Fig. 1717) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha nitens (Audouin)
825' Chelicera without 3 strong apical teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 826
826(825') Chelicera with a simple apically pointed apophysis (a), tooth (t) and T absent, promar-
gin with 12 teeth, uppermost one the largest with subswollen base (Fig. 1718); retromar-
gin lined with 15-16 teeth (Fig. 1719); paracymbium narrowly bilobed at apex (Fig.
1720); distal end of embolus and conductor with a wide gap before apex (Fig. 1721);
both eye rows of equal length and slightly recurved; ocular quad slightly wider behind
(Fig. 1722) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha mandibulata Walckenaer
826' Chelicera with a bifid apophysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 827
827(826') Central ocular quadrangle nearly a square (Fig. 1723); chelicera without an outer
subbasal cusp in the fang; slanting tooth (sl) absent; upper guide (G) tooth prominent
and about half as long as T; tooth (t) rather minute; G and T space encloses 3
retromarginal teeth; promargin with 8 teeth (Fig. 1724), retromargin with 13 (Fig. 1725);
paracymbium slight curved inwards and hooked basally (Fig. 1726); distal end of
conductor and embolus pointed and curved . . . . . . . . . Tetragnatha mavillosa Thorell
827' Central ocular quadrangle not a square (Fig. 1727); chelicera with an outer subbasal
cusp in its fang, upper guide tooth absent but slanting tooth present, T very large, t
moderately high, and promargin with about 8-10 teeth (Fig. 1728); ventrally chelicera
with prominent auxillary fang guide (Ax) and G (Fig. 1729); paracymbium almost
parallel-sided in apical half, tip with small and large tubercles (Fig. 1730) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha praedonia L. Koch
828(824) Fang without an outer cusp; promargin with 8 teeth, uppermost 2 teeth widely spaced
enclosing about 4 retromarginal teeth viewed dorsally (Fig. 1731); retromargin with
12-13 teeth (Fig. 1732); genital fold moderately swollen apically with a truncate tip (Fig.
1733); anterior eye row more recurved than the posterior (Fig. 1734) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha maxillosa Thorell
828' Fang with an outer cusp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 829
829(828') Lower auxillary fang guide prominently very large, even larger than the guide (G),
promargin with 13 teeth (Fig. 1735) and retromargin with 15-16 teeth, fang with an inner
cusp in basal one-third (Fig. 1736); genital fold long and penis-like (Fig. 1737) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha mandibulata Walckenaer
829' Lower auxillary fang guide small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 830
830(829') Promargin with a tooth guide (G) rather robust compared to the distant T (Fig. 1738);
retromargin with 11 with Ax present near G and of equal size, tooth below G with a
swollen base (Fig. 1739); genital fold short with a moderately swollen apex (Fig. 1740);
anterior eye row (AER) narrower than posterior eye row (PER), AER strongly
recurved and PER gently recurved (Fig. 1741) . . . . . . Tetragnatha praedonia L. Koch
830' Promargin with about 9 teeth, guide (G) similar. in size and shape with T (Fig. 1742);
retromargin with 12 teeth, fang with a dorsal cusp viewed laterally (Fig. 1743); genital
fold moderately short with an apical orifice (Fig. 1744) . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragnatha nitens (Audouin)
831(816') Femur with trichobothria (Fig. 1745); paracymbium (Fig. 1746) usually leaf-like and
often divided; abdomen ovoid and occasionally with humps [Metidae] . . . . . . . . . . 832
831' Femur without trichobothria; paracymbium often a sclerotized hook (Fig. 1747); ab-
domen variable often with humeral, dorsal or posterior bumps [Araneidae] . . . . . 835
832(831) Abdomen yellowish with black spots and elevated anterodorsally becoming subtrian-
gular in lateral view (Fig. 1748); epigynum truncate along posterior epigynal margin,
rectangular towards the anterior with a pair of sclerotized circular structures (Fig.
1749); genital bulb subglobular, distal part of embolus and conductor rather bifurcate;
paracymbium with a large tooth directed laterally (Fig. 1750); basal half of femur IV
without a thick layer of trichobothria . . . . . . . . . . . . . . . . . . . Tylorida striata (Thorell)
832' Abdomen moderately elongated, usually greenish yellow; basal half of femur IV with a
double fringe layer of trichobothria (Fig. 1751) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 833
833(832') Abdomen without paired tubercles anterodorsally, uniformly silvery white with lon-
gitudinal black bands (Fig. 1752); midanterior part slightly cleft; epigynum with a
transverse C-shaped cavity facing the posterior margin, median guide narrow, small
notch present below anterolateral corner (Fig. 1753); palpal organ with cymbium and
tibia similar in length, tibia and patella each with a long subapical hair (Fig. 1754) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucauge celebesiana (Walckenaer)
833' Abdomen with 1 or more pairs of tubercles anterodorsally . . . . . . . . . . . . . . . . . . . . 834
834(833') Caudal tubercle present (Fig. 1755); abdomen with one pair of tubercles, tapers
posteriorly and dorsally lined with silvery white and blackish bands (Fig. 1756);
epigynum with a medially constricted guide, top of transversely C-shaped cavity with
rectangular-like band moderately cleft anteromedially (Fig. 1757) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucauge decorata (Blackwall)
834' Caudal tubercle absent; abdomen with 1 pair of anterodorsal tubercles, each tubercle
black on top, entire abdomen yellowish with 3 longitudinal dark bands (Fig. 1758);
epigynum rounded anteriorly with a pair of obliquely converging orifices divided by a
guide (Fig. 1759); palpal organ with an upwardly curved and apically rounded paracym-
bium (Fig. 1760) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucauge blanda (L. Koch)
834'' Caudal tubercle absent; abdomen with 3 pairs of tubercles in the anterodorsal half (Fig.
1761) strongly overlapping the carapace; silvery white dorsally with a median mark of
greyish and blackish patches and a pair of narrow longitudinal black bands laterally (Fig.
1762); epigynum semirectangular with a pair of sclerotized areas divided by a narrow
cavity in the middle (Fig. 1763) . . . . . . . . . . . . . . . . . . . . Leucauge bengalensis Gravely
835(831') Abdomen with lateral and posterior spines; dorsally flattened, hard and provided with
conspicuous yellow and black transverse bands or yellowish white lateral bands; spin-
nerets in an elevated circular area flanged by a ring . . . . . . . . . . . . . . . . . . . . . . . . 836
835' Abdomen without lateral and posterior spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . 837
836(835) Six abdominal spines (4 laterals and 2 posterior) very short; abdomen roughly octagonal,
dorsum with a pair of large yellowish white patches laterally and dark brown medially
(Fig. 1764); carapace with a median conical bulge in the cephalic region; epigynum
transversely oblong with a pair of sclerotized black parts in the middle and moderately
exserted guide (Fig. 1765) . . . . . . . . . . . . . . . . . . . Gasteracantha mammosa C. L. Koch
836' Lateromedian spine of the abdomen conspicuously large; abdominal dorsum with 3
transverse yellowish white bands, median one connects the large spine; sigilla ovoid with
8 in the anterior dark band and 6 in the posterior dark band (Fig. 1766); epigynum
transversely ovate with undulated posterior epigynal margin (Fig. 1767) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gasteracantha sp.
837(835') Posterior eye row strongly procurved; anterior lateral eyes smaller than posterior
laterals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 838
837' Posterior eye row nearly straight or procurved; lateral eyes subequal . . . . . . . . . .841
838(837) Abdomen subglobular with a pair of humps anterodorsally; anterior eyes evenly spaced
and laterals closer to posterior laterals; posterior eye row strongly procurved (Fig.
1768); carapace spread sidewards frontally (Fig. 1769); epigynum subglobular with the
slight V-shaped guide a little below posterior epigynal margin (Fig. 1770) . . .Gea sp.
838' Abdomen moderately elongated oval, broadly ovoid, and usually with a blunt anterior
part; transverse furrow in the foveal region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 839
839(838') Abdomen with alternating transverse yellow and black stripes . . . . . . . . . . . . . . . .840
839' Abdomen with silvery whitish yellow band anteriorly and brown patches of irregular
shapes from the median towards the posterior (Fig. 1771); epigynum with a broad scape
(Fig. 1772) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Argiope catenulata (Doleschall)
840(839) Abdomen broadly oval, dorsum grey-white with network-like black stripes in the
posterior half (Fig. 1773); apical half with 3-4 black almost uninterrupted transverse
stripes; epigynum triangular anteriorly, scape moderately cleft apically (Fig. 1774) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Argiope aemula (Walckenaer)
840' Abdomen elongate-ovate, apico-dorsal black band slightly concave at middle, median
area with transverse black bands reaching margins and a band each above and below it
not extending to the margins (Fig. 1775); apical part of epigynum broadly rounded and
button-like, posterior part truncated (Fig. 1776) . . . . . .Argiope bruennichii (Scopoli)
841(837') Abdomen raised anteriorly and with a pair of shoulder tubercles (Fig. 1777); carapace
flat with a prominent thoracic groove and long cephalic region; ocular quad a little
longer than wide; lateral eyes equal and slightly separated from each other (Fig. 1778)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyrtophora sp.
841' Abdomen not raised anteriorly, with or without shoulder tubercles; carapace not flat,
thoracic groove may or may not be distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 842
842(841') Carapace provided with a U-shaped junction between cephalic and thoracic region;
posterior median eyes very close and almost contiguous; abdomen without median and
paired tubercles antero-dorsally but with a pair of blunt lateral humps posteriorly and a
median caudal one, generally rhombic with broad median silver band and black margins
(Fig. 1779); epigynum with a long narrow scape, tip bent (viewed laterally); bilobed
laterally above it and moderately broad anteriorly (Fig. 1780) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclosa insulana (Costa)
842' Carapace without U-shaped junction; posterior medians not very close to each other
....................................................................... 843
843(842') Abdomen elongated, more than twice longer than wide, a little pointed antero-medially
over carapace but without caudal projection (Fig. 1781); epigynum transverse elongate
with median constriction anteriorly and a globular mark above it (Fig. 1782) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Larinia sp.
843' Abdomen subovate to ovate, less than twice longer than wide . . . . . . . . . . . . . . . . 844
844(843 ') Carapace with a longitudinal furrow (best seen in front-lateral view); ocular quad forms
a trapezium and slightly longer than wide; tibia II of male with macrosetae prolaterally;
epigyne simple tongue-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 845
844' Carapace without a longitudinal furrow, usually a transverse furrow present . . . . 848
845(844) Yellow-bodied spider including legs with about 5 transverse dark bands not extended to
the margins of the abdomen, 3 longitudinal dark stripes in the carapace, and light brown
bands on apices of legs (Fig. 1783); epigynum short tongue-like, narrows before sub-
truncate apex (Fig. 1784); venter of abdomen with a pair of parallel longitudinal white
bands and a transverse one with enlarged tips above spinnerets (Fig. 1785); palpal organ
with a deeply bifurcated median process (Fig. 1786) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoscona doenitzi (Boesenberg & Strand)
845' Brown with whitish bands to dark reddish brown; legs yellowish brown to reddish brown
with or without dark brown bands on apices; abdominal folium a pair of zigzag-like
patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 846
846(845') Carapace with 3 longitudinal dark brown bands becoming fused posteriorly in male
(Fig. 1787), open in the female and with series of constrictions in the median band (band
sometimes dagger-like) of the abdomen (Fig. 1788); epigynum long and tongue-like,
apex rounded and constricted at midhalf before the swollen base (Fig. 1789); venter of
abdomen with a pair of C-shaped white bands facing each other and a pair of ovoid
white spots above sides of spinnerets (Fig. 1790) . . . . .Neoscona theisi (Walckenaer)
846' Carapace without traces of longitudinal dark brown bands . . . . . . . . . . . . . . . . . . . 847
847(846') Apices of legs blackish brown similar to the cephalic region, thoracic area reddish;
abdominal folium variable ranging from white with a red-brown spot, white mottled
brown with a pair of zigzag blackish spots, and brown with white cardiac area and 4 pairs
of blackish brown oblique spots (Figs. 1791-93); epigynum long, upcurved before apex
and bent apically provided with a canal in the middle (Fig. 1794); median apophysis
indented prior to the slightly outcurved structure; tibia rather longer than wide (Fig.
1795) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoscona scylla (Karsch)
847' Tibia of legs with pale transverse subapical and submedian dark bands; carapace
blackish to yellow-brown (sometimes light medially and darker marginally); abdomen
nearly triangular with a tree-like pattern provided with 3 pairs of moderately oblique
black spots posteriorly (Fig. 1796); epigynum projected out of body in lateral view (Fig.
1797) and triangularly tongue-like in ventral view (Fig. 1798) . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoscona nautica (L. Koch)
848(844') Abdominal surface shiny with very few hairs, usually ovoid marked with broad lon-
gitudinal dark bands separated by white areas; legs relatively short . . . . . . . . . . . . 849
848' Abdominal surface dull but clothed with many hairs, usually subtriangular or partly
ovoid with different markings; legs relatively longer . . . . . . . . . . . . . . . . . . . . . . . . . 851
849(848) Anterior medians the largest; median ocular quad wider in front than behind; cephalic
region usually more dark colored than the thoracic area; abdomen chalk white with a
pair of broad longitudinal submedian bands and 4 pairs of light brown globular spots
below cardiac area (Fig. 1799); epigynum with noticeable scape and tubercle-like
protrusions on each side of scape in the posterior epigynal margins (Fig. 1800); tibia of
palpal organ more or less transverse (Fig. 1801) . . . . . . . . . . . . Singa hamata (Clerck)
849' Posterior median eyes the largest; abdomen of female usually with 2 broad longitudinal
darkbands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 850
850(849') Carapace dark brown to black with light lateral margins; femur without median dark
bands; abdomen very dark brown to black with cream or light yellow dorsal stripes, dark
bands on abdomen with 2 moderate swellings laterally and 1 pair of spots inside the
cream band anteriorly (Fig. 1803) or abdomen entirely cream-colored with about 4 light
brown spots (Fig. 1802); epigynum transverse ovate with a broad median piece directed
towards posterior epigynal margin (Fig. 1804); median apophysis curved and tip of
conductor broadly blunt (Fig. 1805) . . . . . . . . . . . . . Hypsosinga pygmaea (Sundevall)
850' Carapace uniform reddish brown without light lateral margins; femur I not darkened;
abdomen with broader dark brown bands without lateral swellings, lateral white bands
sometimes not contiguous posteriorly (Fig. 1806); abdomen of male orange with a pair
of ovoid black spots (Fig. 1807); epigynum transversely E-shaped with median guide
directed towards the anterior margin (Fig. 1808); median apophysis acutely pointed and
narrow, conductor narrowly blunt at tip (Fig. 1809) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypsosinga sanguinea (C. L. Koch)
851(848') Abdomen white with a double V-shaped marking dorsally; carapace yellow-brown (Fig.
1810); epigynum with a moderately long scape extended beyond epigynal margin
posteriorly (Fig. 1811) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Araneus sp.
851' Abdomen without a double V-shaped mark, usually with a pair of moderately oblique
wavy stripes widening anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 852
852(851') Legs uniformly yellow; abdomen blackish with chalk white median band constricted
along lateral margins producing subtriangular marks (Fig. 1812); male with a broad
median band toward eyes (Fig. 1813); epigynum simple with a strongly rounded anterior
and short but wide tongue-like median plate (Fig. 1814) . . . Araneus inustus (L. Koch)
852' Apices of legs dark banded; abdomen with a pair of oblique wavy lines dorsally, tapers
posteriorly (Fig. 1815); scape long, with constricted edges and bent in basal one-third
viewed laterally (Fig. 1816); median apophysis dolphin-head like at tip (Fig. 1817) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Araneus diadematus Clerck
ACKNOWLEDGEMENTS
We thank the following individuals/institutions for the use of their figures: Dr. A. Lewanich,
Entomology and Zoology Division, Department of Agriculture, Bangkok, Thailand (Scirpophaga);
Dr. J. Roffey, Overseas Development Natural Resources Institute, U. K. (locusts and grasshop-
pers); Dr. D. Hollis, British Museum Natural History, U. K. (Oxya), Dr. R. W. Crosskey, BMNH
(Tachinidae) and Trustees of the British Museum (Natural History); Dr. Teiji Kifune, Dept. of
Parasitology, Fukuoka University, Japan (H. thaiae); Dr. I. Miyagi, Medical Laboratory, Ryukyu
University, Okinawa, Japan (Ephydridae); Dr. K. Kamijo, Hokkaido Forest Expt. Station, Hok-
kaido, Japan (Trichomalopsis); Dr. K. Yano, Laboratory of Applied Entomology, Faculty of
Agriculture, Yamaguchi University, Japan (Pipunculidae); Dr. K. A. Sahad, Plant Protection
Division, Dept. of Agricultural Extension, Bangladesh (Mymaridae); Dr. N. F. Johnson, Dept. of
Entomology, College of Biological Sciences, Ohio State University, Columbus, Ohio (Scelionidae-
Telenomus ); Dr. M. Olmi, Institute of Plant Protection, University of Tuscia, Viterbo, Italy
(Dryinidae); and Dr. H. Nagaraja, CIBC, Indian Station, Bangalore, India (Trichogramma and
Trichogrammatoidea) .
We are sincerely grateful to Miss Nonnie Bunyi for typing the manuscript; Mr. Danilo Amalin
for the illustrations; Mrs. M. Austria for providing taxonomic literature of rice pests and their
natural enemies; Misses J. L. Catindig, T. E. Ponce and Mrs. A. Jaballa for library work.
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II. BIOLOGY AND ECOLOGY
3
Insect Pests of the Rice Plant -

Their Biology and Ecology
D. DALE
Kerala Agricultural University
Kerala, India
1. INTRODUCTION
THE RICE PLANT IS an ideal host for many insect species. All of the plant parts are vulnerable to
insect-feeding from the time of sowing till harvest. There are over 800 insect species damaging rice
in one way or another, although the majority of them do very little damage. In tropical Asia only
about 20 species are of major importance and of regular occurrence (Grist and Lever 1969).
Several species that were earlier considered as minor pests have recently become major pests,
whereas the incidence of a few others has considerably declined (Pathak and Dhaliwal 1981). In
addition to direct damage (Fig. I), many insects also act as vectors of dreaded diseases of rice.
Rice is primarily a crop of warm and humid environments. About 60% of the world’s total rice
area is in tropical Asia where insect pest problems are also most severe and numerous. Hence more
pests from this region are extensively covered in this chapter. But still, many major, important and
representative pests from all the other rice growing regions of the world have been included.
Under each pest synonyms, distribution, host plants, symptoms and extent of damage, life cycle
and ecology are given. Mention of a country’s name in the distribution only indicates that the pest
has been reported from some part of the country and not necessarily that the species is distributed
throughout the country. In the case of a few minor pests, little information on their ecology is
available for inclusion in this chapter.
364 RICE INSECTS — BIOLOGY AND ECOLOGY
Fig. 1. Possible relationship between N. lugens feeding and the development of hopperburn
symptoms in rice plants (after Sogawa and Cheng 1979).
2. ROOT AND STEM FEEDERS

There are many insects which feed on the roots and stem of rice plants. Some, such as white grubs and
root aphids, attack exclusively the roots; others (e.g. mealybug, gall midge, rice stem maggot) infest only
the stems while there are some pests such as mole crickets which damage both roots and stems.
The infestation of the rice crop by different insect pests is related also to the growth stage of the
plants. Seedlings are prone to attack by pests such as the rice seedling flies and the rice seed
midges. Insect damage at the early stages of the crop causes the death of the seedlings and leads to
an uneven stand.
2.1 Mole Crickets, Gryllotalpa africana Palisot de Beauvois (Fig. 2.1), Gryllotalpa orientalis
Burmeister (Orthoptera: Gryllotalpidae)
Distribution: Africa, tropical Asia, Europe, Japan
Mole crickets are polyphagous insects. They occur in all rice environments but are most prevalent
in non-flooded upland rice when the fields are damp. In Japan, much of the damage is noted during
spring and autumn when the rice plants are young (Kureha et al. 1974). Severe infestations of these
pests in nursery beds have been reported from India and Japan.
The nymphs and adults attack stems below ground and close to the roots. Sometimes only one
or two tillers are cut and damage is evident only when the tillers begin to dry a few days later. The
entire plant dies if the attack is severe. Dried plants can be seen as patches in the rice field. Young
and newly-planted seedlings are most commonly attacked in the early part of the season before
fields are flooded. Normally damage is greatest near field borders. Unlike field crickets, mole
crickets do not take the cut tillers into their burrows (Tripathi and Shri Ram 1968).
DALE 365
Fig. 2.1
Adult mole crickets are large, light brown in color and 25 to 35 mm long. The forelegs are
highly modified for burrowing soil. Adults cannot survive in the rice fields after flooding. They are
strong fliers, and are attracted to light traps at night. Female crickets burrow bunds and construct
hardened cells in which the eggs are laid. Each cell contains 30 to 50 eggs that hatch in 15-40 days
depending on the temperature. Each female may deposit several hundred eggs during its life span
of more than 6 months.
Feeding by nymphs causes bare patches in the field. They have only limited capacity for
migration and suffer heavy mortality. Cannibalism exists among mole cricket populations. The
nymphal period lasts 3 to 4 months.
There is only one generation a year. Adults overwinter in burrows deep in the soil.
2.2 Rice Root Aphids (Homoptera: Aphididae)

The aphid species that infest rice roots and their distribution are given in Table 1. Some of these
aphids are also occasionally found on the aerial parts of plants.
Fig. 2.2
TABLE 1
World distribution of rice root aphids (modified from Yano et al., 1983)
Aphid species Host plants Distribution

Anoecia corni (Fabricius) Rice Japan
Anoecia fulviabdominalis (Sasaki) Rice, barley, wheat, grasses Japan
Anoecia sp. Rice Japan
Chaetogeoica polychaeta Rice, grasses India
Pal & Raychaudhuri
Forda sp. Rice Japan
Geoica lucifuga (Zehntner) Rice, sugarcane, grasses, Pistacia India, Japan, Malaysia,
Philippines, Taiwan
Geoica setulosa (Passerini) Rice Italy
Geoica sp. Rice Japan
Geoica utricularia setariae Rice Italy
(Passerini)
Paracletus cimiciformis Rice, barley, grasses Japan
von Heyden
Prociphilus sp. Rice, plants belonging to Araceae Japan
and Liliaceae
Rhopalosiphum rufiabdominalis Prunus sp., rice, grasses Argentina, Central America,
(Sasaki) Egypt, Fiji, India, Japan,
Malaysia, Morocco, Surinam,
Taiwan, Thailand, USA, USSR
Tetraneura akinire (Sasaki) Rice Japan
Tetraneura basui Hille Ris Lam- Rice, grasses India
bers
Tetraneura nigriabdominalis Rice, graminaceous weeds Cuba, Fiji, India, Indonesia,
(Sasaki) Japan, Malaysia, New Guinea,
Philippines, Sierra Leone,
Taiwan, Zambia
Tetruneura radicicola Strand Rice, grass Ulmus sp. India
Tetraneura sp. Rice Japan
DALE 367
Rice seems to suffer very little from root aphids. The damage is mostly due to a few species in
isolated areas and that too only at particular seasons of the year. Root aphids occur only in
well-drained soils in rainfed environments. Adults and nymphs remove plant sap and cause the
leaves to turn yellow and become stunted. But in severe cases, which are very rare, plants wilt and
die. In Japan, attacks during the early stage of the crop cause reduction in yield as high as 50%
(Tanaka 1961)
Rhopalosiphum rufiabdominalis (Fig. 2.2) is the most important species among the root aphids.
It is recorded as associated with ants on the roots of various host plants. The species is mostly a pest
of upland rice in Japan and Taiwan and is seldom seen as a threat to irrigated rice anywhere in the
world. In Japan, the aphids move to the drained irrigated fields in late summer or autumn and settle
on the rice stubbles. Both adults and nymphs congregate on the upper parts of rice roots just above
the soil surface when the water is drained from the fields. They sometimes feed on the stem and leaf
bases when the aphid population becomes very high and/or the upper portions of the roots are
submerged.
Root aphids consist only of the female sex. Adult aphids exhibit wing dimorphism. The
apterous female is dark or light green with large red blotches. The nymph-laying period and adult
longevity are usually longer for apterous forms than their alate counterparts. The alate form is
smaller and mainly reddish brown in color. The winged adults are adapted for migration and they
increase in number under stress conditions such as shortage of food and crowding.
The adult aphids reproduce parthenogenetically throughout the year. They start laying nymphs in
a day or two after emergence. Fecundity is higher in summer and fall than in winter and early spring.
The nymphs molt 4 times before they reach adulthood. The nymphal period is about 7 to 9 days. Winged
adults are poor fliers but they often drift long distances by wind or are carried by animals. Although the
aphid could complete 53 to 58 generations in a year under laboratory conditions in Taiwan (Hsieh 1970),
only 20 generations have been recorded in Japan (Yano et al. 1983).
The spring migrants (alate forms) fly from their winter host plants to the rice plants of the first
crop. Once this crop is harvested, the aphids move to either summer grasses or rice plants of the
second crop. During harvest, the alate forms settle on wheat or rice stubbles where they deposit the
overwintering progeny.
Tetraneura nigriabdominalis is a widely distributed species of root aphid. In Japan, it infests
only upland rice but not irrigated fields. Many grasses such as Eleusine indica (L.) Gaertn.,
Pennisetum subangustum Stapf and Hubb., Ischaemum rugosum Salisb. and Paspalum
scrobiculatum L. serve as alternate hosts in Sierra Leone (Akibo-Betts and Raymundo 1978).
The adult aphids are more or less spherical and brown in color. The body is enveloped in a thin
film of white powder. Mature adults measure 3 to 5 mm long. In Japan the infestation starts by
April or May every year.
Anoecia fulviabdominalis is a major aphid species attacking the roots of upland rice in Japan.
In May, the alate spring migrants move from their primary hosts, Cornus spp., to secondary host
plants such as barley, wheat and wild grasses. Later, the aphids are carried by wind or ants to
upland rice fields.
2.3 Rice Mealybug, Brevennia rehi (Lindinger) (Homoptera: Pseudococcidae)
Synonyms: Ripersia oryzae Green

Ripersia sacchari Green
Ripersia rehi Lindinger
Heterococcus rehi Lindinger
Tychea rehi (Lindinger)
Distribution: Australia, Bangladesh, Burma, India, Nepal, Papua New Guinea, Puerto Rico,
Southern USA
Host plants other than rice: Apluda mutica L, Brachiaria reptans (L) Gardner & Hubbard, Chloris
barbata (Sw.), Cymbopogon caesius (Nees) Stapf, Cynodon dactylon (L.) Pers.,
Cyperus rotundus L., Dactyloctenium aegyptium (L.) Willd., Dichanthium an-
nulatum (Forssk.) Stapf, Digitaria sanguinalis (L.) Scop., Echinochloa colona (L.)
Link, Eleusine coracana (L.) Gaertn., Eragrostis interrupta (Lam.) Doell., Fimbris-
tylis miliacea (L.) Vahl, Fimbristylis tenera Roem. & Schult., Imperata cylindrica (L.)
Raeuschel, Isachaemum indicum (Houtt) Merr., Iseilema laxum Hack., Leptochloa
chinensis (L.) Nees, Panicum repens L., Paspalum scrobiculatum L., Saccharum
spontaneum L., Setaria glauca (L.) Beauv.,
The rice mealybug is a sporadic pest causing severe losses to rice in the Indian subcontinent. It
is prevalent in rainfed rice and is not found in irrigated rice.
Mealybugs occur in colonies attached to the stems and leaf sheaths of rice plants. They suck
sap from the plants. White waxy fluff protruding from between the leaf sheath and the stem is the
typical indication of the pest’s presence. As a result of infestation, plants become stunted and the
Fig. 2.3
DALE 369
older leaves turn yellow. High incidence inhibits panicle emergence and plants may even dry.
Infested fields show isolated patches of stunted plants. Damage is severe during drought conditions
when plants can least tolerate the removal of plant sap.
In 1979, when there was a long and severe drought in Bangladesh, a severe outbreak of mealybugs
caused yield losses of up to 100% in some fields (Alam and Karim 1981). Heavy losses caused by
mealybugs have been reported from several parts of India (Banerjee 1956, Mammen 1976).
The adult females are wingless, oblong and 3-4 mm long. They are bulky and remain stationary
on the stems behind leaf sheaths at the base of plants. The body is soft, pinkish, and covered with
white waxy threads. Males, pale yellowish, are seldom found in the colonies. They have a single pair
of wings and a style-like process at the end of the abdomen but lack mouthparts. They are slender
and much smaller than the females. Males migrate freely from plant to plant. Reproduction is
parthenogenetically oviparous as well as viviparous (Alam 1965).
The average preoviposition period is 6 days. The female mealybug lays eggs singly, more of less
in a chain, inside the waxy threads. A single female can lay 60 to 280 eggs and nymphs in about 5
days, after which it dies. Eggs are yellowish white. The incubation period varies from 3 to 6 hours.
Hatching of eggs usually takes place in the morning hours.
Newly-hatched nymphs remain crowded within the waxy threads for 6 to 10 hours before they
disperse to various parts of the same plant or to adjacent rice plants. Nymphs are first yellowish
white but turn dark yellow after a day. They become fixed between the leaf sheath and stem by
driving the proboscis into the stem. The body gets covered with a waxy material in about a day.
There are three nymphal instars which take 12 to 18 days to pass and about 12 generations in a year.
Hot, dry weather favors the survival and multiplication of rice mealybugs. But the population
drops drastically by the beginning of winter. During winter, the pests survive on weeds and
volunteer rice plants. In Nepal, overwintering of mealybugs in both nymphal and adult stages has
been observed (Pradhan 1981). When the weather becomes warm, the insects again appear in the
rice fields.
Although the reasons for the positive relationship between the high incidence of mealybugs and
drought conditions are not precisely known, Fennah and Smith (1954) reported that a low iron
content in the soil during drought upsets the normal metabolism of the plant and makes soluble
nitrogen more available to 'the mealybug Neorhizoecus epicopus. Through simulated drought
condition, Wiggins and Williams (1955) showed that amino acid content in the plant increased
significantly. Availability of extra amino acid favors rapid growth and multiplication of mealybugs.
Fennah (1960) postulated that metabolic upsets in plants caused by water stress accelerates
breakdown of protein and that the altered physiological condition of the plant is more nutritionally
favorable for sap-sucking insects.
2.4 Black Bugs, Scotinophara coarclata (Fabricius), S. lurida (Burmeister), and

S. laliuscula Breddin (Hemiptera: Pentatornidae)
Distribution: S. coarctata: Bangladesh, Burma, India, Indonesia, Kampuchea, Malaysia, Pakistan,
Philippines (Palawan), Sri Lanka, Thailand, Vietnam (Fig. 2.4A)
Fig. 2.4A Fig. 2.4B Fig. 2.4C
S. lurida: Bangladesh, China, India, Indonesia, Japan, Kampuchea, Malaysia,

Pakistan, Papua-New Guinea, Philippines (Mindanao), Sri Lanka, Thailand, Viet-
nam (Fig. 2.4B).
S. latiuscula: Indonesia (Sumatra), Philippines (Luzon) (Fig. 2.4C)

Host plants other than rice: Colocasia esculenta Schott., Hibiscus esculentus L., Hymenachne
pseudointerrupta C. Muell., Panicum amplexicaule Rudge Pl. Guian., Scirpus grossus
L., Scleria sumatrensis Retz., Vigna unguiculanta L., and Zea mays L.
In recent years, many outbreaks of black bugs have been reported even though the insects were
previously considered as minor pests of rice. Scotinophara coarctata appeared in South Palawan,
Philippines in 1982 and a major infestation followed during the subsequent months. In Indonesia
and Malaysia, there were outbreaks in 1978 and 1979 respectively.
Nymphs and adults feed chiefly at the base of stems where they remove plant sap. When the
infestation is at the tillering stage, deadhearts occur but continued feeding results in leaves turning
chlorotic or reddish brown color, and in reduction in tiller number and stunting. Attack during the
booting stage, results in panicles with empty grains similar to the ‘deadhearts’ caused by stem
borers. Direct injury to panicles is also common. Bugs feed on panicles in the milky stage on
overcast days and during night. Injured grains are spotted brown. Heavy infestations may lead to
the death of plants and the whole field appears ‘burned’ similar to that of a hopperburned field.
The effect of black bug feeding on yield components of some rice cultivars was studied by
Heinrichs et al. (1987). They found that yield losses were due to unfilled grains, decrease in tiller
DALE 371
number and less grains per panicle. Percent yield loss at 10 bugs per hill ranged from 14.7% in
resistant cultivars to 23.0% in susceptible cultivars.
Adults are brownish black with a few distinct yellowish spots on the thorax which bears spines
below the anterior angles. They are 8-9mm long. Tibiae and tarsi are pinkish. They give off an
offensive odor typical of stink bugs when disturbed. Adult bugs live for up to 7 months. There is
one generation in a year.
Female bugs deposit eggs on the basal parts of rice plants near the water surface. A female lays
about 200 eggs during her lifetime and guards them until they hatch. Each egg measures 1 mm long,
is greenish when laid and turns pinkish as it matures. The incubation period is 4 to 7 days.
Nymphs are light brown, with a yellowish green abdomen and some black spots. They molt 4
to 5 times and reach the adult stage in 25 to 30 days.
Large numbers of bugs are carried to distant places by strong winds. Adults appear in swarms,
and are strongly attracted to light. Ajuk et al. (1981) and Latif et al. (1982) observed that the flight
activity of adults to light traps coincides with the lunar cycle.
The bugs are able to adapt to a wide variety of conditions and are capable of withstanding
adverse conditions. They hide in cracks in soil during periods of water stress and during the winter.
After overwintering they fly to the rice crop and reproduce over several generations. They again
return to their resting sites after the harvest of the rice crop. The black bug aestivates in the adult
or late nymphal stage in cracks in bunds, in paddy fields or in adjacent higher grounds to a depth of
30 cm., where it remains torpid. It is gregarious during periods of rice fallow.
Populations are generally less on upland than on irrigated and rainfed wetland rice. The insects
prefer continuously cropped irrigated fields to a single-cropped field. Heavy damage by the pest is
usually observed after the heading stage of the rice crop especially when irrigation has been stopped
during thc maturation period.
In the Philippines, the dry season rice crop is damaged more severely than the wet season crop
(Miyamoto et al. 1983). The pest is most abundant in poorly-drained rice fields around marshes.
Asynchronous double cropping of irrigated rice with high levels of nitrogen seems to favor black bug
outbreaks. In Malaysia, the relatively small natural enemy complex of S. coarctata and high
availability of alternate hosts in the vicinity of rice fields are suggested as possible causes for the
frequent black bug outbreaks.
Three species of Scotinophara have been reported from the Philippines, but these species are
distinctly isolated in their distribution; S. coarctata in Palawae S. lurida in Mindanao and S.
latiuscula in Luzon.
2.5 Rice Seed Midges Cricotopus sylvestris (F.), Paralauterborniella subcincta, and
Paratanytarsus sp. (Diptera: Chironomidae)
Distribution: Europe, USA
Host plants other than rice: some weeds
Fig. 2.5
Rice seed midges are common in California rice fields. Serious damage to rice is limited to
germinating seeds and very young seedlings. Maggots feed on primary roots and shoots or may
hollow out embryos, killing plants. They may also feed on floating leaves, causing small holes that
extend completely through the leaves. Once seedlings are fully established, the plants can withstand
damage by the larvae. Larval populations increase daily after flooding because of continued egg
laying, and delay in planting will expose germinating rice seeds to more midge infestations.
Adult flies resemble small mosquitoes as they swarm over rice fields and levees. Masses of eggs
are laid on water in strings held together by a sticky, mucilagenous material that swells in contact
with water to form a protective covering over the eggs. Eggs hatch in one or two days.
The larvae use secreted silk and frass to build tubes on the bottom of the rice paddy and on
submerged vegetation. The tubes serve as larval retreats and also as webs to entrap algae and
diatoms for food. Larvae may also feed on rice seedlings and other plants. Damage to rice is caused
mostly by the third and fourth instar larvae. The larvae pass through 4 instars in 7 to 10 days in the
spring when the water is warm. Pupation takes place in the silk tubes under water. The pupal
period is 2 to 3 days. Three to four generations occur each summer, but only the first two are of
economic concern to rice growers in the USA(Anon. 1983)
Numerous other species of Chironomidae have been reported occasionally infesting rice fields
in various countries. In Egypt, a Chironomus sp. causes damage to rice seedlings grown in saline
soils. The Australian Chironomus tepperi prefers soils with a high organic content following sod
seeding. In the USSR, larvae of a Chironomus sp. feed on floating or submerged rice leaves causing
a severe reduction in shoot numbers.
2.6 Asian Rice Gall Midge, Orseolia oryzae (Wood-Mason) (Diptera: Cecidomyiidae)
Synonym: Pachydiplosis oryzae (Wood-Mason)
Distribution: Bangladesh, Bhutan, Burma, China, India, Indonesia, Kampuchea, Laos, New
Guinea, Pakistan, Sri Lanka, Thailand, Vietnam.
DALE 373
Fig. 26
Host plants other than rice: Cynodort dacfylott (L.) Pers., Echinochloa colorza (L.) Link,
Heteropogon contomts (L.) Beauv., Ischaemunt aristatunt L., Leersia hexrrrtdru Sw.,
Oyza barthii A. Chev., Panicum miliaceum L., Paspalurn scrobiculatum L., Sac-
ciolepis interrufa (Willd.) Stapf
Orseolia oryrae is primarily a pest of lowland irrigated rice but has been rcported in upland and
deepwater rice (Venugopala Rao 1975) also. The extent and severity of gall midge infestations has,
significantly increased during the last two decades. Favorable weather conditions, cultivation of
high-tillering varieties, intensive management practices and low parasitization are conducivc to thc
rapid multiplication’of this pest.
The main external symptom of gall midge attack is a ‘silver shoot’ or ‘gall’ that resembles an
onion leaf. A fully developed gall is a silvery white, hollow tube about 1 cm wide and 10-30 cm long.
Larval feeding suppresses leaf primordial differentiation at the growing tip. This, in turn, induces
the development of radial ridges from the innermost leaf primordium followed by an elongation of
the leaf sheath (Perera and Fernando 1970). Galls appear within a week after the larvae reach the
growing point. Attack of rice seedlings leads to profuse tillering and stunting of the plants. A late
infestation of the plants at the reproductive shoot apex malforms the leaves and panicles. Israel et
al. (1959) reported 0.5% loss in yield for every unit percent increase in incidence. But wide varietal
differences exist in the effect of infestation on crop losses. In certain varicties, gall formation is not
manifested and instead necrosis of the shoot apex results (Picris 1977).
The gall midge attacks rice from the nursery to the end of tillering stage. Young maggots
cannot survive in plants past the vegetative stage as there are no actively growing apical buds for
them to infest.
The periodicity of infestation and extent of damage caused by the gall midge varies in different
countries. In Thailand, the pest has long been reported associated with the rainfed rice of the
North, Northeast and Eastern regions, but in recent years, outbreaks in the irrigated dry season rice
of the Central Plains have also occurred (Katanyukul et al. 1980a). In India, the gall midge was
previously a pest only in the wet season crop but recently it has been observed in the winter crop as
well (Kalode and Kasiviswanathan 1976). In Sri Lanka, high infestations of gall midge were
observed in the southwest monsoon season (April to July) in the wet zone while in the intermediate
and dry zones, they were mostly seen during the northeast monsoon (October-November)
(Kudagamage and Nugaliyadde 1981). The extent of insect damage reported by various workers is
presented in Table 2.
TABLE 2
Damage by O. oryzae in South and Southeast Asia
Country Zone Season Damage Reference

India Tamil Nadu Late 30% hills Natarajan and Chandy 1978
India Uttar Pradesh Late 65% hills Rizvi and Singh 1980
India Madhya Pradesh Second crop 45% hills Kaushik et al. 1979
India Karnataka 50% yield Subramanian 1935
India Nizamabad 30% yield Khan and Murthy 1955
India 15% yield Israel 1959
India 25% yield Lever 1970
Thailand 50% yield Grist 1965
Thailand Northern Wet 33% yield Katanyukul et al. 1980b
Thailand Northeastern Wet 27% tillers Katanyukul et al. 1980b
Sri Lanka Dry & Intermediate Maha 44% tillers Wickremasinghe 1969
Sri Lanka Dry & Intermediate Yala 23% tillers Wickremasinghe 1969
Sri Lanka Dry & Intermediate Maha 100% hills Wickremasinghe 1969
Sri Lanka Dry & Intermediate Yala 18% hills Wickremasinghe 1969
Sri Lanka Wet Yala 80% yield Wickremasinghe 1969
China October 60% hills IRRI 1978
The adult gall midge is similar in appearance to a mosquito. Females have bright red ab-
domens, Mating takes place soon after emergence and egg-laying starts a few hours later. The
female flies mate only once and the unmated ones lay sterile eggs. Adults are nocturnal in habit and
are highly attracted to light traps. Eggs are laid either singly or in groups on the underside near the
base of rice leaves or sometimes on leaf sheaths. A single female lays 100 to 200 eggs either singly
or in groups. The males mostly die in 12 to 18 hours after emergence; females live up to 3 days.
Eggs are long, tubular, and shiny white; sometimes with pink, red or yellow shades. They turn
amber before hatching. Eggs require high humidity for development and hatching. The incubation
period is around 3 to 4 days.
The newly-hatched maggots are grey-white, fairly stout with a pointed anterior end. They
wiggle down the leaf blade in a film of dew and move between the leaf sheath and the stem until they
reach the opening point of the apical or side bud at a node. The larva feeds inside the developing
bud which is a zone of differentiation for new tillers. A hollow chamber called a 'gall' forms around
the larva. As the larva feeds the gall enlarges at the base and elongates having the appearance of an
DALE 375
onion leaf. There are 3 larval instars; total larval period is 15 to 20 days. Generally only one maggot
is found in a tiller.
Pupation takes place inside the gall. The male and female pupae can be easily separated by
their size and color of the abdomen (Panda and Mohanty 1970). Male pupae are small and brown
while the females are larger and pinkish. The color turns darker before adult emergence. The pupa
has several rows of abdominal spines that enable it to move up to the tip of the gall before adult
emergence. The pupa makes a hole at the gall tip with its spines and the midge emerges through
this hole leaving the pupal skin behind. Adult emergence generally takes place at night or early
morning. The pupal period varies from 2 to 8 days. The entire life cycle is completed in about 25
to 38 days.
The gall midge remains inactive as a prepupa in wild rice or weeds during the dry season. At
the onset of the monsoons, it becomes active and completes one or two generations in grasses before
it moves to the rice crop (Reddy 1967). In India pest incidence is high during July and August if
there is rainfall during the daytime. This period coincides with the maximum tillering phase in rice.
The population then declines rapidly in December, primarily, because of the limited availability of
suitable host plants. In some cases, the pest infests rice stubbles left after the first crop.
Wongsiri et al. (1971) reported the ecological conditions that usually prevailed in Thailand at
the times when gall midge populations were high. The sky was cloudy most of the days and the
relative humidity was 75% on an average, the optimum mean temperature for adult emergence
and oviposition ranged from 23 to 27°C. Hidaka et al. (1974) observed that seasonal fluctuations of
gall midge populations were closely related to rainfall. Overcast skies and drizzling rains are
favorable for rapid build-up of pest populations.
Israel et al. (1961) reported an increase in the pest population with high nitrogen levels. The
incidence decreased with high levels of phosphate in combination with nitrogenous fertilizers. The
infestation generally increased with higher levels of potash (Israel et al. 1963). Prakasa Rao (1972)
observed an increased number of absolute tillers and infested tillers per unit area with the applica-
tion of increased levels of nitrogen. Close spacing of transplanted rice resulted in a greater number
of tillers and more leaves per unit area and an increase in relative humidity within the plant canopy
which was suggested as the reason for higher gall midge incidence than under wider spacing
(Prakasa Rao 1975).
Variable reactions of differential rice varieties to O. Oryzae have provided evidence of different
biotypes in South and Southeast Asia. The variability of the species is indicated by three biotypes
in different regions of India.
2.7 The African Rice Gall Midge, Orseolia oryzivora Harris and Gagne
(Cecidomyiidae: Diptera)
Distribution: Burkina Faso, Cameroon, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia,
Mali, Niger, Nigeria, Senegal, Sierra Leone, Sudan, Togo
Fig. 2.7
This species had previously been misidentified and considered as O. oryzae. It has been
recently shown by Harris and Gagne (1982) that the specimens from Asia and Africa are mor-
phologically distinct in the larval, pupal and adult stages.
The distribution of this pest appears to be increasing in the rice growing regions of Africa.
Although the gall midge has been reported from several countries, it has not attained the status of
a major pest in most of them. In Upper Volta, O. oryzivora is one of the most serious and
widespread pests of irrigated, rainfed and lowland rice (Bonzi 1980).
The biology of the pest is similar to that of O. oryzae. Infestation is most severe when the first
rains are early and are followed by a relatively long drought, which delays rice planting. The first
generation occurs on grasses and when rice becomes available, it is attacked. The extent of
infestation may be related to the availability of host plants, the amount of rain, especially at the
beginning of the season, and the cropping pattern at a given locality. Little is known on the pest-host
plant relations (Agyen-Sampong 1982).
2.8 Stalk-eyed Borer, Diopsis macrophthalma Dalman (Diptera: Diopsidae)

Synonym: Diopsis thoracica Westwood
Distribution: Benin, Burkina Faso, Cameroon, Ghana, Ivory Coast, Liberia, Mali, Nigeria,
Senegal, Sierra Leone, Swaziland
DALE 377
Fig. 2.8
Diopsis macrophthalma is a major pest of rice in many parts of tropical Africa. It causes severe
damage in the valley of Benue in North Cameroon, Swaziland and Sierra Leone. In Senegal, it is
found only in the south. In Burkina Faso and Mali, it is a sporadic pest in areas with sufficient
humidity in the dry season. The pest infests irrigated rice fields in Ivory Coast and Benin, and is less
frequent in rainfed areas (Breniere 1976). In Ghana, Abu (1972) noted that it could cause about
9% yield loss. Alghali (1981) reported grain losses from 1.9 to 97% in southern Nigeria.
The stalk-eyed borer is not known to attack crop plants other than cultivated rice. However, it
seems probable that the pest has alternate hosts among wild rice and grasses. Cyperus difformis, a
weed found commonly in rice fields, on which eggs, larvae, pupae or adults are sometimes found,
may be a host plant during non-rice cropping seasons (Alghali 1979).
The pest infestation delays booting and leads to a decline in the number of panicles. If the
attack comes early, it may cause total crop loss; if it occus later, some varieties compensate for
damage by producing new tillers. Studies conducted in Nigeria (Alghali and Osisanya 1982) have
shown strong correlation existing between the number of eggs laid in a plant and number of tillers,
and the number of eggs and deadhearts.
Adult flies are typical diopsids easily recognized by their characteristic eyes borne on the tip of
stalks. Flies are often found in areas with water throughout the year. In the dry season, they occur
in swarms in the shade near streams and canals.
Gravid females lay eggs singly on the upper surface of young leaves, normally in the groove of
the midrib. Boat-shaped, striated eggs with a characteristic anterior projection are fixed to the leaf
with a cement that prevents them from being washed off in heavy rains. Eggs are creamy white when
laid but later darken to tan. Each female fly lays about 20 eggs over period of 10 days. Peak
oviposition occurs 30-40 days after transplanting and almost terminates by the end of the tillering
stage. Alghali and Osisanya (1981) reported that virtually no eggs were laid and no deadhearts
developed on 60-day-old plants.
The larva is a yellowish maggot, about 18 mm long and 3 mm wide, easily recognizable by the
two long abdominal extensions ending in black hooks pointed forward. The maggot moves down
inside the leaf sheath and feeds above the meristem on the central spindle of young leaves causing
a deadheart symptom. Every tiller attacked is lost. In its life, a single larva can destroy up to ten
neighboring tillers (Feijen 1979). Later generations of larvae feed on the developing flower head.
The larval stage lasts for 25 to 33 days.
Pupae are red with brown dorsal bands and clear segmentation. They are flat and almost
triangular in section because of the compression inside the stem. During later stages, the imago can
be seen developing inside the pupal case. Adults emerge after a 10-12 day pupal stadium. There are
two principal generations between June and October, and a third less prominent one in the
off-season.
Diopsis apicalis Dalman is a polyphagous species often encountered in rice fields. It is
distinguished from D. macrophthalma by its small size and an apical smoky spot at the tip of each
wing. The larvae feed on healthy plants or on tissue decomposed after a caterpillar attack. They
may even eat caterpillars or the maggots of D. macrophthalma. However, its useful role as a
predator cannot make up for the damage it causes to the rice crop (Breniere 1976).
2.9 Rice Stem Maggot, Chlorops oryzae Matsumura (Diptera: Chloropidae)
Distribution: Indonesia, Japan, Korea, Vietnam
Host plants other than rice: Alopecurus fulvus J. E. Sm., Hordeum vulgare L., Secale cereale L.,
Triticum aestivum L.
Fig. 2.9
DALE 379
The rice stem maggot is an important pest of rice in Japan and Vietnam where crop losses up to
30% have been reported. In Vietnam, damage is more serious in the colder regions than in the
warm areas. Upland rice and early varieties are more prone to infestation than irrigated and
late-maturing varieties.
The maggots are located near the growing point of the rice plant and feed on leaf blades.
Tillering is reduced and stunting results from early attack. Larvae of later generations feed on
developing flowers and this reduces yield considerably. The type of damage differs in north and
south Japan. In the north, where there are only two generations, the first brood larvae attack young
leaves and growing tips of rice plants. But in the south, where there are three generations, the first
brood maggots infest young leaves while the second brood larvae attack only leaf tips (Uyeda et al.
1962).
The adult which looks like a small housefly is grey with three black longitudinal stripes on the
thorax. Female flies lay 50-100 small white, elongated eggs singly on the leaf blades of rice seedlings
in the nursery. The incubation period is 7 days. The adult flies live for a fortnight.
The maggots are white, translucent, and about 1 mm long. The anal segment is bifid. The
larvae migrate to the central whorl of the rice plant and start feeding. The larval stage lasts about 6
weeks. Pupation takes place between the leaf sheath and stem and the pupal period is 2 weeks.
There are remarkable variations in the growth period of the insect in different regions of Japan
(Tamura et al. 1959). The flies of the later generations oviposit on grasses where the emerging
larvae overwinter.
2.10 Rice Seedling Flies, Atherigona exigua Stein, and Atherigona oryzae Malloch (Fig. 2.10)
(Diptera: Muscidae)
Distribution: Bangladesh, India, Indonesia, Japan, Malaysia, Pakistan, Papua New Guinea,
Philippines, Singapore, Sri Lanka, Thailand
Host plants other than rice: Cynodon dactylon (L.) Pers., Panicum repens L., Sorghum bicolor (L.)
Moench., Zea mays L.
Fig. 2.10
Damage by seedling maggots is highly seasonal in occurrence. It is restricted to upland rice and
does not occur in flooded wetlands.
The maggots feed in the central shoot of rice seedlings by tunnelling downward to the base.
They move their mouth hooks back and forth in a rasping motion. They produce typical symptoms
of twisted deadhearts similar to those caused by lepidopterous stem borers. The attacked plants
turn yellow, become stunted and may subsequently die. Maturity of plants which survive is delayed
by 7 to 10 days.
The adult is a small fly, about 3 to 3.5 mm long, with a grey thorax, yellow spotted abdomen and
yellow legs. The flies have an angular head with deep set antennae. Adults are strong fliers and are
active only during the daytime. They are attracted to plants less than a month old for oviposition and
a female may lay 100 eggs during its lifetime. Eggs are laid singly on the underside of the leaves 3
to 5 days after mating, usually in the evening hours. Eggs are pure white, elongated and very large
(1.5 mm long) in comparison with the adult. Incubation period is 3 days.
The newly-emerged larva is creamy white and moves down the leaf blade on a film of dew in the
early morning. The maggots then burrow the leaf sheath towards the base of the stalk. They molt
three times in 6 to 10 days and then pupate inside the stem or in the soil within a small yellow
puparium. Pupae are barrel-shaped and reddish brown in color and turn to dark brown at the later
stages. The pupal period lasts 8 days. Adult emergence can take place any time of the day, but it
occurs mostly in the afternoon (Rodriguez 1969). Adult longevity is 10-12 days.
Damaging infestation levels occur several months after the beginning of the rainy season. Peak
incidence of the pest in India was reported during the months of July and August (Senapati and
Satpathy 1983). The infestation of A. oryzae was most serious on rice after a prolonged period of
drought (Fernando and Manickavasagar 1957).
2.11 White Grubs (Coleoptera: Scarabaeidae)

White grubs, the large larvae of scarab beetles, are becoming important pests of upland rice in many
parts of the world. As they are mostly polyphagous, various other crop plants are also attacked by
these insects.
White grub species are divided into two groups: (1) ‘chafers’ in which adults feed on the foliage
of trees and larvae feed on roots of living rice plants, and (2) the ‘black beetles’ in which adults feed
on roots and larvae survive on organic matter and do not attack living roots.
The larvae of chafers feed on roots and other subterranean plant portions of rice and many
other crop plants. As the grubs grow, they become voracious feeders causing enormous damage to
crops. In case of heavy root injury, the whole plant is affected, while partial damage leads to poor
growth of host plants. A single white grub is capable of destroying a number of plants. However,
in the older crops, plants are not killed outright. In such cases, yellowing followed by withering of
the leaves takes place. Clustered distribution is a common feature of white grubs and infestations
often tend to occur in the fields year after year (Veeresh 1977).
DALE 381
Fig. 2.11
Sporadic occurrence of chafers such as Anomala dimidiata, Holotrichia seticollis, H. longipen-

nis and Popillia cupricollis has been reported on rice crops from India. Grubs of P. cupricollis damage
roots of rice seedlings while adult beetles chew the rice grains at the dough stage. Severe outbreaks
occurred over large areas in northern India in 1981 and more than 50% of the crop was damaged
by white grubs. Leucopholis irrorata (Fig. 2.11), another chafer species, caused intensive but highly
localized damage to upland rice in southern Luzon, Philippines, in 1974 (IRRI 1976). The distribu-
tion of this pest in the Philippines is closely related to sugarcane cultivation (Litsinger et al. 1983).
Heteronychus lioderes, a black beetle, is a specific pest on rice in Uttar Pradesh, India (Garg
and Shah 1983). H. plebeins damages the rice crop in Madagascar in humid soil during the dry
season. H. mosambicus was recorded as damaging the roots of rice plants in Nyasaland. It occurs
only in those fields from which water has receded and the surface remains mostly damp.
Rain at the beginning of the rainy season stimulates the overwintering adults of most species
back to activity. But adults of Leucopholis irrorata in the Philippines revert to normal activity only
a month after rains begin (Litsinger et al. 1983). They emerge from the soil leaving behind a small
hole on the surface. The beetles then undertake feeding and a mating flight during night and return
back to the soil before dawn. They are found in soil up to a depth of 0.3 meter (Rai et al., 1969).
Adult chafer beetles are grey, tan, dark brown or black. The males are considerably smaller
than females. The male Leucopholis irrorata is 25 mm long and 14 mm broad; the corresponding
measurements for the female being 30 mm and 16 mm respectively (Otanes 1924). The beetles feed
on leaves and fruits of various trees. Besides being a source of food, these trees serve as mating
sites. Beetles, after mating, remain in the trees during the day and at night fly short distances to lay
eggs in nearby fields. Hence more egg laying and a higher level of infestation are noticed in fields
near trees. Adults are attracted to light traps and catches are the largest during new moon nights.
Eggs are laid in the soil at a depth of 5 to 15 cm. They are creamy white and oval when freshly
laid and measure on an average 3 mm long and 1.7 mm broad. The total number of eggs laid per
female varies from 50 to 60. Color changes as the development advances and the egg swells up to 4
mm in diameter, one or two days before hatching. Eggs are highly sensitive to dry weather and must
be in moist soil to hatch. The egg stage lasts for 1 to 3 weeks.
The newly hatched grub is creamy white in color. The head turns brown and the larva becomes
active in a few hours. The first instar larva can feed on the organic matter available in the soil but
prefers plant roots. The second and third instar grubs require roots as their food for normal
development and pupation. Larvae prefer moist soil. In moist soil, the grubs are seen near the soil
surface whereas in dry soil they remain close to a location in the soil where moisture is suitable. The
larva can survive for more than 110 days within the earthen cell under low soil moisture levels but
cannot survive for more than six days under water-saturated conditions (Veeresh 1977). In nature,
continuous rain or a water-logged condition in fallow land for 3 to 4 days forces the grubs to come
out of the soil where they are exposed to dehydration and predation by birds, The uneven
distribution of white grub larvae within a field is due to their strict moisture requirement.
There are three larval instars. The stadia are 1 to 2 weeks, 3 to 4 weeks and 4 to 8 weeks for
first, second and third instars respectively. In the tropics, the grubs feed on the roots of host plants
from July to September. The maximum damage is usually done from the middle of August to early
September when the grubs are fully grown. Mature larvae descend to a depth of 0.3 to 2.0 m in the
soil during the dry season where they prepare an earthen cell and pupate. The pupal cells protect
the dark brown pupae from drying out. Adult emergence starts by the last week of September and
is completed by the end of November.
The adult beetles remain inactive within their pupal cells till the next April when they begin to
leave the soil. There is only one generation of the pest in a year. The life cycle is completed in 11 to
16 weeks in the tropics. In temperate areas 2-year life cycles are common.
The larvae are unable to tolerate low temperatures. The third instar grubs kept at constant
temperatures of 4, 10, 15, 20, 25 and 30°C did not survive for more than a month except at 20 and
25°C (Veeresh 1977).
2.12 Grape Colaspis, Colaspis flavida (Say) (Coleoptera: Chrysomelidae)

Distribution: Southern USA
Host plants other than rice: Chenopodium album L., Digitaria sanguinalis (L.) Scop., Fragaria
vesca L., Glycine max (L.) Merr., Iva Ciliata Willd., Lespedeza striata (Thunb.) Hook
& Arn., Muhlenbergia mexicana (1.) Trin., Paspalum laeve Michx., Poa compressa
L., Polygonum lapathifolium L., Potentilla bicolor L., Pyracantha coccina Roem.,
Rumex crispus L., Vitis vinifera L.
Adults and grubs of C. flavida are polyphagous and a number of important cultivated plants are
their hosts. The insect first drew the attention of economic entomologists as a pest of grapes, and
thence the common name. Rice is the spring host of the overwintering larvae. Damage is mostly
DALE 383
Fig. 2.12
confined to dry-seeded fields. In the spring the overwintering grubs feed on germinating seeds,
roots and seedlings reducing the crop stand and reducing tillering.
First brood beetles appear in the field in early June and are abundant up to early July. They
mate soon after emergence and may mate several times during their life. The average longevities of
the males and females are 7.8 and 13.4 days respectively (Rolston and Rouse 1965). The preoviposi-
tion period lasts for 3 to 5 days. Eggs are laid in the soil around the roots of host plants. The
incubation period is 6 to 9 days.
Larvae spend up to 7 months in cells pressed slightly into or on the soil. Pupation occurs in the
soil and the pupae remain in the soil for 3 to 7 days until adults emerge. Highest larval populations
are seen in less compact soils. Beetles are unable to come out from dry, hard soil.
The grape colaspis overwinters as larvae near the boundary of the subsoil and topsoil. The
larvae suffer considerable mortality from spring tillage. The surviving larvae move upward and start
feeding about the first week of May. Pupation begins in late May and continues until late June. The
first brood of adults starts ovipositing from about the beginning of June until mid-August. Some
larvae may not pupate until the following spring; others pupate in late July or August and produce
a second brood of adults. These adults are responsible for the overwintering larvae that develop in
late fall.
CoIaspis occidentalis (L.) from Guyana and C. prasina (F.) from Brazil are of relatively minor
importance.
2.13 Rice Root Weevil, Echinocnemus oryzae Marshall (Coleoptera: Curculionidae)

Distribution: India
Host plants other than rice: FimbristyIis tenera Roem. & Schult. and many wild grasses
The rice root weevil, E. oryzae, previously a minor pest of paddy has become an important pest in
some river basins in India. Adults feed on leaves of newly transplanted rice, but seldom cause
economic damage. Larvae feed on roots during the wet season. Plants attacked during tillering
show more damage symptoms than plants damaged after tillering. Yield losses may reach to 10%.
Fig. 2.13
It is probable that the insect is primarily a pest of grasses and that it has only secondarily taken to
rice. The distribution of the pest is rather restricted to deltaic regions that have recently been
brought under rice farming (Singh and Kalkat 1956). Clay and heavy loam soils favor the incidence.
The insect does not seem to occur in sandy soil (Thirumala Rao 1952).
Eggs are laid at the base of plants. Grubs devour the fibrous roots of rice plants and attacked
plants become stunted and tillering is reduced. Damage is maximum in the newly transplanted crop
during July-August. In severe cases the plants wither and the field has many patches that have to be
filled by transplanting fresh seedlings. Once the plant is established, the chances of withering are
low.
Adult weevils are black or piceous and densely covered with grey scales. They emerge with the
onset of the monsoon towards the end of May. The beetles are usually found on grasses and after
pairing, give rise to a new generation of grubs that can be seen among rice roots from July to
September.
Larvae are translucent white and about 5 mm long with six pairs of prominent tubercles on the
dorsal side of the abdomen. These structures serve a respiratory function by taking in oxygen from
the roots. The larvae burrow down into deeper layers of soil to remain in a resting condition and
later pupate. There are two generations per year.
2.14 Rice Plant Weevil, Echinocnemus squameus Billberg (Coleoptera: Curculionidae)

Synonym: Echinocnemus bipunctatus Roelofs
Distribution: China, Indonesia, Japan, Korea, Ryuku Island, Taiwan
Host plants other than rice: Alisma plantago-aquatica L., Callitriche verna L., Cyperus rotundus L.,
Echinochloa colona (L.) Link, Potamogeton natans L.
E. squameus is a sporadic minor pest of rice. Adult weevils feed on the leaves of young rice plants
at or near the water surface. Feeding damage is evident by a few longitudinal scars that are visible
on the leaves when they open. Maximum damage is caused to plants 3 to 4 weeks after transplant-
DALE 385
Fig. 2.14
ing. The infected plants are stunted and do not tiller normally. Sometimes irregular and delayed
flowering also occurs (Kojima et al. 1981). In severe cases, the affected plants break at the site of
insect feeding due to wind or submergence. Larval injury on roots is seldom noticed (Kuwayama
1963).
The adult weevils emerge from hibernation and migrate to rice fields by the end of May. In
Japan migration to the rice fields coincides with the transplanting time. Plants adjacent to levees
are most prone to damage by the beetles. The beetles walk on the surface of water and dive under
water or swim. Females dive under water and lay their eggs singly on soil close to the rice plant.
Water is indispensable for the development and hatching of eggs (Kuwayama 1963). The incuba-
tion period is 6 to 10 days.
The newly hatched grubs burrow into the soil. The spiracles are modified to six paired dorsal
hooks on the lateral side of the larvae which under flooded conditions, are thrust into roots to
obtain oxygen. Larval feeding occurs on decayed organic matter and rarely on plant roots. The
full-grown grubs usually pupate by the end of September, but are unable to pupate under flooded
conditions (Oya and Sato 1978). Overwintering sites are low-lying areas and stubbles in rice fields.
But adults often hibernate on the roots of grasses on levees. There is one generation annually.
2.15 Paddy Root Weevil, Hydronomidius molitor Faust (Coleoptera: Curculionidae)

Distribution: India
This pest is a serious problem to rice cultivation in many parts of Bihar, Gujarat and Haryana states
in India. Damage up to 30 to 50% has been reported from Haryana (Kushwaha and Sharma 1980).
Even though adult weevils have been observed feeding on rice leaves, the damage caused by them
is negligible.
Fig. 2.15
Grubs feed on the roots of young transplanted rice plants. Regenerating roots are also
subsequently destroyed by larval feeding. The attacked plants become stunted and tillering is
reduced. The leaves turn yellow, develop a rusty appearance and in severe cases, plants are even
killed (Kushwaha et al. 1983). But once plants are established, larvae cause little damage.
The adults start emerging after the first showers in June. The adult weevil measures 4.5 mm in
length and 2.2 mm in width. Freshly emerged weevils are light brown in color, changing to dark
brown to black at later stages. Adults are semi-aquatic and are capable of swimming in water. They
mate 3 to 4 hours after emergence and females lay their eggs on the soil near the roots of grasses.
The eggs hatch in 7 days.
Grubs begin hatching in early July. They are translucent white in color. Grubs crawl down to
the root zone and first feed on the root hairs and later on the roots. The maximum larval population
is observed in late July which coincides with the main transplanting season. The active feeding stage
lasts from August to September, after which the larvae move down into deeper layers of soil where
they remain in an inactive resting stage during the winter and summer. Most of the grubs occur at a
depth of 25 to 30 cm (Singh and Chaudhary 1968).
There is a short pupal stage of 10 to 15 days before adult emergence in June. There appears to
be only one generation of this insect in a year.
2.16 Rice Water Weevil, Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae)

Distribution: Japan, Korea, USA
Host plants other than rice: Many plants belonging to the families Alismataceae, Commelinaceae,
Cyperaceae, Gramineae and Juncaceae
The rice water weevil is the most important pest of rice in the USA. A parthenogenetic strain of the
pest is believed to have been introduced into Japan from California in 1976. Now L. oryzophilus is
regarded as one of the most destructive pests of rice in Japan and probably one of the most difficult
to control (Okada 1982).
DALE 387
Fig. 2.16
The adult feeds on the leaf epidermis of young rice plants. Adult damage is generally of minor
importance and is usually confined to the edge of fields, although Ingram (1927) has reported plant
mortality due to it in some late-planted rice fields in the USA. Weevils have also been reported to
feed on panicles, consuming floral parts or the endosperm of the developing rice grain. The larvae
cut the roots of rice plants. This results in the stunting of plants, delayed maturity and loss in grain
yield. Heavy larval infestation may reduce plant vigor and cause lodging of plants prior to harvest.
Yield losses up to 75% have been reported (Newsom and Swanson 1962, Grigarick 1963) due to the
infestation of the water weevil.
The adult weevil is about 5 mm long, olive-grey to tan, with a dark V-shaped area on the elytra.
In the females, this area is more distinct and the abdomen more swollen.
Oviposition begins after the field is flooded. Semi-aquatic adults usually fly into the standing
rice crop at night and then move from plant to plant by swimming just below the water surface. The
gravid female moves down the stem and lays eggs in or under the basal submerged portion of leaf
sheath tissue and, rarely in the roots. Maximum egg-laying occurs one or two weeks after flooding.
The egg is white, elongate and slightly curved. It is about 0.8 mm long. The egg stadium lasts
for 4 to 9 days depending on temperature (Raksarart and Tugwell 1975).
Larvae are white, very small and legless. The head is brown, and small in relation to the rest of
the body. The first instar grubs mine in the leaf sheath for about a day and then move down the plant
to the soil where they start feeding on roots. Larvae possess dorsal hooks that are modified
abdominal spiracles. These structures facilitate larval movement in the soil and help in the
acquisition of oxygen from the root aerenchyma (Everett 1966). There are four larval instars (Cave
and Smith 1983). Larvae attain a maximum length of 8 mm in a period of approximately 21 days.
Pupation takes place in oval mud cells lined with a water-tight material which is attached to the
roots. The pupa is white and of the same size as the adult weevil.
In Japan, only one or sometimes two generations are recorded in a year (Tsuzuki et al. 1982).
But in southern Louisiana, USA under optimal conditions, up to 4 generations occur. Successive
generations take place within the same rice field only when there is no crop in the seedling stage in
the area.
By the end of August, the weevils move to overwintering sites such as woodland leaf litter,
clumps of grass, or rice stubbles. Adult overwintering populations are generally higher in moist
places than at dry sites. The weevils come out of overwintering areas and start feeding on grasses
from late April to early May. They then fly into rice fields that are transplanted in May. The period
of oviposition of the water weevil is mostly governed by the duration of day light in the locality.
The population densities in all stages of the pest are higher in those rice fields with standing
water throughout the cropping season than in fields that are only intermittently irrigated (Yasuda
et al. 1979). Shimohata and Kano (1982) observed that the weevils feed more and produce more
eggs on younger rice seedlings.
The critical minimum temperature for adult female survival in Japan was found to be as low as
–5 to –10°C (Tsuzuki et al. 1979). This demonstrates the ability of the pest to survive in low-
temperature regions and to cause damage to rice crops there.
Lissorhoptrus brevirostris, another species of water weevil, has been reported from Cuba
(Carbonell and Ravelo 1979). The damage caused by adults is not economically important. But the
larvae can destroy up to 83% root tissue and cause 54% yield loss (Carbonell 1982). The pest
population is the highest during May to October when the temperature ranges between 20" and
27.5°C and the rainfall is higher than 100 mm. There are two generations in a year with peak
populations in June and September.
3. STEM BORERS
Rice stem borers are a key group of insect pests, mostly belonging to the two lepidopteran families
of Pyralidae and Noctuidae. According to Pathak (1968), pyralid borers are the most common and
destructive of all stem borers and usually exhibit a high degree of host plant specificity. The noctuid
borers, on the other hand, are polyphagous and only occasionally cause economic losses. A list of
common stem borers is provided in Table 3.
in Asia, Scirpophaga incertuIas and Chilo suppressalis are the major stem borers and are widely
distributed from India to Japan. Maliarpha separatella, Chilo zacconius and Sesamia calamistis are
the most common species in Africa. Diatraea saccharalis and Rupela albinella are the important
stem borers of rice in the western hemisphere even though their damage is less severe. In Australia,
rice is attacked by a species of Phragmatiphila (Hely 1958).
Stem borer larvae start their attack by boring the inner portion of the leaf sheaths. At this early
stage, the symptom of damage is the longitudinal yellowish white patches at the feeding sites on the
leaf sheath. The subsequent boring of the stem by caterpillars often leads to the severing of the
apical plant parts from the base. As a result, the central leaf whorl does not open, turns brown, and
DALE 389
TABLE 3
Common species of rice stem borers (Pathak 1975)
Scientific name Common name Distribution
Ancylolomia chrysographella Koll. – Southeast Asia

Chilo agamemnon Blez. Small purple-lined borer Africa
Chilo auricilius (Dudgeon) Gold-fringed rice borer Indian subcontinent, Southeast Asia
Sugarcane internodal borer
Chilo loftini (Dyar) – Mexico, USA
Chilo partellus (Swinhoe) Pink borer Oriental and Ethiopian regions
Stem borer
Sorghum stem borer
Chilo phaeosoma (Martin) – Ethiopian region
Chilo plejadellus (Zinken) Rice stalk borer Mexico, USA
American rice stem borer
Chilo polychrysus (Meyrick) Dark-headed stem borer Indian subcontinent, Southeast Asia
Rice stalk borer
Chilo suppressalis (Wlk.) Striped stem borer Asia, Italy, Spain
Rice striped borer
Pale-headed striped borer
Rice stem borer
Asiatic rice borer
Chilo zacconius Blezynski African rice borer Africa
Diatraea saccharalis (Fabricius) Sugarcane borer Southern USA, Central and South
American sugar cane borer America
Small moth borer
Elasmopalpus lignosellus (Zeller) Lesser corn stalk borer Neotropical, Trinidad, Brazil
Maliarpha separatella (Rogonot) White borer Africa
Niphadoses gilviberbis (Zeller) – Burma
Niphadoses palleucus Common – Australia
Phragmatiphila sp. – Australia
Proceras indicus (Kapur) Internodal borer Indian subcontinent
Rupela albinella (Cramer) South American white borer South America
Scirpophaga incertulas (Wlk.) Yellow stem borer Asia
Yellow rice borer
Yellow borer
Scirpophaga innotata (Wlk.) White rice borer Asia, Australia
White stem borer
Sesamia botanephaga – Ethiopian region
(Tams & Bowden)
Sesamia calamistis (Hampson) African pink borer Africa
Mauritius pink borer
Sesamia inferens (Walker) Pink stem borer Asia
Ragi stem borer
Purple stem borer
Violet stem borer
Zeadiartaea lineolata (Wlk.) Neotropical corn borer Venezuela
Maize borer
dries. The lower leaves, however, may still look green and healthy. This condition is known as the
“dead heart” symptom and the affected tillers do not bear panicles.
If the detachment of growing plant parts takes place after panicle initiation, the panicles may
not emerge. Those that have already emerged may not produce grains. Such panicles are con-
spicuous in the field by bearing empty grains, being erect and white. They are called “white heads.”
The plants can compensate for a low percentage of early dead hearts by putting forth additional
tillers. But for every percent of white head, 1 to 3% loss in yield may be expected (Pathak 1975).
The bionomics and ecology of a few important stem borer pests are dealt herein.
3.1 Gold-Fringed Rice Borer, Chilo auricilius Dudgeon (Lepiodoptera: Pyralidae)

Synonyms: Chilotraea auricilia (Dudgeon),
Diatraea auricilia (Dudgeon)
Argyria sticticraspis Hamps.
Distribution: Bangladesh, Bhutan, Burma, China, Hong Kong, India, Indonesia, Malaysia, Nepal,
Philippines, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Saccharum officinarum L., Sorghum bicolor (L.) Moench., many grasses
Chilo auricilius is a serious pest of sugarcane in India, Taiwan and China. It has recently become
more abundant on rice plants and is becoming a major pest in Orissa and West Bengal in India and
in Bangladesh., Damage caused by this pest was reported to be around 30% loss in grain yield in
Orissa (Rao 1964) and 20% in Bangladesh (Alam et al. 1964). The following description on the
bionomics of the stem borer is mostly based on studies conducted by Rao and Rao (1980) in India.
Moths mate between 8 and 9 p.m. Gravid females lay eggs in masses on foliage, mostly on the
undersurface of rice plants. Occasionally a few egg masses are laid on leaf sheaths also. Average
oviposition period is 3 days and the maximum number of eggs is laid on the first day of egg-laying.
The average number of eggs laid per female was 123 during the entire oviposition period in an
average of 9.5 egg masses. Eggs hatch in 5 to 7 days.
Larvae have only four stripes (the dorsal stripe being absent), and their body is proportionately
wider than in C. polychrysus, and C. suppressalis from which the species is separated only by slight
differences in the thoracic chaetotaxy. The caterpillar molts six times before becoming a pupa. The
larval period is about 30-32 days.
Pupation takes place in the affected stems. The postero-lateral and postero-dorsal spines on
the cremaster of the pupa are more prominent than on the other Chilo species. The pupal period
lasts for 6 days.
Four generations occur in Taiwan and China, and 5 to 7 in India. The insect is active during
and just after the monsoon.
3.2 Spotted Stem Borer, Chilo partellus (Swinh.) (Lepidoptera: Pyralidae)

Synonym: Chilo zonellus (Swinh.)
DALE 391
Fig. 3.2
Distribution: Afghanistan, Comoro Islands, India, Kenya, Malawi, Nepal, Pakistan, Sikkim, Sri
Lanka, Sudan, Tanzania, Thailand, Uganda
Host plants other than rice: Eleusine coracana (L.) Gaertn., Saccharum officinarum L., Sorghum
bicolor (L.) Moench., Triticum aestivum L, Zea mays L., wild grasses
The insect is primarily a pest of maize and sorghum. But as a rice pest it is particularly important
in eastern India. It rarely occurs in lowland irrigated conditions, but in upland rainfed conditions it
is often abundant.
Moths are yellowish to yellowish brown with a slender body. Forewings terminate with an acute
tip, and are straw-colored with one or two transverse rows of small dark brown dots. Hindwings are
white with a marginal fringe. Usually male moths are darker and smaller than the female.
Eggs are laid in two overlapping rows on all parts of the plant. They are oval, flat and whitish.
About 300 eggs are laid by a single female. Hatching begins after 3 to 5 days.
Young larvae feed on leaves for about 5-6 days after which they penetrate the midribs of leaves.
The larvae are dirty white with setae borne on each segment. There are two dorsal stripes which
may merge to form a wide band and lateral stripes. The larvae enter the stem directly or through
the growing point. They feed inside the stem until they become fully grown after 18-30 days.
Pupation takes place within the stem and lasts for 6-12 days. Pupae are slender, 10-15 mm long
and shining brown. The moths emerge usually just after sunset.
During the winter, older larvae remain dormant in the stems and stubbles. In the tropics,
continuous development occurs during the rainy season. In places where there is a distinct dry
season, the pest enters a larval diapause after the rainy season. Diapausing larvae pupate after the
first showers and the moths emerge a few days later.
3.3 American Rice Stalk Borer, Chilo plejadellus Zincken (Lepidoptera: Pyralidae)
Distribution: Mexico, USA (Arkansas, Louisiana, Texas)
Host plants other than rice: Spartina cynosuroides (L.) Roth., Zizania aquatica L., Zizaniopsis
miliacea (Michx.) Doell and Aschers
Fig. 3.3
Dr. L. O. Howard first observed this pest on rice near Savannah, Georgia, in 1881 and described it
as a new species (Riley 1883). The stalk borer has a restricted distribution and serious infestations
are only sporadic.
Moths have pale straw-colored forewings with a sprinkling of minute black dots. The hind wings
are white. Moths are lighter in color than Diatraea saccharalis. Eggs are deposited in clusters of 50 or
more with the individual eggs overlapping. They are oval, flat and nearly white when freshly laid.
The larva is yellowish white and is marked on each side with a brown stripe running the entire
length of its body and a fainter brown stripe below it. The larva is about 25 mm long when full
grown. The larvae feed on the inner tissues of rice stem causing deadhearts and whiteheads. Newly
hatched larvae have also been known to attack at the uppermost node after the panicle has already
emerged and partially developed. These panicles may break at the point of injury.
Pupation takes place in the spring, the larvae having spent the winter in the stubble. The pupa
is light to dark brown. It is smooth, tapers to a point at the rear and is nearly always found enclosed
in a heavy web inside the stem.
3.4 Dark-headed Stem Borer, Chilo polychrysus (Meyrick) (Lepidoptera: Pyralidae)

Synonyms: Chilo polychrysa (Meyrick)
Chilotraea polychrysa (Meyrick)
Diatraea polychrysa (Meyrick)
Proceras polychrysa (Meyrick)
Distribution: Burma, China, India, Kampuchea, Laos, Malaysia, Philippines, Thailand, Vietnam
Host plants other than rice: Echinochloa crus-galli (L.) Beauv., Hymenachne acutigluma (Steud.)
Gilliland, Saccharum officinarum L., Scirpus grossus L., Setaria pallide-fusca
(Schumach.) Stapf and C., E, Hubb., Zea mays L.
Chilo polychrysus is a major pest of rice in Malaysia and parts of India. Before 1960, it was the most
dominant species in Peninsular Malaysia but since the introduction of double cropping, the yellow
DALE 393
Fig. 3.4
borer is gradually taking its place. In South India, it was first seen attacking the rice crop during the
1955-56 cultivation season and many serious infestations subsequently followed.
The caterpillars attack all stages of the rice crop. The young plants, when attacked, are killed
in a short period having no chance of recovery. In the case of mature plants, the infestation results
in the formation of deadhearts and whiteheads. A heavy attack of this borer may cause a loss in
yield of up to 60%.
The adult moth is light brown with 6-7 tiny black dots and silver sides at the tip of the forewings.
The hind wings are yellowish white. It has a body length of 10-13 mm and wing span of 16 to 25 mm.
The moths usually live for 2 to 5 days.
The moths are active during the night and oviposition takes place between 7 to 11 p.m. Eggs
are laid in batches of 20 to 150 in longitudinal rows along shallow furrows on both surfaces at the
basal portions of the leaves. They overlap each other and are scale-like in appearance. A single
female moth can lay up to 480 eggs in 3 days. The egg is white when laid but turns black as it nears
hatching. The fully-developed embryo eats its way out of the egg shell, usually in the morning hours.
The incubation period is 4 to 7 days.
The newly-hatched larvae are greyish white and 1.3 mm long. They are distinguishable from
other species by the head and prothoracic shield which are distinctly black. The first instar
caterpillar moves down to the leaf base and bores into the outer leaf sheath. Sometimes the larva
enters the midrib of a leaf and tunnels down into the leaf sheath. The attacked leaf sheath first
appears yellow and then gradually dries up. If the rice plant is thick, stout and succulent, the whole
larval and pupal existence are completed within the peripheral leaf sheaths sparing the central culm.
On the other hand, if the outer leaf sheaths are thin or dry, the caterpillar bores into the central
shoot. One larva is usually found in a single stem. If the host plant is dead as a result of infestation
or when it becomes congested with larvae, the larvae migrate to neighbouring plants. In such cases,
the points of entry are indicated by plugs of fecal matter.
The last instar larva measures about 21 mm in length. The head is brownish black while the body
is creamy white with five distinct brown stripes along the back of the body. The head and body are
sparsely clothed with inconspicuous hairs. The full-grown caterpillar, just prior to pupation, makes
an exit hole below which the feeding tunnel is cleared and lined inside with a loose layer of silken
frass. There are 6 larval instars, and the larval period ranges from 20 to 41 days.
The newly-formed pupa is dirty white to light straw-colored with the larval body stripes clearly
visible. The color subsequently turns deep brown to black. The pupal period lasts for 4 to 6 days.
No dormancy or diapause has been reported. The number of generations in a year varies up to
12 depending on the climate and availability of host plants. Overlapping generations may occur
throughout the year. The second and succeeding generations are usually more destructive. In the
absence of rice plants, the larvae move on to alternate hosts.
3.5 Rice Striped Borer, Chilo suppressalis (Walker) (Lepidoptera: Pyralidae)

Synonyms: Crambus suppressalis Wlk.
Chilo simplex (Butl)
Chilo oryzae Fletcher
Distribution: Bangladesh, Bhutan, Burma, China, Hawaii, India, Indonesia, Iran, Italy, Japan,
Kampuchea, Korea, Laos, Malaysia, Nepal, Pakistan, Papua New Guinea, Philip-
pines, Spain, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Echinochloa colona (L.) Link, Echinochloa crus-galli (L.) Beauv.,
Phragmites australis (Cav.) Trin.ex Steud., Sclerostachya fusca (Roxb.) A. Camus,
Typha latifolia L., Wild rices, Zea mays L., Zizania aquatica L.
The rice striped borer, Chilo suppressalis has been considered as one of the most serious pests
of rice in temperate and subtropical Asia. However, its importance is gradually declining in many
countries. The pest mainly attacks rice in the middle or late stage of plant growth. In addition to
causing dead hearts and white heads, larval feeding can also cause reduced plant vigor, fewer tillers,
unfilled grains and lodging of plants. High grain yield losses have been reported for this pest; up to
100% grain loss has been reported from Japan (Logothetis 1951), 50% in the Philippines (Otanes
and Sison 1947) and 20% in Taiwan (Ou 1959). In Spain, Chilo suppressalis, coupled with Sesamia
sp., produces an estimated crop loss of about 2% (Grist 1965), but in Malaysia, no serious damage
has been reported (Van Vreden and Ahmadzabidi, unpublished).
Fig. 3.5
DALE 395
The adult moths are 13 to 16 mm long and with straw to light brown forewings. There are a
number of silvery scales and usually five black dots at the tip of the forewing; the hind wing is
yellowish white. The females are lighter in color than the males. Moths emerge at about 7 to 8 pm
and become active in the early morning. During the day they remain in dense foliage. Moths are
capable of flying long distances; sometimes winds carry them to distant places.
Male moths are strongly attracted to virgin females, the attraction being maximum on the
evening of their emergence but declining as days pass. Mating usually takes place during the night
of emergence, but egg-laying starts only on the next night. The mating frequency is maximum among
2- or 3-days old moths and it then decreases rapidly with age (Kanno and Sato 1978). Through
laboratory studies, it has been shown that the threshold light intensity for mating initiation varies
with temperature (Kanno 1980). Male and female moths mate as often as 8 and 4 times respectively,
in their life times.
A female moth may lay 100 to 550 eggs in batches of 50 to 80, depositing one batch each night
over a 3 to 5 day period. Eggs are naked and scale-like, pale to dark yellow, laid in masses formed
of overlapping rows. Egg masses are found on the basal half of the leaves or occasionally on the leaf
sheaths along the midrib of either the upper or lower surface. The eggs hatch in 3 to 5 days.
Maximum hatching takes place in morning between 5 to 6 a.m. Generally, all eggs of a mass hatch
together.
The larvae live gregariously during the first three instars (Nozato 1981). If the young larvae are
isolated from each other, they suffer high mortality. The newly emerged larvae crawl up the rice
plant and then congregate beneath the leaf sheath. All larvae enter the stem through a common
hole. Larval feeding generally occurs in the middle of the stem. The fully-grown larva is ap-
proximately 26 mm long and 2.5 mm wide. It has a yellowish brown head and has three dorsal and
two lateral brownish abdominal stripes. The middle dorsal stripe is lighter in color. The larval stage
normally lasts from 20 to 48 days.
Nozato (1982) has shown that larval mortality in the first generation is density-dependent. High
mortality among older larvae may be due to the destruction of rice stems through feeding.
Pupation takes place within the stem. The last instar larva makes an exit hole in the internode
for the moth to emerge. The pupa is reddish brown in color and is without a silken cocoon.
In temperate regions, the larvae enter into diapause in the winter and development resumes in
the spring with warmer temperatures. Temperature, daylength and the stage of growth of rice
plants are principal factors inducing diapause. Exposure of larvae to short daylengths (8 to 14
hours) induces diapause while longer days (14.5 to 16 hours) prevents it.
The rice striped borer is well adapted to a temperate climate. All stages of the pest can
withstand low temperatures and development proceeds normally in cold habitats of temperate
regions. The threshold temperature for egg development is 10 to 12°C. The incubation period
decreases with an increase in temperature up to 30°C. At 35°C, the eggs die. The optimum
temperature for larval development is between 22 and 33°C even though larvae are able to withstand
temperatures as low as -14°C for short periods of time. Under ideal ecological conditions, there are
5 to 6 larval instars while under adverse conditions, as many as 9 instars have been recorded. The
rate of pupal development increases with temperature from 15 to 30°C. Beyond 35°C, the pupae
suffer high mortality and the emerging moths are mostly deformed.
The pest can have one to five generations per year depending on the availability of host plants
and the occurrence of favorable temperature conditions. The longer the period of host plant
availability, the more the population is built up through increased generations in a year.
In Japan, distinct ecotypes of C. suppressalis have been reported. They are ‘Shonai’ in the north,
‘Saigoku’ in the southwest and ‘Tosa’ from Kochi prefecture. The Shonai ecotype is more tolerant
to lower temperature than the Saigoku ecotype. The stem borer population between the areas
distinctly occupied by these ecotypes is intermediate in nature.
3.6 African Rice Borer, Chilo zacconius Blezynski (Lepidoptera: Pyralidae)

Synonym: Proceras africana Auriv.
Distribution: Benin, Burkina Faso, Cameroon, Ivory Coast, Mali, Niger, Nigeria, Senegal
Host plants other than rice: Echinochloa crus-galli (L.) Beauv., Oryza barthii A. Chev., Sorghum
arundinaceum (Desv.) Stapf.
Chilo zacconius is a polyphagous pest of rice commonly found in West Africa. Adult moths have
pale yellow forewings with irregular black spots. Males are darker than the females. Moths are
nocturnal and hide during the day.
Oviposition takes place on the upper or middle leaves. Pale yellow eggs are laid in two or three
overlapping longitudinal rows. Eggs are oval, flat and imbricate like fish scales. They hatch after 4
days.
The newly-hatched larva moves around actively on the plant, feeds a little from the leaf and
bores into the stem through the leaf sheath. From flowering, the larvae bore into the flower stalk
Fig. 3.6
DALE 397
causing whiteheads. Then they descend into the central stem which they perforate at different
levels. The larvae can live for several months without food.
The larva has an ivory body with a dark brown head. There are seven longitudinal pink stripes
on the body, the ventral ones being incomplete and sometimes indistinct. The larvae pass through
5 instars in a period of 4 weeks before they pupate.
Pupation takes place in the stem or in the leaf sheaths. The chrysalis is dark brown; its abdomen
ends in a four pointed crest with another two-pointed crest at the back. The pupal period is 6 to 7
days.
There are 5 to 7 generations per year. The number is mostly governed by the duration of the
dry season and the availability of host plants. Generally, two successive generations occur in the
same rice field; the first generation larvae attack the stem and the second mostly infest the panicle
stalks.
A very similar species, Chilo diffusilineus J. de Joannis, has also been reported from the humid
equatorial Sudanese areas (Breniere 1976, Bonzi 1982). It differs from C. zacconius only in the
shape of genitalia in both sexes. The habits and appearance of the larvae are identical. C. dif-
fusilineus infests irrigated and rainfed rice plants. The exact distribution of the species is not known.
3.7 Sugarcane Borer, Diatraea saccharalis (Fabricius) (Lepidoptera: Pyralidae)

Distribution: Southern USA, Central and South America to Argentina, West Indian Islands
Host plants other than rice: Brachiaria mutica (Forssk.) Stapf, Leptochloa filiformis (Lam.)
Beauv., Paspalum urvillei Steud., Saccharum officinarum L., Sorghum halepense
(L.) Pers., Vetiveria zizanioides (L.) Nash, Zea mays L.
Diatraea saccharalis is the most damaging and the widely distributed stem borer in Latin America.
It is more numerous in Texas than the rice stalk borer Chilo plejadellus (Bowling 1967).
Adult moths have a wing span of 20 to 26 mm. The forewings are light brown to grey with less
continuous lines and a dark discal dot. The moths hide during the daytime under foliage. They are
Fig. 3.7
positively attracted to light. The adult population is maximum when the rice plants are at about 30
days after transplanting.
Oviposition takes place dying night. Eggs are laid on young leaves on both the upper and
lower surfaces. They are creamy white when laid but turn yellowish as the incubation period
advances. Eggs are deposited in groups of 2 to 100 or more, although the number of eggs usually
found in a single cluster is less than 50.
The larva is yellowish white with brown spots. In the winter these spots are absent and the color
becomes more deeply yellow. The newly hatched larvae move about and feed on rice leaves for 24
to 48 hours. They then crawl into the space between the leaf sheath and stem to make the entry hole.
Sometimes these larvae come out of the initial tunnels and make new attacks on the same plant or
search for other plants to infest. Normally 5 to 6 larval instars can be distinguished. The fully-grown
larva is 25-35 mm long and makes an exit hole for the adult moth to emerge.
Pupation takes place within the stem. The pupa is light brown and 10-12 mm long. It is naked
without being enclosed in a silken web.
The insect spends the winter as a larva in the stubble of host plants. The larvae pupate and
moths emerge in spring. The insects breed on alternate host plants until rice plants are big enough
to feed upon.
Four and a partial fifth generation appear each year in the cool northern and southern limits of
the pest’s distribution. In the tropics, as many as 7 generations develop.
3.8 White Borer, Maliarpha separatella Ragonot (Pyralidae: Lepidoptera)

Synonyms: Anerastia pallidicosta (Hampson)
Enosima vectiferella (Ragonot)
Ampycodes pallidicosta Hampson
Rhinaphe vectiferella Ragonot (Hampson)
R. pallidicosta (Hampson)
Distribution: African countries, Burma and China
Host plants other than rice: Andropogon tectorum Schum. & Thonn., Sorghum bicolor (L.)
Moench.
Among the four stem borers usually found attacking rice in Africa, the white borer M. separatella is
considered to be the most common. Due to its peculiar habit of remaining at the lower internodes
of the rice plant, the larva does not usually produce ‘dead hearts’ or ‘white earheads’. The growing
apical portion of the plant is not cut off from the base and hence panicles can be initiated at the last
node. The infestation mainly leads to a reduction in plant vigor and tiller number and a high
percentage of unfilled grains. The level of M. separatella infestation does not correlate with the
ultimate damage of the crop. At 75 days after transplanting, 97% hills were infested and 71% tillers
were damaged by the larvae. However, the incidence of ‘dead heart’ was as low as only 1% (Ho et
al. 1983). Apart from the direct damage, Pollet (1978) has reported a synergistic interaction
DALE 399
Fig. 3.8
between M. separatella and Pyricularia oryzae, the organism causing rice blast disease where the
fungus preferentially attacks plants already infested with the stem borer.
The adult moth is slender and straw colored with a prominent red-brown line along the anterior
margin of the forewing. The hind wings are yellowish white, with a metallic sheen, fringed with long
yellow hairs. The body length varies in the two sexes; 11 to 13 cm and 13 to 15 cm in male and
female, respectively.
The moths start laying eggs as soon as the rice seedlings are established at about 2-3 weeks after
transplanting. The eggs are laid on rice leaves in 3 to 6 parallel rows, each batch containing 30 to 95
eggs. Sometimes eggs are seen deposited within the youngest unopened leaf. The eggs are glued
on to the upper surface of the leaves by a sticky secretion which, on drying, completely encloses the
eggs. The eggs, white when they are laid, turn yellowish and darken before hatching. The incubation
period is 7 to 10 days.
The newly hatched larvae are white with dark brown heads. The caterpillars are very active and
disperse immediately after eclosion. They move briskly to the tip of the leaf blade from where they
suspend themselves by means of silken threads which they produce. Larvae drift in the wind and
attach themselves to the base of a rice plant and they bore into the stem. Dispersal is so efficient
that in about 24 hours after hatching, no more than one larva in each stem can be seen. The larvae
begin to feed on the inner tissues of the stem once they bore into the first or second internode. They
do not migrate from one tiller to another once the larvae have securely lodged themselves within the
stems. The larvae are not voracious feeders and throughout the larval stage they confine themselves
to one or two internodes. They usually can not survive in plants without internodes. The larva
passes through 5 to 7 instars before it pupates. The entire larval period which lasts for 35 to 50 days
is spent within a single rice stem.
Before pupation, the larva moves from the feeding site towards the upper part of the stem. An
exit hole is cut in the stem before the larvae spin a web channel connecting the inner wall of the stem
and the exit hole. The head region is placed upwards at a point 1 to 2 cm below the exit hole.
An important character of M. separatella is that it undergoes larval diapause lasting as long as
251 days (Akinsola and Agyen-Sampong 1984). In southern Nigeria, the resting stage starts in
July/August and lasts until March of the next year. Rainfall and duration of sunshine play a major
role in the initiation of diapause. Induction of diapause usually occurs before the rice plant dies and
starts drying. Diapausing larvae are sluggish, milky white and their body is wrinkled while non-
diapausing larvae are agile, yellowish white with a rather smooth body.
Among the different stem borers of rice, M. separatella is the major species in the upland rice
growing areas of Africa. In upland rice, the larvae are located more towards the base of the plant
while in flooded and swampy conditions, they tend to be located in the upper portion. In deep water
rice it is believed that the initial infestations occur before flooding. The infested tillers are detached
from their base (Akinsola 1980). The stem borer attack coupled with water pressure around the
base of the plant leads to the detachment of the affected part from the plant. There are usually 3 or
4 generations per year.
3.9 Yellow Stem Borer, Scirpophaga incertulas (Walker) (Lepidoptera: Pyralidae)

Synonyms: Tryporyza incertulus (Wlk.)
Schoenobius incertulas (Wlk.)
Distribution: Afghanistan, Bangladesh, Bhutan, Burma, China, Hong Kong, India, Indonesia,
Japan, Kampuchea, Laos, Malaysia, Nepal, Pakistan, Philippines, Ryuku Islands,
Sikkim, Sri Lanka, Taiwan, Thailand, Vietnam
The yellow stem borer Scirpophaga incertulas is a serious pest of rice throughout the Orient. It is
regarded as monophagous with exclusive host specificity to rice. However, some recent studies
(Zaheruddeen and Prakasa Rao 1983a, b) have shown that wild rices Oryza rufipogon, O. nivara,
Fig. 3.9
DALE 401
O. latifolia and O. glaberrima, and a grass weed Leptochloa panicoides can well sustain the borer
throughout its development and hence are to be considered as potential alternate host plants of
S. incertulas.
The yellow stem borer is the dominant and the most destructive stem borer species in India
(Walker 1975, Panda et al. 1976), Sri Lanka (Fernando 1967), Pakistan (Moiz and Rizvi 1971),
Bangladesh (Catling and Alam 1977), Thailand (Yasumatsu 1976) and Malaysia (Chang 1981).
However, in Japan, this species is currently limited to the southernmost regions of Kyushu (Kiritani
1981). In Taiwan, it is no longer considered as the most serious pest of rice, probably due to the
extensive use of organophosphate insecticides to control the borer (Chu 1971).
The extent of crop losses vary in time and space. S. incertulas was reported to cause 1 to 19%
yield loss in early-planted and 38 to 80% in late-planted rice crops in India (Khan 1967). A loss of
5 to 10% was reported in the Philippines (Rowan 1923). According to Walker (1975), the stem
borer causes 10 to 30% crop loss in Taiwan. In Malaysia, the yield reduction may amount to 10%.
Adult moths often exhibit sexual dimorphism. The female moth is bigger than the male and its
forewings are bright yellowish brown with a distinct black spot in the center. The abdomen is wide,
the tip being covered with tufts of yellowish hairs. The male moth is pale yellow; the abdomen is
slender and the anal end has a thin hairy covering dorsally. Spots on the forewing are not
conspicuous.
The moths usually emerge at night between 7 to 9 p.m. They are not active during the daytime,
but can be seen resting on stems and leaves. At dusk, especially on still, warm, humid nights, after a
rain, the moths mate. Mating generally occurs between 7 to 9 p.m. The female moths lay eggs early
at night in small masses covered with hairs and scales derived from the anal tuft. In the tropics, the
moths are short-lived and die 2 to 3 days after oviposition. But in Taiwan and Japan, they are
reported to live longer (Shiraki 1917); the longevity ranges from 5.3 to 8.8 days in females and 4.5 to
8.6 days in males.
Eggs, numbering 100 to 150, are laid in masses near the tip of leaf blades. They are creamy
white, flattened, oval and scale-like and covered with a tuft of tan anal hairs from the female moth.
Before hatching in 5 to 8 days, the eggs darken to a purplish tinge.
The first instar larvae are about 1.5 mm long and 0.5 mm wide. The body is pale yellow, the
prothoracic shield dark brown and prominent and the head orange. The newly-emerged larvae
show a strong tendency to disperse. They move downward and wander about on the plant surface
for one or two hours. They can also be seen hanging down by a silken thread to be carried by wind
to adjacent rice plants. During this roaming period, many larvae die. The survivors enter between
the stem and the leaf sheath and feed on green tissues of the leaf sheath for two to three days. The
caterpillars then start boring in the stem, often at the nodal region and feed on the inner tissues of
the plant.
Often the larvae leave the first host plant after a week. They move in search of other plants in
cylindrical cases made up of rolled leaf tips in which they drift on the irrigation water. After
reaching a suitable host plant, the larvae bore in, leaving their cases sticking to the stem at right
angles. In mature rice plants, the caterpillars bore into the stalk at the top region just below the
earhead. At this stage, a number of larvae are found in the same stalk (Banerjee and Pramanik
1967).
The full-grown sixth instar larvae are 25 mm long, white or yellowish white, and with a
well-divided prothoracic shield. They make a thin silken case over themselves inside the stem soon
after the prepupal molt. The larval period usually lasts for 30 days. Before pupation, the larvae
make an exit hole through which the adult moth later escapes.
Pupation takes place inside the stem, mostly in the lowest node of the plant or just above the
water level. Puttarudriah (1945) found that in seedlings, the larvae pupate in the root region only.
Pupae are pale at first and turn darker brown after some time. They are partly exarate, and the tips
of their appendages are free (Isaac and Venkatraman 1941). The pupal period is usually 6 to 10
days but may be prolonged in cooler months.
During periods when there is no rice crop and the temperature is not optimal for larval
development, mature larvae diapause in rice stubble. These larvae are vulnerable to low tempera-
ture and high moisture. Because the larvae cannot survive in the cut straw, they go down into the
stubble to diapause. In Japan, where plant stems are cut near the soil level at harvest, the stubble
harbors far fewer borers than in Taiwan, where longer stubbles are left when rice is harvested. In
Japan, as soon as warm weather begins in April-May, the larvae start pupating (Kawada et al. 1934).
In single cropping areas of tropical south India, some larvae enter into diapause after harvest in
December. These larvae pupate and emerge as moths after the monsoon rains begin in June or July
(Perraju and Reddy 1965).
Although eggs show some development at 13°C, they normally develop only at or above 16°C.
The optimum temperature is between 24 and 29°C. The humidity requirement for egg development
is between 90 and 100%, and hatching is drastically reduced when it falls below 70% (Doke 1936).
The threshold minimum temperature for larval development is 16°C. When reared at 12°C, the
second and third instar larvae do not molt and they die. The rate of larval development is positively
correlated within a temperature range from 17 to 35°C (Lin et al. 1959). The threshold for pupal
development is 15 to 16°C.
S. incertulas occurs year-round in Sri Lanka, south India and Malaysia where rice is cultivated
in succession. The number of generations in a year is governed mostly by the ecological conditions
and the availability of rice plants and varies from 2 to 6. In temperate countries like Japan, the pest
has only three broods. The third brood moths are generally more numerous than the first and the
second (Ishikura and Nakatsuka 1955). The first brood moths appear as early as December-
January in subtropical areas such as Hainan Island, China and Taiwan, where six generations occur
annually (Kiritani and Iwao 1967). In the cooler areas of China, the fourth brood is usually partial,
whereas in warmer areas such as Chunchi, the fourth brood is the largest (Tsai 1936). Three broods
are reported to occur during the 6 month growing season in Sri Lanka (Fernando 1967).
Various biotic factors operate in the suppression of yellow stem borer populations at their
different stages of growth. Eggs often have a high parasitization rate. In Sri Lanka, as many as 70%
of the larvae die mostly because of the activity of parasites and adverse ecological conditions
DALE 403
(Dayawathie 1983). Patel (1970) reported that during dry months the yellow stem borer occurs only
as a minor pest in Central India.
The seasonal abundance of S. incertulas is highly variable from year to year and region to region.
However, some definite trends are discernible. There are two well-defined peaks of pest abun-
dance, (i.e. April-May and September-October) in tropical countries such as India (Banerjee and
Mondal 1983), Bangladesh (Lippold 1971) and Pakistan (Ahmad and Afzal 1982).
In tropical rainy climatic zones as in West Bengal, India, high maximum temperatures (average
of 39.4°C), low humidity (average 53% in summer months), and low minimum temperature (average
12.7°C in winter months) are associated with low moth populations in these two seasons. In
temperate regions such as Kochi in Japan, a prominent population peak occurs in July instead of
the one in April-May in the tropics. In subtropical transitional areas, such as Taiwan, many seasonal
upsurges of the pest are noted.
High wind, especially above 8 km per hour, interrupts the flight activity of moths. The
nocturnal flight activity shows a clear lunar effect, with large catches at or near the new moon and
small catches during the full moon, possibly resulting from the effect of relative illumination. The
depression in the light-trap catches during the full moon phase showed a proportional inverse
relation with the duration and intensity of moonlight (Banerjee and Mondal l983)
The stage of host plant growth and soil fertility are factors that influence oviposition and larval
development in the yellow stem borer. Generally, rice plants in the vegetative and early heading
stages receive more eyes than those nearing maturity. Plants receiving more nitrogenous fertilizers
are preferred for oviposition to those treated with lower rates and provide a better nutritional
source for larval growth (Saroja and Raju 1981). However, the application of high amounts of
potash reduces the incidence of stem borer, probably by inducing resistance in host plants (John
and Thomas 1980, Vaithilingam 1975).
The yellow stem borer is a notorious pest of deep water rice in the main flooding period. In
Bangladesh, at harvest, more than 90% of white heads were found to be due to S. incertulas (Catling
1980). In Thailand, it comprised more than 95% of the borer population in attacked stems. The
average incidence increased with the stage of crop, from early elongation phase to ripening through
late elongation and booting-heading stages (Catling 1982). Stem damage and borer activity are
apparently very low during the preflood stage from May to mid-July. Stem damage increases
gradually during the late preflooding stage and culminates in maximum damage at harvest.
3.10 White Stem Borer, Scirpophaga innotata (Walker) (Lepidoptera: Pyralidae)

Synonym: Tryporyza innotata (Walker)
Distribution: Australia, India, Indonesia, Laos, Malaysia, Pakistan, Philippines, Taiwan, Vietnam
The white stem borer is a major pest of rice in South and Southeast Asia and Australia. It occurs
predominantly in areas where there is only one wet season crop per year and the stubble is left
undisturbed during the dry season. The pest is especially important in Indonesia, where it occurs in
Fig. 3.10
low-lying areas up to 200 m above sea level. It is a dominant species in the upland rice areas of
Sarawak. The insect does not occur in regions with high rainfall, as the larvae cannot survive
extremely wet situations (Soenardi 1967). Grain yield losses due to the white stem borer of up to
95% have sometimes been recorded in Indonesia. Heavy losses, however, do not occur every year.
The adult white stem borer is similar to the yellow stem borer in appearance. In contrast to the
yellow stem borer the female moth of the white stem borer does not have the characteristic black
spot on the wing and adults of the two white stem borer sexes are similar in appearance. The white
moth with an orange anal tuft is commonly seen in the field especially in the early stages of the crop.
The adult female has a wing span of 26-30 mm and the male of 18-24 mm.
Eggs are laid in clusters of 70-260 usually on the underside of young leaves. The egg mass is
covered with silky hairs from the anal portion of the female moth and is similar to that of the yellow
stem borer. The incubation period is 4-9 days.
The young larva penetrates the leaf sheath and bores down into the stem. It is milky white and
grows to a length of 25 mm. The age of host plant has a strong effect on the duration of larval period.
It varies from 19 to 31 days.
The full-grown larva pupates within the stem after making an exit hole for the moth to emerge.
Pupae are soft-bodied, pale and 12 to 15 mm long. They are more white-colored than those of the
yellow stem borer. Pupation is completed in 7-11 days.
Larvae undergo diapause during the dry season. After the crop has been harvested, the
caterpillars remain inactive in the lowest internodes of the stubble. Pupation does not take place
until the first rains at the beginning of the next season. With the rains, the larvae become active,
pupate, and the moths emerge. The threshold rainfall which terminates diapause is about 10 mm
(Soenardi 1967).
Adult moths are seen in rice fields only during the wet season. Three to five generations are
produced on the rice crop, depending upon the duration of the variety, time of sowing and
transplanting. The first 3 generations cause deadhearts in nurseries and in the young crop, while
feeding of the two subsequent generations causes whiteheads.
DALE 405
3.11 South American White Borer, Rupela albinella (Cram.) (Lepidoptera: Schoenobiidae)
Synonym: Scirpophaga albinella (Cram.)
Distribution: Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Hon-
duras, Mexico, Nicaragua, Panama, Peru, Surinam, Trinidad, Venezuela
Rupela albinella is not normally a major pest of irrigated rice even though sometimes high popula-
tions occur. It can be a serious problem to upland rice, attacking plants at ground level and feeding
upward in the stem. The damage is caused by the entry of larvae into the leaf sheath and tunnelling
into the nodes. Yellowish stained patches below the axis of leaf sheaths indicate the white borer
infestation.
Fig. 3.11
The shining white moths are commonly seen resting on the uppermost leaf tips or flying near
rice plants. The anal tuft is white in the male, while in the female moth it is yellow. Male moths live
for 4-6 days and females 5-8 days.
Conspicuous yellow clusters of eggs are laid on the leaves. Eggs are oval and yellowish green and
are covered with white silken hairs. They are laid in batches of 80 to 120. They hatch in 8-9 days.
Larvae are very agile, cream colored, and are usually dispersed by wind. The head is small and
reddish brown. There is a longitudinal dorsal brown stripe on the body. The activity of the
caterpillars is mostly restricted to the lower portion of the rice plant. The larvae make characteristic
holes in the stem before they pupate. The larval period is 35 to 50 days.
Pupation takes place inside the stem in a white silken cocoon, The pupa is yellowish brown in
color. The pupal period is 10 to 13 days. The adult mothcomes out through the hole previously cut
by the larva. In Surinam, there are two main periods of moth emergence in a year. Diapause occurs
only when the plant has flowered and it is also related to temperature.
3.12 African Pink Borer, Sesamia calamistis Hmps. (Lepidoptera: Noctuidae)

Synonym: Sesamia vuteria
Distribution: Angola, Burkina Faso, Burundi, Cameroon, Congo, Gambia, Ghana, Ivory Coast,
Madagascar, Mauritius, Reunion, Ruanda, Senegal, Sudan, Tanzania
Fig. 3.12
Host plants other than rice: Chrysopogon aciculatus (Retz.) Trin., Coix lachryma-jobi L.,
Hemarthria compressa (L. f.) R. Br., Hyparrhenia rufa (Nees) Stapf, Paspalum
conjugatum Berg., Pennisetum pupureum Schumach., Pennisetum subangustum
Stapf & Hubb., Rottboellia exaltata L. f., Saccharum officinarum L., Setaria splen-
dida Stapf, Sorghum bicolor (L.) Moench, Sorghum halepense (L.) Pers., Tripsacum
laxum Nash, Triticum aestivum L., Vossia cuspidata (Roxb.) Griff., Zea mays L.
Sesamia calamistis is a polyphagous pest and its infestation in the rice crop is highest at the
flowering stage. The young caterpillars occupy the stalk at the base of the panicle, which later
becomes a whitehead. Older larvae feed lower down in the stalk. The yield loss is directly related
to plant damage. In general, irrigated rice suffers less than the rainfed lowland crop. Often upland
rice is also severely attacked by this pest (Ho et al. 1983).
The adult moth is light beige with brown stripes. The margin of the anterior wings is wide,
whitish and partly smoky. The posterior wings are white with a yellowish margin. The moths are
nocturnal in habit and are able to travel long distances. Mating usually takes place as early as the
first night after emergence and egg-laying begins the same night. Eggs are thrust within the leaf
sheath surrounding the upper internodes. They are spherical, flat at the poles, and with numerous
longitudinal striations. The egg stage lasts for 7-10 days.
The larvae are smooth, uniformly yellowish pink on their dorsum, with greyish lateral and
dorsal stripes. The head capsule and thoracic plate are dark, shiny brown. The caterpillars first
feed within the tissues of the leaf sheath, subsequently entering the culm through a horizontal cavity
and then moving downward, sometimes through several internodes. They stay within the sheaths or
stems. The frass which fills the vertical galleries is partly thrown out through openings in the sheath.
The larval period is 4 to 5 weeks.
Pupation takes place within the base of the stem or in the folds of thewithered leaf sheaths. The
pupal stage lasts for 10 to 14 days.
DALE 407
In wet climates the life cycle is practically uninterrupted. But drought and cool conditions
induce a reduction in the rate of development. Mature larvae become inactive from the start of the
dry season until the rains return or the temperature increases. Usually it takes 2 months for the
completion of one generation, but towards the end of the crop season, duration of the life cycle may
be extended (Breniere 1977).
3.13 Pink Stem Borer, Sesamia inferens Walker (Lepidoptera: Noctuidae)

Synonyms: Leucania inferens
Nonagria inferens
Distribution: Bangladesh, Bhutan, Brunei, Burma, China, Hong Kong, India, Indonesia, Japan,
Kampuchea, Korea., Laos, Malaysia, Nepal, Pakistan, Papua-New Guinea, Philip-
pines, Ryuku Islands, Singapore, Solomon Islands, Sri Lanka, Taiwan, Thailand,
Vietnam
Host plants other than rice: Beckmania crucaefonnis (L.) Host., Cymbopogon nardus (L.) Rendle,
Cyperus japonicus Houtt., Echinochloa crus-galli (L.) Beauv., Echinochloa frumen-
tacea Link, Eleusine coracana (L.) Gaertn., Hemarthria compressa (L. f.) R. Br.,
Hordeum vulgare L., Panicum maximum Jacq., Paspalum scrobiculatum L.,
Paspalum thunbergii Kunth ex Steud., Phragmites karka (Retz.) Trin ex Steud.,
Polypogon fugax Nees, Saccharum arundinaceum Retz., Saccharum officinarum L.,
Saccharum spontaneum L., Scirpus maritimus L., Sclerostachya fusca (Roxb.) A.
Camus, Setaria italica (L.) Beauv., Sorghum bicolor (L.) Moench., Triticum aestivum
L., Zea mays (L.)
The pink borer is probably the least destructive pest among the rice stem borers. This may be due
to its extreme polyphagy, and outbreaks in rice usually result from a population spill-over from
adjacent sugarcane fields or other alternate hosts.
Fig. 3.13
The adult moth is fawn, the forewings being tan with dark brown markings. From a centered
point in the forewing, grey-black lines radiate toward the wing tips ending in a thin terminal line of
dark spots. There are tufts of hair on the pronotum. The hind wings are white. The wing span is
30 - 35 mm in the female and 20 - 30 mm in the male. The body length is 14 - 17 mm. Male and
female moths can be easily distinguished by observing their antennae; they are pectinate in the male
but filiform in the female (Pate1 and Rajesh Verma 1980).
Bead-like eggs are laid in rows between the leaf sheath and stem, at 30 to 100 eggs per batch.
The female is known to lay over 400 eggs. They are creamy white to dark and are naked. Eggs hatch
in about a week.
The larva has an orange-red head capsule and its body is purplish pink dorsally and white
ventrally. The head is relatively large compared to the rest of the body. The body is distinctly
segmented with no stripes and it tapers towards the abdominal tip. The caterpillars do not tend to
congregate but disperse early. They often come out of the stem of one host plant and bore into
neighboring rice plants thus destroying several plants. The larva becomes full-grown in 4 - 5 weeks
undergoing 5 - 7 molts. At this stage, it is 20 - 26 mm long. As the larvae are highly polyphagous,
they can move to adjacent fields or border areas to complete their development even after the rice
has been harvested.
Pupation usually takes place inside the larval tunnel within the stem but occasionally occurs
outside the stem between the leaf sheath and the stem. The pupa is dark brown with a tinge of
purple on the cephalic region. It is 18 mm long and 4 mm wide. The pupal period is about a week.
In the north Indian hilly tracts, low infestation on rice occurs in early July, when larvae cause
deadhearts in seedlings. Severe infestations occur in September, with maximum whiteheads of up
to 50-60% in the field (Garg and Tandon 1983).
There are up to six generations in a year. The ecology of the pink borer has received little
research attention.
4. RICE GRASSHOPPERS
A number of species of grasshoppers are occasionally found in rice fields but rarely cause significant
damage other than along the field margins. They chew angular holes in leaves, causing an injury
similar to that caused by leaffolders or armyworms. These insects are polyphagous; and besides
rice, they feed on maize, sugarcane, millets and many grasses. A list of rice grasshoppers and their
general distribution is given in Table 4.
4.1 Large Rice Grasshoppers, Hieroglyphus banian (Fabricius) (Fig. 4.1), and Hieroglyphus
nigrorepletus (I. Biol.) (Orthoptera: Acrididae)
Synonym: Hieroglyphus furcifer (Serv.)
Distribution: Afganistan, Bhutan, Burma, China, India, Nepal, Pakistan, Sri Lanka, Thailand,
Vietnam
DALE 409
TABLE 4
Rice grasshoppers and their distribution
Species General distribution
Acrida turricata K. India
Acrida exaltata Walker India
Acrotylus humbertiana S. India
Aeolopus affinis B. India
Aeolopus tamulus F. South and Southeast Asia
Atractomorpha crenulata F. India, Malaysia
Chrotogonus sp. India
Colemania sphenarioides Bol. India
Gastrimargus spp. Southeast Asia
Heteropternis respondens W. India, Taiwan
Hieroglyphus banian (F.) South and Southeast Asia, China
Hieroglyphus daganensis K. Africa
Hieroglyphus furcifer Sw. India
Hieroglyphus nigrorepletus (I. Bol.) South and Southeast Asia, China
Hieroglyphus oryzivorus Carl India, Pakistan
Oxya chinensis T. India to north Australia and up through
Southeast Asia to Japan
Oxya intricata (Stal) Africa, South and Southeast Asia
Oxya japonica T. South and Southeast Asia, Japan
Pyrgomorpha conica 01. Cyprus, India
Host plants other than rice: Gossypium hirsutum L., Pennisetum typhoides (Burm. f.) Stapf and C.
E. Hubb., Saccharum officinarum L., Sorghum halepense (L.) Pers., Setaria italica
(L.) Beauv., Zea mays L.
These grasshoppers are specific pests, which occasionally reach economic status, in marshy and
humid habitats. Serious outbreaks have been reported from many parts of India (Nair 1978, Singh
and Dhamdhere 1982).
Fig. 4.1
The nymphs eat newly germinated rice seedlings and cause them to wither. Adult grasshoppers
feed on the leaves and shoots and sometimes cut the earheads. If the emerging inflorescence is
attacked, the resulting grains become chaffy (Chatterjee and Debgoswami 1981).
The adult grasshopper is of medium size, the female measuring 34 - 54 mm and the male, 28 -
40 mm in length. It is dull green or yellow-brown. The lower body surface is brownish black. The
adults mate after a short pre-copulation period of 1 - 3 days. The longevity of the grasshopper
ranges from 1 to 8 months. Adults of H. nigrorepletus are more commonly found in the brachyp-
terous form than macropterous form. Egg pods are laid in soil, each pod containing about 35 eggs.
The total number of eggs laid per female ranges from 100 to 150. In wet sandy soil, eggs are laid at
a depth of 3-5 cm, while in dry sandy soil, they are laid on the soil surface. Oviposition occurs from
October to December and the eggs remain in soil till rains begin the following June-July. Individual
eggs are yellowish and covered with a gummy substance that hardens into a waterproof coating.
Nymphs hatch out shortly after the rainy season begins. If the rains are poor, the eggs do not
hatch and at temperatures above 40°C, there is heavy egg mortality. Young hoppers are brownish
yellow and later change to dull green. Nymphs which hatch from egg pods buried in compact soil
at a depth of more than 5 cm are unable to come out (Pradhan 1969). Like the adults, nymphs hide
from birds during the day and feed on rice foliage at night. The usual number of instars is 5 to 6 for
males and 7 for females. There is only one generation a year.
4.2 Rice Grasshopper, Hieroglyphus daganensis Krauss (Orthoptera: Acrididae)

Distribution: Chad, Ghana, Nigeria, Sudan
Host plants other than rice: Andropogon gayanus Kunth., Arachis hypogaea L., Chrysopogon
aciculatus (Retz.) Trin., Diectomis fastigiata (Sw.) Kunth., Hyparrhenia chrysargyrea
(Stapf) Stapf, Pennisetum subangustum Stapf & Hubb., Rottboellia exaltata L. f.,
Setaria sphacelata (Schumach.) Stapf & C. E. Hubb., Sorghum bicolor (L.) Moench.
The rice grasshopper occurs mainly in swampy grassy areas. In Ghana, it has appeared in large
numbers since 1972 and damages many hectares of rice crops. It is estimated that yields are
reduced by about 10% in outbreak areas. The following description of the pest is based mainly on
the paper by Agyen-Sampong (1975).
Both nymphs and adults feed on leaves but when the population density is high, panicles and
the stems are also attacked. The grasshopper population is higher in rice fields than on the grasses
during September-October, but the reverse is true during the end of October to early December.
The adult grasshopper is pale greenish yellow or light orange with yellow patches. The head is
ochraceous with yellowish brown eyes, pale greenish mandibles and a black band under each of the
genae. The male is medium-sized and slender and 3.1 to 4.2 cm long. The female is larger and
measures from 4.1 to 6.1 cm in length. Adults are more commonly found in the brachypterous form
than in the macropterous form.
Mating and oviposition begin from October to early December. The eggs are laid as pods in
the soil and they hatch in the beginning of the rainy season in July of the following year.
DALE 411
The grasshoppers start appearing in rice fields in August and the population reaches its peak
during September-October which coincides with booting and panicle initiation stages of the rice
plant. At the end of October when the crop begins to mature and leaves start turning yellow, the
grasshoppers leave for other grassy host plants.
H. africanus Uvarov is another grasshopper pest found in Africa. It is more robust than H.
daganensis.
4.3 Small Rice Grasshopper, Oxya chinensis (Thunberg) (Orthoptera: Acrididae)

Synonyms: Oxya chinensis Walk
Oxya velox (F.)
Distribution: Africa, Burma, China, India, Indonesia, Japan, Korea, Madagascar, Malaysia,
Taiwan
Host plants other than rice: Cyperus rotundus L., Saccharum officinarum L., Zea mays L.
This grasshopper is mostly a minor pest of rice. But in Korea up to 20% of the crop may be lost
during dry weather in some years.
The nymphs and adults feed on rice leaves, and nurseries suffer the most when attacked. Adults
may feed at the bases of maturing earheads causing them to dry up (Ramakrishna Ayyar 1963).
Fig. 4.3
The grasshopper is green or pale brown with a dark stripe running laterally from each eye
through the thorax to the base of wings. The male grasshopper measures 20 mm long while the
female measures 30 mm.
Eggs are laid in masses of 10 to 30 among rice stems, leaf folds and grass clumps above water
level. In dry conditions eggs are deposited just below the soil surface. Eggs are enclosed in a gummy
froth that hardens to form an ootheca. The eggs hatch in 2 - 3 weeks. There are usually 6 larval
instars with many females passing through an extra molt. Nymphal development usually takes about
100 days.
In the tropics, pest activity is continuous throughout the year, especially where both early and
late varieties are grown. In northern areas, there is only one generation a year and the insect
overwinters in the egg stage. In India, the grasshopper is seen in the rice fields all through the year
but is more abundant from August to November.
5. LEAFHOPPERS AND PLANTHOPPERS

The rice leaf- and planthoppers have gained great economic significance in recent years as their
infestations very often assume epidemic proportions. The leafhoppers (Cicadellidae) attack all the
aerial parts of the rice plant. But planthoppers (Delphacidae) confine themselves to the basal
portions.
Oman (1949) classified the damage done to rice plants by these insects into four categories:
(1) removal of plant sap by direct feeding; (2) injury in the plant tissue and deformation of leaves
and stems through defecation and oviposition; (3) transmission of various diseases; and (4)
predisposition of plants to diseases such as tungro, rice dwarf, orange leaf, and grassy stunt that are
transmitted by leaf- and planthoppers. The biology and ecology and of a few important leafhoppers
and planthoppers are herein given.
5.1 White Rice Leafhopper, Cofana spectra Distant (Homoptera: Cicadellidae)

Synonyms: Cicadella spectra
Tettigella spectra
Tettigoniella spectra
Distribution: Africa, South and Southeast Asian countries such as India, Philippines, Sri Lanka,
and Taiwan
Host plants other than rice: Cyperus rotundus L., Echinochloa colona (L.) Link., Fimbristylis
miliacea (L.) Vahl., Saccharum officinarum L., Scirpus articulatus L., Sorghum
bicolor (L.) Moench., Zea mays L.
It is a minor pest of rice rarely occurring at populations causing yield loss and does not transmit
any virus disease. Feeding by a large number of nymphs and adults may, however, cause typical sap
loss. Leaf tips first dry up and later the leaf turns orange and curls. The pest causes stunting and
yellowing of plants, and severe infestations cause plant death (Sam and Chelliah 1984). C. spectra
occurs in all rice environments but is most common in rainfed wetland rice at the end of the rainy
season.
The body of C. spectra is yellowish, the forewings grey-white with prominent veins and the head
bears four black spots. It is the largest among the species of leafhoppers and planthoppers. The
female, including tegmina, measures about 9.5 mm while the male is shorter and around 7.5 mm
DALE 413
Fig. 5.1
long. The adult longevity recorded in India for males and females is 7 and 10 days respectively (Sam
and Chelliah 1984).
Adults rest on the lower surface of the leaves or on tillers at the base of the plant. They are
highly attracted to lights at night. During oviposition, the female makes a cut parallel to the long
axis of the leaf sheath with its saw-like ovipositor. Eggs are laid in rows of 10 to 15 across the slit at
the base of the plant above the paddy water. The number of eggs laid per female averages about 50.
They hatch in 5 to 12 days.
Adults appear in transplanted rice fields in July-August and multiply and reach a peak
population during September-October. Then the population gradually declines through the cold
months of November and December. Many overwinter as eggs, though a few adults can also be seen
at this time. The overwintering eggs hatch in December-January, develop in grass hosts and the
resulting adults appear in January-February. Another population peak is reached during March.
Then the population again declines gradually.
5.2 Green Leafhoppers, Nephotettix spp. (Homoptera: Cicadellidae)

Green leafhoppers are important insect pests of rice throughout Asia. They cause either direct
damage to the rice crop by sucking the sap and injecting toxic chemicals or indirectly by transmitting
virus diseases such as dwarf, transitory yellowing, tungro, yellow dwarf and yellow-orange leaf.
The genus Nephotettix is widely distributed all over Asia and Africa. The important Oriental
species are N. nigropictus, N. virescens, N. cincticeps, N. malayanus and N. parvus. Their distribution
and the virus diseases they transmit, are given in Table 5. The genus in Africa and Madagascar has
been found to be represented by two species – N. modulatus and N. afer (Ghauri 1968).
414 RICE INSECTS — BIOLOGY A N D ECOLOGY
TABLE5
Distribution of green leafhopper vectors of rice virus diseases
Leafhopper Distribution Virus disease Transmitted

N. nigropictus Australia, Bangladesh, Bhutan, Burma, Dwarf, transitory yellowing, tungro, yellow
China, Hong Kong, India, Indonesia, Japan, dwarf, orange leaf
Kampuchea, Korea, Laos, Malaysia, Nepal,
Pakistan, Papua-New Guinea, Philippines,
Sri Lanka, Taiwan, Thailand
N. virescens Bangladesh, Burma, Hong Kong, India, In- Tungro, leaf yellowing, transitory yellowing,
donesia, Laos, Malaysia, Pakistan, Philippines, yellow dwarf, orange leaf
Taiwan
N. cincticeps China, Japan, Korea, Taiwan Transitory yellowing, yellow dwarf,
dwarf
N. malayanus Burma, China, India, Malaysia, Philippines, Waika, tungro
Sri Lanka, Taiwan
N. parvus India, Malaysia, Sri Lanka Tungro, yellow dwarf
Feeding by green leafhoppers is confined mostly to the leaf and leafsheath of rice. Mild
infestations may reduce the vigor of the plants and the number of productive tillers. Heavy
infestations cause withering and complete drying of the crop.
Among the diseases transmitted by Nephotettix spp., tungro is the most destructive. The
infection occurs primarily in the early stages of crop growth. It is the only known non-persistent or
transitory rice virus (Ling and Tiongco 1979). When the epidemic is severe, 100% yield loss can
occur (Heinrichs 1979). Heavy epidemics of tungro occurred in Indonesia, Thailand, Malaysia,
Bangladesh and India during the last two decades. In the Philippines, the disease has been a serious
threat ever since it first appeared as an epidemic in 1957.
Nephotettix spp. were known only as minor pests in Bangladesh until 1955 (Alam and Islam
1959). But since then, 50 to 80% damage has been reported from different regions of the country.
The emergence of green leafhoppers into a major pest has been commonly attributed to the
introduction of high yielding rice cultivars and the accompanying high nitrogen applications (Karim
and Pathak 1979).
The optimum temperature for the development of green leafhoppers ranges between 25 and
30°C. Insects exposed to higher temperatures lay less eggs, some of which may not hatch. The high
incidence of Nephotettix spp. is attributed to favorable temperature, low rainfall and good sunshine.
A positive correlation has been obtained in Japan between the hopper population and the amount
of sunlight while a negative correlation exists between the population build-up and relative
humidity.
Acoustic behavior of green leafhoppers during courting has been well established as a function
of communication between the two sexes. Ichikawa (1976) first reported the vibration signals
emitted from the abdomen of male and female N. cincticeps prior to mating. He could also show
that male sounds are the essential stimuli for the female to receive an appropriate mating partner.
Later Inoue (1982) demonstrated that these calling sounds in Nephotettix spp. are species-specific
DALE 415
and may act as one of the mechanisms of reproductive isolation among the different species of green
leafhoppers.
There are five closely related species of the genus Nephotettix in Asia. The distribution of some
of these species is symmetric in the tropics and subtropics. Further, some are not isolated seasonal-
ly and have a common host plant: N. virescens and N. nigropictus on rice in the tropics, N. virescens,
N. nigropictus and N. malayanus on the weed Leersia hexandra in the tropics and the subtropics.
The genus Nephotettix is represented in Africa and Madagascar by two species N. modulatus
and N. afer (Ghauri 1968). Both are largely symmetric and widely distributed throughout Africa.
Information regarding their bionomics and their possible role as vectors of rice virus diseases is
Scanty.
5.2.1 Nephotettix nigropictus (Stål)

Synonym: Nephotettix apicalis Ishihara et Kawase
Host plants other than rice: Echinochloa colona (L.) Link, Echinochloa crus-galli (L.) Beauv.,
Eleusine indica (L.) Gaertn., Ischaemum indicum (Houtt.) Merr., Leersia hexandra
Sw., Paspalum scrobiculatum L., Saccharum officinarum L., wild rices, Zea mays L.
The adult is green, the male possessing a median black wedge-shaped mark on the forewings. In
the female, this area is brown in color. Both nymphs and adults are very active in summer, especially
on hot sunny days. With the onset of cold weather, they become somewhat sluggish.
The premating period ranges between 2 to 5 days after the final molt. Females start laying eggs
1 to 3 days after mating and prefer to oviposit in the basal portions of leaf sheaths of young rice
seedlings. The eggs are laid in batches of 10 to 15 into the epidermis and cortex and sometimes in
the air cavities of the leaf sheath with the saw-like ovipositor. They are arranged in a single row,
along the length of the leaf sheath. Most eggs are laid during the afternoon hours (Dhawan and
Fig. 5.21
Sajjan 1976). The ovipositional sites can be seen as slightly bulged portions and are lighter green in
color. The number of eggs laid starts declining with age after the second week of mating. In India,
the average adult longevities for males and females are 56 to 62.3 days respectively during the
months of June to October. But the period extends during the winter months (Misra 1980).
Eggs are somewhat crescent-shaped, pale yellow and measure 1 × 0.3 mm. Reddish eye spots
are visible just before hatching. The incubation period varies from 8 to 14 days depending on the
temperature.
Nymphs are creamy white at first and later turn green. The color darkens as the nymphs pass
through five successive molts. The nymphal period is 2 to 3 weeks in summer and longer in winter.
Nasu (1967) observed a longer nymphal period in the case of females than in males. Nymphal
survival was found higher on the weed Leersia sp. than on rice cultivars (Razzaque 1984).
Seasonal occurrence is based on studies conducted in India (Misra 1980). The leafhoppers
start appearing in the rice fields in August. They multiply and reach a peak population during
October-November. Then the population declines drastically due mainly to the fall in temperature.
Surviving adults live on grasses found near irrigation canals and levees. The ratoon and self-sown
rice also harbor the pest. During December-January, the hoppers migrate to rice nurseries and
then in February-March to the transplanted crop. A rise in population takes place during this time
even though the number is comparatively smaller than that in October-November. Again the
hoppers move on to grasses during summer for feeding and egg-laying. The adults emerging from
these grasses migrate to the rice fields during late July or early August.
5.2.2 Nephotettix virescens (Distant)

Synonyms: Nephotettix bipunctatus Distant
Nephotettix impicticeps Ishihara et Kawase
Host plants other than rice: Cynodon dactylon (L.) Pers., Echinochloa crus-galli (L.) Beauv.,
Eleusine indica (L.) Gaertn., Leersia hexandra Sw., Panicum ramosum L., Sac-
charum officinarum L., wild rices, Zea mays L.
The newly-emerged adult is yellowish in color. It gradually turns yellowish green and then green in
about 3 hours after emergence. Most of the adults emerge early in the morning. The adult green
leafhopper is about 4 mm long. It has a pointed vertex and a green head. Males have green
forewings with a small dark brown or black band in the middle while in females there is no such
band.
The premating period ranges from 1 to 2 days. Eggs are laid in small slits made in the soft parts
of the leaf sheath. The number of eggs in a batch seldom exceeds 30 and the total eggs laid during
the life cycle is around 350. Unmated females lay sterile eggs randomly on the leaf sheaths, while
mated females lay fertile eggs which are inserted in the leaf sheath in an arranged manner.
Newly-laid eggs are barely visible and are oblong, bent and pale yellow. The incubation period
varies from 6 to 12 days.
DALE 417
Fig. 5.2.2
Young nymphs are creamy white with black longitudinal stripes on the sides of the body. They
turn yellow or yellow-green in about an hour after molting. First instar nymphs are more numerous
on the lower surface of older leaf blades, but from the second instar onwards, they distribute
themselves rather evenly on all leaves. Nymphs and adults suck sap from leaf sheaths and blades.
The rate of nymphal development is faster at higher temperatures. Cheng and Pathak (1971)
reported that the average larval duration was only 14.1 days at 35°C while it was 37.3 days at 20°C.
The male nymphal period is usually shorter than that of females. Rice plants at the tillering and
panicle initiation stages are most favorable for the rapid build-up of pest populations. The nymphal
stage is greatly prolonged during winter. The insect can withstand long periods of starvation (Basu
et a1 ., 1976).
Ho and Chen (1968) observed 8 generations of the pest in Taiwan whereas in India, occurrence
of up to 11 generations has been recorded. Throughout India, the population of N. virescens is
higher than that of N. nigropictus (Mathur and Chaturvedi 1981).
Leafhoppers move from one rice crop to another and during the intervening periods they feed
and breed on grasses found in the rice ecosystem. There are two population peaks in a year — one
in March during the first crop season (February to May/June) and another in October-November
during the second crop (June to December).
It Vietnam, it was reported that during a year when there is little rain with low temperature and
high humidity, there will be a heavy incidence of green leafhopper (Quyen 1963). Similar conditions
lead to sudden N. virescens outbreaks in India (Ghosh et al., 1960).
5.2.3 Nephotettix cincticeps (Uhler)
Host plants other than rice: Alopecurus aequalis Sobol., Avena fatua L., Echinochloa crus-galli
(L.) Beauv., Leersia japonica Honda, Leersia oryzoides (L.) Sw. var japonica,
418 RICE INSECTS—BIOLOGY AND ECOLOGY
Fig. 5.2.3
Phalaris arundirtacea L., Phleum paniculatum Huds., Phragmites australis (Cav.)

Trin. ex Steud., Poa annua L.
Nephotettix cincticeps is the only species related to the incidence of the yellow dwarf disease in
Japan. Yield loses as high as 75% have been reported due to this disease.
Life history and habits are similar to those given for N. nigropictus. But there are differences in
the seasonal occurrence caused primarily due to the ecological differences in the pest habitats. The
following description is based on the study of Hirao and Inoue (1978) conducted in Japan.
N. cincticeps passes the winter as a half-grown, mostly fourth instar, diapausing nymph. During
the off-season of rice cultivation, a winter grass, water-foxtail Alopecurus aequalis serves as the main
host plant. This grass is widely seen in fallow fields after rice harvest. Adults migrate from the wild
host vegetation to paddy fields in early summer immediately after transplanting. Afterwards, the
hoppers spend three generations in paddy fields. The adult emergence of the last generation is
interrupted by the nymphal diapause induced by short day length of autumn. There are four
generations of the pest in most regions of Japan. But there may be a fifth or even a partial sixth
generation in the warmer southern parts of Kyushu.
A positive relationship exists in Japan between low populations of N. cincticeps and heavy
snowfall although this factor is not the only one in checking pest outbreaks (Otake 1966).
5.2.4 Nephotettix malayanus Ishihara et Kawase

This is one of the two Malayan species of the genus Nephotettix reported by Ishihara and Kawase in
1968. It formed 3.9% of the hopper specimens collected from North Krian District, Perak of the
Malay Peninsula during 1960-62. N. malayanus is less of a threat to rice cultivation than the other
green leafhoppers because a weed, Leerisia hexandra is the most preferred host plant.
DALE 419
Fig.5.2.4
N. malayanus adults are mostly greenish although the head is tinted with yellow. It is smaller in
size than the closely allied species N. cincticeps. Body lengths, including tegmina, of the male and
female are 3.8-4.6 mm and 4.3-5.0 mm respectively. Adults usually emerge in the morning.
Eggs, which are deposited in leaf sheaths, hatch in 7 to 10 days. The newly hatched nymphs
remain on the leaf sheath for about an hour and then disperse to the leaf blades. There are five
nymphal instars. The nymphal period varies from 13 to 19 days.
The effect of temperature on the development of N. malayanus on L. hexandra was studied by Kim
(1985). He found that the developmental period was longest at the lowest temperature. At 25°C, the
incubation period of the eggs was 10.2 days while at 33°C, it decreased to 7.1 days. Similarly, the life span
of the adult hoppers was longest (23.3 days) at 25°C while it was only 17.3 days at 33°C.
N. malayanus is the vector of two virus diseases of rice - waika and tungro. Both adults and
nymphs of N. malayanus transmit rice waika virus and transmission efficiency is higher than in N.
cincticeps. But in the case of tungro, transmission efficiency is generally lower with N. malayanus
than with N. virescens (Kim 1985).
5.2.5 Nephotettix parvus Ishihara et Kawase

This species was also established by lshihara and Kawase (1968) from their insect collections from
the Malay Peninsula. It formed 1.7% of specimens belonging to the genus Nephotettix.
N. parvus is the smallest among the green leafhoppers. Females closely resemble those of
N. nigropictus but the crown is more prolonged anteriorly and the body is comparatively shorter
and narrower.
5.2.6 Nephotettix modulatus Melichar

Distribution: Angola, Congo, Egypt, Ethiopia, Israel, Ghana, Ivory Coast, Kenya, Madagascar,
Malawi, Morocco, Nigeria, Senegal, Sudan, Tanzania, Tunisia, Uganda, Zambia
The usual color pattern is almost like that of N. nigropictus with which N. modulatus is very often
confused. It is lighter in color than N. afer. Specimens of N. modulatus are slightly larger than those
of N. afer and differ in that the anterior margin of vertex is conically produced, ovipositor is longer,
and dorsal setae of male pygofers are longer.
5.2.7 Nephotettix afer Ghauri

Distribution: Angola, Congo, Ethiopia, Kenya, Madagascar, Nigeria, Rhodesia, South Africa,
Sudan, Tanzania, Uganda, Zambia
The color pattern of N. afer is similar to that of N. nigropictus, even more so than N. modulatus.
Three transverse black bands, marginal and submarginal on the vertex, are well developed in most
specimens.
5.3 Zigzag Leafhopper, Recilia dorsalis (Motschulsky) (Hornoptera: Cicadellidae)

Synonyms: Inazuma dorsalis
Dellocephalus dorsalis
Distribution: Australia, Bangladesh, Bhutan, Burma, China, India, Indonesia, Japan, Kam-
puchea, Korea, Laos, Malaysia, Pakistan, Philippines, Sri Lanka, Taiwan, Thailand,
Vietnam
Host plants other than rice: Cynodon dactylon (L.) Pers., Cyperus rotundus L., Echinachloa
colona (L.) Link., Hordeum vulgare L., Leersia hexandra Sw., Saccharum of-
ficinarum L., Triticum aestivum L.
R. dorsalis is one of the vectors of rice dwarf, orange-leaf and tungro viruses in many parts of Asia.
Only the female is viruliferous. Both nymphs and adults suck sap from the leaves and leaf sheaths.
Leaf tips dry and leaf margins become orange. Later the entire leaf becomes orange and leaf
Fig. 5.3
DALE 421
margins curl. Damage appears first on older leaves. Young seedlings wilt and die when R. dorsalis
is abundant. However, it usually occurs in low population densities and seldom causes significant
damage to the rice crop by removal of plant sap. It occurs in all rice environments but transmits
virus diseases only to wetland rice. Adults are highly mobile and enter rice fields in the early growth
stages.
The forewings of the adult insect are white with light brown bands taking the shape of a ‘W’ and
giving the wing a zig-zagged pattern. Body length is 3.5-4.0 mm.
The premating period, after the final molt, ranges from 2 to 4 days. Egg are laid in rows within
the leaf sheaths. The average number of eggs laid per female is 98 (Misra 1980) and eggs hatch in
7 - 9 days. The ovipositional sites can be located with difficulty as small brownish spots; sometimes
micropylar ends of eggs protrude from the leaf surface. The adult hoppers live for 10-14 days.
There are five nymphal stages lasting for 16 days. The nymphs are yellowish brown in color
varying from 1.0 to 3.0 mm in body length. They are found both on leaves in the upper parts of the
rice plant and on tillers near the base.
In India, the adults make their first appearance during the rainy season, June-July, in the rice
nurseries and newly-planted fields. Their number gradually increases to reach a peak during
August-September. The population then gradually declines in October-November. At that time
some adults are still found on plants on the leaves of paddy fields. These overwintering adults again
appear in December-January in the seedbed of the first crop. In the newly-transplanted crop, they
multiply rapidly and reach a peak during February-March. Then the population starts declining.
R. dorsalis in southern Japan has four generations in a year and hibernates in the egg stage.
5.4 Smaller Brown Planthopper, Laodelphax striatellus (Fallen) (Hornoptera: Delphacidae)

Synonyms: Delphacodes striatellus
Laodelphax striatella
Distribution: China, Europe, Japan, Korea, Philippines, Siberia, Taiwan
Host plants other than rice: Wheat, barley, maize, oats, sugarcane, millets and various grasses
Laodelphar striatellus is important as a vector of rice-stripe and black-streaked dwarf virus in China
and Japan (Anon. 1976). Once it has acquired the virus it remains a vector throughout its life.
Although the nymphs and adults suck plant sap, feeding damage rarely causes yield losses. It is
abundant in temperate subtropical regions.
Adults have two wing forms, brachypterous and macropterous. The adult males are usually 3.5
mm long, while the females are smaller, about 2.0 mm long. Sometimes the insects have red eyes.
The female lays 60-260 white eggs in masses in the leaf midrib or leaf sheath near the base of
the plant. The eggs hatch in 5-15 days.
Fig. 5.4
The nymphs are light to dark brown and are usually smaller than those of other species at
corresponding instars. There are five larval instars. At a temperature of 25°C, the nymphal period
is about two weeks.
The hoppers congregate on the lower portions of the plant just above the water level in irrigated
fields, and in the interior of the clustered bases of the tillers, strongly suggesting that the insects
prefer humid conditions. Close planting and weedy rice fields favor the incidence of the pest.
There are six to seven generations in a year. In Japan, this species hibernates as last instar
nymphs in winter wheat. The emerging adults then move into the transplanted rice in late May and
early June. Usually 5% of these hoppers carry stripe virus. By July-August, they infest the second
rice crop.
The smaller brown planthopper harbors yeast-like symbiotes in the fat body, which are trans-
mitted to the next generation through eggs (Noda 1977). High temperature (35°C) destroys the
symbiotes and leads to a poor growth of the host insects (Noda and Saito 1979).
5.5 Brown Planthopper, Nilaparvata lugens (Stål) (Homoptera: Delphacidae)

Synonyms: Nilaparvata oryzae Esaki et Hashimota
Hikona formosana Matsumura
Nilaparvata sordescens Kuwayama
Distribution: Australia, Bangladesh, Bhutan, Burma, China, Fiji, India, Indonesia, Japan, Kam-
puchea, Korea, Laos, Malaysia, Nepal, Pakistan, Papua New Guinea, Philippines,
Solomon Islands, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Leersia hexandra Sw.
The brown planthopper became a threat to rice production throughout South and Southeast Asia
in the early 1970s. Outbreaks have coincided with the large-scale release and cultivation of high
yielding varieties. The brown planthopper damages the plant through the removal of plant sap and
as a vector of rice viruses. The honey dew excreted by the nymphs and adults at the base of the
DALE 423
Fig. 5.5
plants is covered with sooty mold. Even the introduction of rice cultivars with genetic resistance to
the brown planthopper has not provided a panacea because of the selection for biotypes that arc
capable of surviving on these cultivars (Pathak and Khush 1979).
The insect prefers rainfed and irrigated wetland fields to upland rice, and direct-sown fields to
transplanted fields. It is known only to feed on rice and the weed Leersia Hexandra (Heinrichs and
Medrano 1984). It infests the rice crop at all stages of plant growth. As a result of feeding by both
nymphs and adults at the base of the tillers, plants turn yellow and dry up rapidly. At early
infestation, round yellow patches appear which soon turn brownish due to the drying up of the
plants. This condition is called ‘hopperburn.’ The patches of infestation then may spread out and
cover the entire field. Crop loss is usually considerable and complete destruction of the crop occurs
in severe cases. N. lugens is a vector of the virus diseases - grassy stunt, ragged stunt, and wilted
stunt (Hibino 1979, Chen and Chiu 1981).
Adult emergence takes place at the basal part of the host plant. It begins at dawn and continues
for 4-5 hours. The adult planthoppers are ochraceous brown dorsally and brown ventrally. The
tegmina are subhyaline with a dull yellowish tint. The female measures about 5 mm and the male
4.5 mm in length. Adult longevity differs considerably between laboratory and field conditions
(migrants), maximum values being 36.6 and 9.0 days respectively.
The brown planthopper is dimorphic, with fully-winged ‘macropterous’ and truncate-winged
‘brachypterous’ forms. The macropters are potentially migrants and are responsible for colonizing
new fields. At the time of colonization, the macropterous forms dominate in a rice field both under
temperate (Kisimoto 1956,1965; Kusakabe 1979) and tropical conditions (Dyck et al., 1979). After
settling down on rice plants, they produce the next generation in which most of the female insects
develop as brachypters and males as macropters. There are many factors which have been sug-
gested to be responsible for wing morphism in the brown planthopper. Crowding during the larval
stage and reduction in the quality and quantity of insect food induce macroptery (Kisimoto 1965).
Short daylength and low temperature also favor the production of macropterous forms (Johno 1963,
Kisimoto 1965). Saxena et al., (1981) have shown a signifcant increase in macropterous forms
among progenies reared on senescent and hopperburned rice plants.
Sexual communication between the adults of the brown planthopper operates primarily by
means of acoustic signals transmitted through the substrate (Ichikawa and Ishii 1974, Ichikawa et
al., 1975). These signals resemble most of the smaller Auchenorrhyncha (Claridge 1983).
Courtship behavior may be initiated by either sex. Both virgin females and mature males start
emitting signals within a few minutes of settling on a host plant. The female call consists of simple
intermittent pulses produced by a visible vibration of the abdomen. On receiving the signal, the
male walks rapidly towards the female. If the female stops signalling, the male stops walking and
starts sending its own signals until they are reciprocated by the female. The male call has a more
complicated structure (Claridge et al., 1985). In consists of repeated sections which themselves
consist typically of three phases: (a) a series of 3 to 10 complex pulses, (b) a series of regularly and
rapidly repeated pulses, and (c) further complex groups of pulses. Thus an alternation of male and
female calls eventually leads the male to make contact with the receptive female and mating may
take place. Claridge et al., (1984) studied populations of N. lugens from different geographically
defined areas and found some to differ significantly in the pulse repetition frequencies of male calls.
The biggest differences were found between populations from Australia and those from the
Solomon Islands.
Adults usually mate on the day of emergence and the female starts laying eggs from the day
following mating. Brachypterous females lay 300 to 350 eggs, while macropters lay less eggs. The
eggs are thrust in a straight line generally along the mid-region of the leaf sheath, though sometimes
eggs are laid on the leaf midribs. Eggs are covered with a dome-shaped egg plug secreted by the
female. Only the tips of eggs protrude from the plant surface. The egg-laying sites appear as
brownish streaks. The number of eggs laid at a site has been reported to vary in different
countries – 2 to 3 in Japan, 4 to 10 in the Philippines, and 2 to 12 in India. Red eye spots appear at
one end of the egg before hatching. Eggs hatch in about 6 to 9 days.
The newly hatched nymph is cottony white and turns purple brown within an hour. Feeding on
the plant sap, it undergoes five instars to become an adult. The nymphal period varies widely
depending on the food conditions, density during development, and other environmental factors.
In the tropics, it takes about 10 to 18 days from hatching until the first instar nymph reaches
adulthood.
Temperature is a critical factor in the life activities of the insect. The threshold temperatures
of embryonic and post-embryonic development of the brown planthopper are 10.8 and 9.8°C
respectively (Suenaga 1963). The hatchability and survival rate are the highest around 25°C. Eggs
are very sensitive to desiccation and soon shrivel when the host plant starts wilting (Kisimoto 1977).
The temperature conditions in the nymphal stage affect the longevity and oviposition of adult
hoppers (Mochida 1964). The population growth of N. lugens is maximum at a temperature range
of 28 to 30°C in the daytime and slightly lower at night. In warm and humid climates the planthop-
pers remain active throughout the year and their population fluctuates according to the availability
of host plants, activity of natural enemies and other environmental factors prevailing in the locality.
DALE 425
There are many reports (Miyake and Fujiwara 1962, Okamura 1963, Sugimoto 1967) on the
incidence of diapause in the egg stage of N. lugens. Low temperature coupled with short
photoperiod and rice plants at the ripening stage were factors reported to induce females to deposit
diapausing eggs. However, it is questionable to consider this as actual diapause because none of
these reports defined diapausing eggs either morphologically or physiologically.
N. lugens is a long-range migrant. The brown planthoppers make wind-assisted migratory
flights each year to colonize the summer rice growing areas of China. By capture of marked
hoppers the insects are found to migrate 200 to 300 km in mainland China (Tu 1979). The hoppers
are carried to Korea and Japan by prevailing winds from permanent breeding grounds in southern
China. Influxes of the brown planthopper occur from late June to mid-July every year coinciding
with the arrival of low pressure fronts from the south. In the tropics, migrations also occur. Saxena
and Justo (1984) collected hopper adults in two successive inter-island voyages indicating that
migrant planthoppers are reaching the Philippine Archipelago from certain rice-growing areas
lying to the southwest of the Philippines in the Indian Ocean. The inflowing warm and humid air
currents facilitate this migration. Observations indicate that the brown planthoppers take off at
dusk and that some are capable of continuously flying up to 26h if the temperature is more than 17°C
(Rosenberg and Magor 1983). Others may fly for shorter durations giving the transient population
a chance of dispersal and colonization on rice plants on the way.
Several biotypes of N. lugens are known to occur throughout tropical Asia. They are distin-
guishable by their differential virulence, honeydew production and ability to survive on various rice
varieties. The susceptibility of IR26, the first brown planthopper-resistant variety released by IRRI,
in India signalled the evolution of biotypes among hopper populations.
The seasonal prevalence of the pest is mostly dependent on the availability of host plants. In
the warm and humid tropics, planthoppers remain active throughout the year. In India, in most
regions, the peak population is observed during the late rainy season from October to November.
Another peak appears during the dry season from April to May in the regions where double
cropping is widely practiced. In Japan and Korea, macropterous adults immigrate into rice fields
from late June to early July every year. Afterwards, the pest spends several generations on rice and
moves or dies at the end of cropping season.
N. lugens can have several generations during one cropping season on irrigated rice in the Asian
tropics depending on the length of the crop. It has 5 generations on one rice crop in southern Japan
(Mochida 1964), 5-6 in the central China (Lei and Wang 1958) and 4 to 6 in Indonesia (Mochida et
al., 1977). There are usually three generations on the modern high yielding intermediate duration
varieties in the tropics (Heinrichs et al., 1986).
N. lugens is a phloem-feeding insect. Removal of translocating nutrients from the plant sap
reduces the net photosynthate available to the plant for sustenance and storage. The biological
effects which collectively lead to plant damage have been detailed by Sogawa and Cheng (1977) and
Chen and Cheng (1979). The effects and their interactions are schematically presented in Fig. 1(see
Page 364).
Many factors have been attributed to the rapid development of brown planthopper popula-
tions. Laboratory studies of Sogawa (1971) have shown that the hoppers prefer and feed more on
plants receiving high nitrogen rates. This leads to faster insect development and higher fecundity.
These findings were confirmed in the work of Heinrichs and Medrano (1985) where results clearly
showed that insect weight, feeding rate and population growth increased with the application of N
fertilizer. Observations from the Philippines (IRRI 1979), Indonesia (Mochida et al. 1979) and
Taiwan (Cheng 1977) all concur that intensification of rice production through continuous irriga-
tion favors outbreaks. Insecticides have been suggested as the major factor involved in brown
planthopper outbreaks due to the poor control and reproductive stimulation of the hopper at
sublethal rates and destruction of natural enemies (Heinrichs and Mochida 1984).
5.6 Whitebacked Planthopper, Sogatella furcifera (Horvath) (Homoptera: Delphacide)

Synonyms: Delphax furcifera
Delphacodes furcifera
Sogata furcifera
Sogata distincta
Sogata pallescens
Megamelus furcifera
Chloriona furcifera
Related species infesting rice:
Sogatella kolophon
Sogatella nigeriensis
Sogatella vibix
Fig. 5.6
DALE 427
Distribution: Afghanistan, Australia, Bangladesh, Bhutan, Burma, China, India, Indonesia,

Japan, Kampuchea, Korea, Laos, Malaysia, Mongolia, Nepal, Pakistan, Papua New
Guinea, Philippines, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Millets, maize and many grasses
Sogatella furcifera has a long history as an insect pest of rice. It was first reported from Japan in 1899
and from India in 1903 (Chaudhary et al., 1968). The whitebacked planthopper has emerged as a
major pest of rice in many Asian countries where it has destroyed sizeable proportions of the crop
in certain years. The whitebacked planthopper occurs in both upland and wetland rice environ-
ments.
Long-winged adults enter the field at about 30 days after sowing and usually complete fewer
generations on a crop than does the brown planthopper. The whitebacked planthopper prefers a
young crop and often produces long-winged migratory forms before the plants flower. Both adults
and nymphs suck sap primarily at the base of the rice plants which leads to yellowing of the lower
leaves, reduced vigor and stunting of plants. Because seedlings are attacked in the nursery,
infestation is often carried through eggs into the transplanted crop. Severely attacked seedlings do
not grow; are stunted, wilt and eventually die. If the infestation is at the panicle initiation stage, the
number of grains and the panicle length decrease. But when attacked later, during the maturation
period, grains do not fill fully and ripening is delayed. When the hoppers are present in large
numbers late in the crop growth stage they are seen infesting the flag leaves and panicles. Gravid
females cause additional damage by making oviposition punctures in leaf sheaths. Feeding points
and wounds caused by egg-laying may later become potential sites for the invasion of bacteria and
fungi. Moreover, the honeydew produced by the hoppers serves as a medium for mold growth. The
pest has not been proven to be a vector of any rice virus disease.
The rice plants affected by the whitebacked planthopper appear uniformly in large areas
throughout the field rather than as localized hopperburned patches as in the damage by the brown
planthopper. This may be due to the difference in the distribution patterns of the two planthopper
species.
Under favorable conditions, the insect produces several generations and inflicts heavy damage
on the rice crop. Serious outbreaks of the pest in the tropics have been reported in Bangladesh,
India, Indonesia, Malaysia, Nepal, Pakistan, and Vietnam (Mochida et al., 1982). In Japan, the
grain loss caused by this pest has reached up to 90% in some fields (Suenaga 1971). A study on the
percentage loss of paddy caused by the artificial infestation of the whitebacked planthopper was
conducted by Khatri et al. (1983). The percentages of grain loss varied from 11 to 39 when 15 insects
per hill were released at varying stages of plant growth.
The adult hopper is 3.5 - 4.0 mmn long. Forewings are almost uniformly subhyaline with dark
veins. There is a prominent white band between the junctures of the wings. The body is creamy
white with the mesonotum and abdomen black dorsally and the legs, ochraceous brown. There is a
conspicuous black dot at the middle of the posterior margin of each forewing which meets when the
forewings come together. Macropterous males and females and brachypterous females are com-
428 RICE INSECTS - BIOLOGY AND ECOLOGY
monly found in the rice crop whereas brachypterous males are very rare. The planthoppers,
especially adults, prefer to stay at the upper portion of rice stems. Adults are positively phototropic
and hence are attracted to light traps.
The preoviposition period ranges between 3 to 8 days. Eggs are laid in longitudinal rows within
the leaf midribs. On an average, a female hopper laid 164 eggs in tests in India (Vaidya and Kalode
1981). But in Japan, Suenaga (1963) noted that the total number of eggs laid per female ranged
between 300 and 350. Eggs are similar in size and shape to those of the brown planthopper but have
a longer, more pointed, egg plug. The eggs hatch in about 6 days. Nymphs are white to a strongly
mottled dark grey or black and white in color. They reach adulthood in 12 to 17 days passing
through five instars.
In Japan (Miyake and Fujiwara 1961) the macropterous males and females immigrate into rice
fields during late June and early July. At this time, the hoppers show distinct ovipositional
preference for rice plants. During summer, the number of brachypterous adults increases. The
pest population declines in the rice fields in late August. Then there is a shift in ovipositional
preference from rice to grass. During October-November, the adults emerging on grasses start
laying diapaused eggs which hatch by the middle of April. Nymphs prefer weeds for feeding. The
adult hoppers which develop from these nymphs infest the rice fields in June-July and the cycle
repeats.
The ecological conditions and factors reported to be conducive for a population explosion of
the whitebacked planthopper differ in various regions. In Vietnam, daylength, temperature and
humidity play important roles in planthopper epidemics. According to Tao and Ngoan (1970),
macropterous and brachypterous adults are usually abundant during the wet season when day and
night temperatures do not vary much and the humidity remains around 85%. A prolonged monsoon
with intermittent rain and application of heavy doses of nitrogenous fertilizers were suggested as the
cause for an epidemic in Pakistan in 1978 (Majid et al., 1979).
S. furcifera is known to migrate across mainland China (Wang et al., 1982) and from China to
Japan and Korea (Mochida et al., 1982). Saxena and Justo (1984) collected airborne hoppers in
traps set aboard inter-island vessels sailing in the Philippine archipelago.
The importance of the whitebacked planthopper as a rice pest has tremendously increased
during the last decade both in upland and lowland areas, particularly where varieties resistant to
brown planthoppers are grown. It may be that the ecological niche vacated by N. lugens is gradually
being occupied by S. furcifera (Khan and Saxena 1985).
5.7 Rice Delphacid, Tagosodes orizicolus (Muir) (Homoptera: Delphacidae)

Distribution: Argentina, Brazil, Caribbean Islands, Central America, Colombia, Mexico,
southern United States, Surinam
Host plants other than rice: Grass weeds such as Echinochloa crus-galli (L.) Beauv.
DALE 429
Fig. 5.7
The rice delphacid causes severe damage to the rice crop as the principal vector of hoja blanca or
white leaf disease. It is also often present in sufficient populations to destroy entire rice fields
through feeding damage. Adults and nymphs suck sap from rice leaves, stems, and also from
developing panicles during the flowering stage. The insects excrete honeydew which forms the
substrate for sooty mold on leaves and stems. The pest feeds on both rice and weeds even though
it prefers rice. These insects are rather sedentary in habit but are distributed by heavy winds and
flowing water.
Adults are 3 to 4 mm long, brownish red with a light median stripe dorsally. The body darkens
towards the abdomen. Males are smaller and darker than the females. Nymphs have two black
stripes running the entire length of the body. Adults are of two forms — macropterous and
brachypterous. The wings are yellowish, the veins yellow except for some in the apical area and the
white costal veins that form the dorsal stripe when the wings are folded over the body.
The females lay 300 to 500 eggs in batches of 7 in the midribs of rice leaves. About 95% of the
eggs are laid on the upper side of the leaves. Eggs are 0.1 mm long, cylindrical, and slightly curved.
They are white when laid but darken subsequently.
The pest is prevalent in Brazil throughout the year with three population peaks in April-May,
September and December-January (Ferreira and Silveira Neto 1979). Studies conducted in Cuba
(Sousa et al., 1977) have shown that the development and multiplication of T. orizicolus are greater
on young than on mature rice plants.
This delphacid can diapause in the egg stage, but nymphs usually hatch in 4 to 8 days. The
population increases as the rice crop matures.
6. LEAF-FEEDING THRIPS
6.1 Rice Thrips, Stenchaetothrips biformis (Bagnall) Thysanoptera: Thripidae)
Synonyms: Bagnallia biformis Bagnall
Thrips oryzae Williams
Chloethrips oryzae (Williams)
Thrips holorphnus Karny
Thrips dobrogensis Knechtel
Baliothrips biformis (Bagnall)
Distribution: Afghanistan, Bangladesh, Bhutan, Burma, China, Europe, India, Indonesia, Japan,
Kampuchea, Korea, Laos, Malaysia, Nepal, Papua-New Guinea, Philippines, South
America, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Agropyron kamojo Ohwi, Alopecurus aequalis Sobol., Arundinella
hirta Tanaka, Cyperus iria L., Cyperus rotundus L., Digitaria ciliaris (Retz.) Koel.,
Echinochloa colona (L.) Link, Echinochloa crus-galli (L.) Beauv., Eragrostis fer-
ruginea Beauv., Festuca parvigluma Steud., Imperata cylindrica (L.) Raeuschel.
Leersia japonica Honda, Panicum repens L., Paspalum scrobiculatum L., Pert-
nisetum japonicum Trinius, Phragmites australis (Cav.) Trin. ex Steud., Zea mays L.,
Zoysia japonica Steud.
Stenchaetothrips biformis is a sporadic pest of rice infesting mostly young plants. It can damage large
areas of paddy fields within a few days because it builds up high populations due to a comparatively
short life cycle. The pest infests both upland and wetland rice crops. In years of a delayed monsoon
or low rainfall during June-July, severe outbreaks of rice thrips have often been reported from India
(Mammen and Nair 1977, Nath and Sen 1978, Velusamy and Chelliah 1980, Gubbaiah 1984),
Bangladesh (Husain 1982) and China (Zu-yin et al., 1978).
Larvae and adults have rasping mouthparts that lacerate the green tissue of leaves. They have
only one mandible that is used to puncture leaf tissue. The maxillae and mouth cone, which form a
Fig. 6.1
DALE 431
tube, are used to suck leaf sap. Young plants, usually of 1 to 2 weeks after transplanting, are the
most affected. Damage becomes evident as fine yellowish lines or silvery streaks on the leaves which
later curl from the margin towards the midrib. The lower leaves are often killed. In severe
infestations, the plants become stunted and wither. But the plants that survive early infestation
usually recover and are able to bear normal grains.
Adult thrips are minute elongate insects, about 1 mm long, dark brown and with pronounced
seven-jointed antennae. They can be either winged or apterous. Wings when present, are long,
narrow, and fringed with fine hairs and lie along the insects’ backs when they are at rest. At the base
of the forewing is a light spot. The males, which are smaller and more slender than the females, are
seldom observed. Reproduction is believed to be parthenogenetic. Adults are day-flying insects
and are not attracted to light. Despite their small size and fragile appearance, thrips can travel long
distances. They migrate during the day and seek out newly planted rice fields or other grassy hosts.
But they prefer to confine themselves to rolled leaves most of the time (Nugaliyadde and Heinrichs
1984). Adult thrips live for about 20 days.
Eggs are reniform and are inserted singly in tissues of the youngest leaves on the side facing the
stem. They are hyaline when laid but turn pale yellow as they mature. A female lays about 25 eggs
in its lifetime. Eggs hatch after 3 days.
The newly hatched larvae are almost transparent but they turn yellowish white after the first
molt. The legs, head and antennae are darker. After a brief resting period, the larvae start feeding
actively on leaves and on the basal parts of leaf sheath. If young rice plants are not available, the
thrips may even feed on rice flowers. The larvae remain on the same plant in which they hatched.
There are 3 larval instars which last for a total of 6 to 8 days.
The prepupae are usually seen in groups. Pupation takes place inside rolled leaves. At this
stage, the appendages are clearly visible and the wings reach up to the fourth abdominal segment.
Prepupal and pupal stages together last for 3-4 days.
Thrips are normally abundant during the dry season when there is little or no rainfall. There-
fore, plants that are transplanted preceeding the rains become the main target of thrips attack.
Prolonged rainy periods and heavy rains reduce the pest population (Zu-yin et al., 1978). The
outbreak of thrips in Bangladesh in 1982 was attributed to a long winter followed by a Summer
drought (Husain 1982).
In China, the thrips have 10 generations a year. The threshold temperature for temperate
regions is 11.5 ± 12°C; the optimum temperature for development ranges from 15 to 25°C. Adults
of the overwintering generation emerge in the middle of April and lay eggs on weeds. During the
early half of May, adults of the second generation invade the fields and infest rice plants. Maximum
crop damage, however, is caused by the second and fourth generations from the middle of May to
the middle of July. The peak population is attained in the middle of June.
Seasonal abundance in tropical countries is slightly different probably due to the change in
seasons and planting times. In northern India, the thrips appear on rice plants the first week of
August. The population reaches its peak by the third week and declines thereafter. The crop that
is planted in early July has less infestation compared to that planted in late July or early August.
Older plants, particularly those at 30 days after transplanting, have very low populations (Senapati
and Satpathy 1982).
Nugaliyadde and Heinrichs (1984) studied the effects of plant age, adult density and tempera-
ture on fecundity and population growth of the rice thrips. Plant age did not influence female
longevity. However, male longevity was significantly less on the two-leaf-stage plants than on the
six-leaf-stage plants. Daily mean fecundity was significantly higher on rice seedlings at the third,
fourth, and fifth-leaf stages than on seedlings at the first, second, cr sixth-leaf stages. Females on
plants with low population density levels produced significantly more progeny. The total progeny
produced per female over 14 days of the early reproductive period was not affected by the
temperatures tested in the study (23, 26, 29 and 32°C).
7. LEAF-FEEDING LEPIDOPTERA
7.1 Rice Leaffolder, Cnaphalocrocis medinalis (Guenee) (Lepidoptera: Pyralidae)
Synonym: Cnaphalocrocis jolinalis Led.
Distribution: Afghanistan, Australia, Bangladesh, Bhutan, Brunei, Burma, China, India, In-
donesia, Japan, Kampuchea, Korea, Laos, Madagascar, Malaysia, Nepal, Pakistan,
Papua-New Guinea, Philippines, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Brachiaria mutica (Forssk.) Stapf, Echinochloa colona (L.) Link,
Eleusine coracana (L.) Gaertn., Isachne dispar Trin., Leersia hexandra Sw., Pen-
nisetum pedicellatum Trin., Saccharum officinarum L., Sorghum bicolor (L.)
Moench., Triticum aestivum L., Zea mays L.
The rice leaffolder, Cnaphalocrocis medinalis, earlier considered as a minor and sporadic pest of
rice in many Asian countries, appears to have become increasingly important with the spread of
high-yielding rice varieties and accompanying changes in cultural practices. Misuse of insecticides
Fig. 7.1
DALE 433
and excessive use of nitrogenous fertilizers have been cited as the cause for high leaffolder
populations (Dhaliwal et al., 1979). Outbreaks have been reported from India (Yadava et al., 1972,
Chatterjee 1979), southern Japan (Hirao 1981) and Vietnam (Bautista et al., 1984)
Infestation usually occurs during late growth stages of the rice crop. The larvae fold the leaves
and scrape the green tissues of the leaves from within and cause scorching and leaf drying. Each
larva is capable of destroying several leaves by its feeding. Under heavy infestation, each rice plant
may have several rolled leaves, which severely restricts its photosynthetic activity. When plants are
attacked in the bootleaf stage, grains are partially filled. A close correlation between the intensity
of leaffolder attack and loss in grain yield very often exists (Upadhyay et al., 1975). Bautista et al.,
(1984) have clearly shown that yield loss due to rice leaffolder is positively related to the percentage
of damaged leaves. In their studies, yield was significantly decreased at 17.5% damaged leaves
resulting in 16.5% yield loss, and a 21.3% yield loss occurred with 26.6% damaged leaves. Sellamal
Murugesan and Chelliah (1983) reported that a 10% increase in flag leaf damage by the leaffolder
reduces grain yield by 0.13 g per tiller and the number of fully-filled grains by 4.5%. However, rice
plants sometimes compensate for flag leaf damage by increasing the photosynthesis rate of the leaf
adjacent to the flag leaf (Derui 1984). Leaf feeding also predisposes the plants to fungal and
bacterial infections.
Moths are nocturnal and in the day time, they are usually seen resting on the under surface of
leaves and on stems. During the early morning hours, moths are often active but their activity
diminishes as the sun increases in severity. The attraction of moths to light is rather poor.
Adult moths are yellow-brown in color, small, and are 10 to 12 mm long with a wing expanse of
13 to 15 mm. The wings take the shape of an equal sided triangle when at rest. The forewings have
three dark oblique lines of varying lengths. The hind wings have a broad anal area. The tibiae are
tufted with black hairs which are prominent in the male but not in the female. The tip of the
abdomen is blunt in the male but pointed in the female (Lingappa 1972). The female attracts its
mate with a pheromone and generally mates between dusk and midnight. Adults hide on the host
plants during the day to escape predation by buds and only take short flights when disturbed. Adult
longevity is about a week.
Egg-laying starts one or two days after mating. Flat, oval and whitish yellow eggs are laid singly
or in rows, parallel to a midrib on both surfaces of young leaves and rarely on stems. The eggs are
laid in batches of 10 to 12, the biggest batches being laid on the fourth to seventh nights after the
emergence of moths. Each female moth may lay about 300 eggs during its lifetime. The incubation
period varies from 3 to 6 days.
The newly hatched larva is white translucent with a light-brown head. The body, however, turns
green once the larva starts feeding. After hatching, it crawls to the base of the youngest unopened
leaf and begins to feed. The second instar migrates to an older leaf and folds the leaf together.
Some of the newly hatched larvae suspend themselves by silken threads from the leaf tip and
disperse to other plants by wind (Velusamy and Subramaniam 1974). Mostly, only one larva is
found in a leaf fold (Yadava et al., 1972). Leaf tubes are generally made by folding a single leaf but
cases are also observed where two or three leaves are used for making one fold. There are generally
five larval instars in tropical countries. The frequency of larval molting is affected more by the
growth stage of the rice plant than by the temperature in temperate regions (Wada 1979).
Details regarding the mode of construction of leaf folds by larvae have been reported by
Ramasubbaiah et al. (1980) and Fraenkel and Fallil (l981). The larva selects a point on a leaf about
15 cm from the tip and lines the surface for about 6 cm with a salivary secretion as a flooring for the
leaf fold. Then the larva swings its head in quick succession between two opposite leaf margins,
affixing a silk thread at each touch-down. The tiny threads from about 100 such movements fuse
together to a firm band which on drying contracts. Ten to 25 such bands are produced on a leaf at
distance of a few millimeters. The entire process requires about 45 minutes, the larva resting several
times during the period. The nature of the folds depends on the age of the crop (Rajamma and Das
1969). In rice seedlings and young plants, 3 to 4 leaves of adjacent plants are webbed together
longitudinally to make a tubular fold. In the more mature plants, folds are made out of single leaves
that are folded either longitudinally and rarely transversely. Only fully grown larvae make a
transverse leaf roll. In the earhead stage, the larvae make a shelter by webbing together the bootleaf
and the earhead.
The full grown larva is about 16 mm long and 1.7 mm wide across the thorax. It is yellowish
green in color with a dark brown head and prothoracic shield. Mature larvae jump or wiggle rapidly
when touched. The larval period lasts for 15 to 25 days.
Pupation takes place inside the leaf roll in loosely woven strands of silk. The newly formed
pupa is slender and greenish brown and turns brown later. The moth emerges after 6 to 8 days.
Overwintering has been confirmed only in Okinawa, Japan (Miyahara 1981). As the insect does
not hibernate, low temperature in winter affects overwintering survival. Both eggs and larvae
cannot survive winter in Japan (Sato and Kishino 1978). Very little is known about the cold-hardi-
ness of the leaffolder.
The leaffolder moths have been recorded in tropical countries throughout the year, although
they are most abundant during the wet season. In countries such as Japan, with a cold winter, moths
are first seen in the paddy field in mid-June (Wada et al. 1980). Then their population gradually
increases until late August after which the larval density decreases and very few eggs are seen on
rice leaves. At the end of the season, the generations overlap each other as indicated by the various
instars seen in a field at the same time.
C. medinalis moths migrate long distances. In 1977, many moths were captured on a vessel in
thc East China Sea far from land (Kisimoto 1978). During migration, most of the female moths
remain unmated and once they settle on rice plants, they immediately mate and start laying eggs
(Miyahara 1980). The insect is believed to take the same migration route from China to Japan and
Korca as Sogatella and Nilaparvata do (Miyahara 1981).
High humidity and optimum temperatures are conducive ecological factors for the rapid
multiplication of the leaffolder (Pathak 1975). The threshold temperatures of development for egg,
larva and pupa are 12.5, 12.2 and 142°C respectively (Wada and Kobayashi 1980). The larval period
is significantly extended when the larvae are reared on leaves of rice plants at the ripening stage.
DALE 435
There are many other factors that play a role in the development and population increase of the
rice leaffolder. It has been shown by Hanifa et al. (1974) that silica in host plants plays an important
role in the feeding activity of leaffolder larvae. Arrangement of silica chains in the leaf varies in
resistant and susceptible cultivars. Closer silica chains, high epidermal silica deposition, heavy
deposition of silica in the intercostal area, and single or double rows of silica in resistant varieties
provide a sufficient mechanical barrier to the feeding by caterpillars.
There are varying reports on the effect of nitrogenous fertilizers on rice leaffolder incidence.
Many workers (Regupathy and Subramanian 1972, Michael Raj and Morachan 1973, Subbaia and
Morachan 1974, Chandramohan and Jayaraj 1977, Upadhyay et al. 1981) have shown that the
incidence of leaffolder is positively correlated to the level of nitrogenous fertilizers. A combined
application of high doses of N, P and K has been shown to alter the mineral metabolism of rice
plants in favor of leaffolder development and multiplication (Regupathy and Subramanian 1972).
But when K was applied alone, leaffolder infestation was significantly reduced (Subramanian and
Balasubramanian 1976).
It is interesting to note that the damage is more severe in shady areas particularly near the
levees (Velusamy and Subramaniam 1974). Close crop growth consequent to heavy manuring is also
conducive for leaffolder activity.
Adult feeding influences the fecundity and subsequent population build-up of the pest. The
female moths require an exogenous source of sugar for the production of eggs. The increase in
fecundity under laboratory conditions resulting from the utilization of hopper-excreted honeydew
indicates that adult moths can use honeydew as a source of sugar in rice fields (Waldbauer et al.
1980)
7.2 Rice Leaffolder, Marasmia exigua (Butler) (Lepidoptera: Pyralidae)

Synonym: Susumia exigua (Butler)
Distribution: Australia, Bangladesh, Brunei, Fiji, Guam, India, Japan, Korea, Malaysia, Solomon
Islands
Fig. 7.2A
Three Marasmia species, M. exigua, M. patnalis (Bradley) and M. ruralis (Walker), attack rice. They
are distinguished by wing markings. M. exigua (Fig. 7.2A) is the most common pyralid pest of rice
in Fiji. The habits and the biology of the insect are very similar to those of Cnaphalocrocis medinalis.
The adult moth has three long, dark bands on the forewings while C. medinalis has two long and one
short band. Earlier the species was misidentified as Marasmia venilalis (Walker).
The adult moth is about 7 mm long with a wing span of 13 mm. Its forewings have light yellow
and brown markings. The gravid female moths prefer to lay eggs on the green parts of tall rice
plants. Eggs are laid overlapping one another, in a row parallel to the midrib of the leaf. The eggs
in a cluster average 3 but may be as high as 11. They are flat and translucent. The incubation period
is about 5 days.
The newly-hatched larvae tend to be gregarious for some time and then start feeding on the
leaves. The second instars draw the leaf margins together and stitch them with contractile silk.
Both ends of the leaf roll are left open. The larva feeds on the green tissue leaving conspicuous
white scars on the leaf blade. There are six larval instars and development takes approximately 4
weeks. The larva differs from that of C. medinalis in that the apex of the pronotum is convex while
it is straight in C. medinalis.
Pupation takes place within the leaf roll. The pupa is not enclosed in a cocoon but is surrounded
with silken frass and fecal plugs, Pupae are 8 mm long and 1.7 mm wide. The pupal period is about
9 days (Hinckley 1963). The insect is most abundant in Fiji during April and May.
Marasmia patnalis (Fig. 7.2B), a closely related species, has recently been reported as a pest of
rice in Southeast Asia (Bradley 1981). The species is shade-loving and is considered as a major pest
of rice in Sri Lanka. Forewings are pale yellow with a greyish marginal band. Hind wings are also
pale yellow but whitish basally. The male moth is readily distinguished by the presence of a
prominent patch of dark brown and shining adroconical scales at the middle of the forewing costa.
The life cycle of M. patnalis was studied in the Philippines by Joshi et al. (1985). Eggs are laid
singly or in groups of 2 to 9 on the upper side of the leaves but sometimes eggs are also seen on leaf
Fig. 7.2B
DALE 437
sheath. The incubation period is 4 days. First instar larvae scrape the leaf surface. The second
instar larvae fold leaves and start feeding from within. The larval period is about 23 days.
The pupal period is 9 days and takes place within a silken cocoon, most commonly between
leaves that have been stitched together. Adult emergence takes place at night.
7.3 Rice Caseworm, Nymphula depunctalis (Guenee) (Lepidoptera: Pyralidae)

Synonym. Nymphula stagnalis (Zeller)
Distribution: Argentina, Bangladesh, Bhutan, Brazil, Burma, China, Gambia, Ghana, India,
Indonesia, Kampuchea, Laos, Madagascar, Malaysia, Malawi, Mauritius, Mozam-
bique, Nepal, Nigeria, Papua-New Guinea, Philippines, Sri Lanka, Taiwan,
Uruguay, Venezuela, Vietnam, Zaire
Host plants other than rice: Brachiaria murica (Forssk.) Stapf, Brachiaria ramosa (L.) Stapf,
Cynodon dactylon (L.) Pers., Cyperus iria L., Cyperus rotundus L., Cyrtococcum
patens (L.) A. Camus, Echinochloa colona (L.) Link, Isachne dispar Trin., Leersia
hexandra Sw., Panicum repens L., Paspalum conjugatum Berg.
Nymphula depunctalis is an important pest of irrigated and rainfed wetland rice in the Orient. Even
though rice is its preferred host plant, the insect infests various other grasses as well. The pest
occurs regularly in low populations, but sporadic increases in population result in intense defolia-
tion of plants. An extensive outbreak occurred in Kerala, south India in 1968 (Joseph 1969). The
infestation is more severe on dwarf, compact, heavy-tillering, high-yielding varieties (Kittur and
Chauhan 1974).
Damage is caused by larvae cutting off the leaf tips for making the characteristic leaf cases and
by the removal of the green tissue. Defoliation occurs in rice plants before the maximum tillering
stage. Plants that have recently been transplanted are preferred but the larvae may infest the
nursery. It does not occur after maximum tillering. Larvae scrape the leaf tissues, leaving only the
papery upper epidermis. Generally, several larvae attack the same rice plant and cut off most of the
Fig. 7.3
leaf tips for constructing larval cases and scrape the remaining leaves. Caseworm damage can be
distinguished from that of other pests by the ladder-like appearance of the removed leaf tissue,
resulting from the back and forth motion of the head during feeding. Also, the pattern of damage
is not uniform because the larvae in the cases are carried to one side of the paddy by winds or water
currents. Attacked plants often become stunted.
Many workers have studied the bionomics of the rice caseworm in India (Srivastava et al. 1970,
Kittur and Chauhan 1974, Viraktamath et al. 1974, Pillai and Nair 1979) and in the Philippines
(Sison 1938).
Moths are nocturnal in habit and are attracted to light. They are delicate, white with fuscous
markings and black specks on wings. Female moths are larger than the males. Mating mostly takes
place during the night; egg laying occurs the next night. Usually all the eggs are laid during a single
night. Each female lays about 60 eggs and may die 2 - 3 days after oviposition. The male moths live
for 4-5 days.
The freshly laid eggs are light yellow, smooth and spherical. They are laid in batches of about
20 on the underside of leaves floating on the water. The eggs turn dark yellow as they mature. The
incubation period lasts for 4 days.
Larvae are transparent green in color with light brownish orange heads. The first instar larva
starts feeding on the green tissues by scraping the leaf surface. It then moves to the leaf tip and cuts
a slit on the margin at a point 2 - 3 cm below the tip and makes a fold. Then the larva makes another
cut about 1 cm below the first and rolls the margin with silk to form a tubular case. The inside of
the case is lined with silk to hold a thin film of water which is essential for both respiration and
preventing desiccation of the larva. A new case is constructed after every larval molt. The case that
is carried along by the caterpillar whenever it moves helps the larva to float on the water surface.
During the daytime, larvae hide in cases while at night they crawl up rice plants and feed with their
head and legs protruding from the case. Feeding is reduced under dry conditions and the presence
of water droplets on the leaves is helpful for larval feeding. Larvae are 20 mm long when fully
grown, and are semi-aquatic in habit with lateral, slender, filamentous gills.. The larval stage lasts
for 15 - 30 days.
Pupation takes place inside the larval case with both ends closed and attached to the rice stem
just above the water level. The case has an inner silk lining with a slit-like opening at the anterior
end. The fresh pupa is milky white which gradually changes to light yellow. The adult moth
emerges in 4 - 7 days.
In south India, moth populations reach two peaks, one during November-December and
another in May-June. These peaks coincide with periods of high rainfall and high humidity.
Variations in temperature do not appear to be correlated with pest populations (Pillai and Nair
1979). In northern India, the infestation is more severe in the months of August - November
(Srivastava et al. 1970). The insect occurs in overlapping generations and does not hibernate.
N. nympheata, commonly called China mark moth, is a related species of N. depunctalis
distributed mainly in Italy and Hungary. Five generations occur per year in Italy, the first being the
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most important as it attacks very young plants. A smaller rice caseworm, N. vittalis Bremir has been
reported from Japan, Korea, Manchuria, and USSR. The larvae fold leaves of the plant and feed
from within.
7.4 Green Horned Caterpillar, Melanitis leda ismene Cramer (Lepidoptera: Satyridae)
Distribution: Bangladesh, Bhutan, Burma, China, India, Japan, Kampuchea, Laos, Malaysia,
Nepal, Pakistan, Philippines, Thailand, Vietnam, West Africa
Host plants other than rice: Imperata cylindrica (L.) Raeuschel, Panicum maximum Jacq., Sac-
charum officinarum L., Sorghum verticilliflorum (Steud.) Stapf
Melanitis leda ismene is a minor pest of rice. The pest occurs in all rice environments but is more
prevalent in rainfed areas. Deep water rice in Bangladesh is also often infested by this insect (Alam
1974).
Fig. 7.5
The adult is a large, dark brown butterfly measuring about 7.5 cm across expanded wings and
with a body 2.2 cm long. It flies at dusk making darting movements among the rice plants. There
are large, round spots on the upper surface of the forewings. The underside is ochreous brown with
dark transverse stripes and target-like spots; hind wings have six prominent spots on their ventral
side. The wings are folded above the body when the insect is at rest.
Pearl-like eggs are laid singly or in rows on rice leaves and are 1 mm in diameter. Each female
lays 50-100 eggs in a lifetime of about 2 weeks. They hatch in about 4 days. The caterpillars feed on
the margins and tips of leaves and remove leaf tissue and veins. Damage in similar to that of
grasshoppers and armyworms. The yellow-green larva blends into the rice foliage, and is difficult to
see, in spite of its large size. The body is covered with small, yellow hairs. The head is flat and
quadrangular with a pair of brown horns; at the posterior extremity of the body are two slender
processes. Pupation takes place on the rice leaf. The pupa is dark green, smooth, and hangs by its
anal extremity from the leaf. The pupal period lasts for 10 days. In Bangladesh, the pest is observed
to reach its peak population during October (Alam 1974).
7.5 Rice Skipper, Parnara guttata (Bremer et Grey) (Lepidoptera: Hesperiidae)

Distribution: Bangladesh, China, India, Indonesia, Japan, Kampuchea, Korea, Laos, Malaysia,
Pakistan, Philippines, Taiwan, Thailand, Vietnam
Host plants other than rice: Carex olivacea Boott., Phragmites communis Trin.
Parnara gutatta (Bremer and Gray) and Pelopidas mathias (F.) are the most common rice field
species of the day-flying skippers.
Fig. 7.5
The rice skipper, P. gutatta, is one of the major pests of rice in China (Chiang 1977). Outbreaks
of the pest also often occur in the paddy fields of central and western Japan. P. guttata is the most
dominant species among the rice skippers in Japan and China (Masuzawa et al. 1983). It occurs in
all rice environments but is mostly a pest of irrigated rice.
The caterpillars feed on leaves from the margins inwards and then parallel to the midrib which
is often left uneaten. In addition, the larvae tie, with silken threads, the two edges of the same leaf
or two adjacent leaves together to form a tube in which they live. Damage is severe in young
transplanted rice seedlings and the attack may continue till the plants mature. In severe cases, the
plants do not recover. Grain quality is adversely affected by the pest.
Adult butterflies are olive brown and 10 to 15 mm long with a wing span of 20-44 mm .There
are 5-9 small whitish specks on the forewings and similar spots, 2 to 4 in number, on the hindwings.
The butterflies are swift fliers being active only during the day.
The white spherical eggs are glued singly on leaf blades by the female moths. The eggs hatch in
3-4 days. Larvae rest at the base of the plants during the day and feed on leaf blades at night. The
full grown caterpillar is pale green, and has a dark head with a W-shaped black mark on the rear.
The mature larvae often move from one rice plant to adjacent plants. There are 5 larval instars in
Taiwan, but the fifth instar larvae molt once more in Japan (Ishii 1980). The larval period is about
a month. Pupation takes place inside the leaf tubes. The light brown or light green pupa rests in a
pad of silk and has a pointed end which is attached to the folded leaf. Adults emerge after 8 to 10
days.
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Adult polymorphism is exhibited by the skipper butterflies. Hasegawa (1975) has reported that
the species has two seasonal forms; the yellowish brown ‘summer form’ and the dark brown ‘autumn
form’. Conditions of temperature and photoperiod prevailing during the larval stage mostly deter-
mine the morphological characters of the adult butterfly. A long photoperiod induces pale wing
coloration and small size of adults. But low temperature induces larger size, stronger melanization
and more conspicuous spots on the wings (Ishii and Hidaka 1979). The ecological factors such as
photoperiod and temperature may also affect the physiology of the migrating butterflies. Females
reared under 14-hour photoperiod, which corresponds to the migratory adults growing in early
autumn, have higher flight capacity, lower fecundity and a longer pre-ovipositional period than
those under a 16-hour photoperiod which corresponds to the non-migratory adults in summer (Ono
and Nakasuji 1980).
Parnara guttata is well known as a migrant butterfly in central and western Japan. Second
generation butterflies fly en masse to the warmer southwest in early autumn for hibernation. Eggs
of the third generation are laid on grasses and the larvae develop on those grasses in autumn
(Nakasuji et al. 1981). Average migration distance is less than 100 km (Nakasuji and Ishii 1983).
However, some of the migrants fly more than 500 km over the Pacific Ocean (Asahina and Turuoka
1969) and may thus reach the southwest islands of Japan or even Taiwan.
7.6 Rice Skipper, Pelopidas mathias (F.) (Lepidoptera: Hesperiidae)
Distribution: Bangladesh, China, India, Indonesia, Japan, Kampuchea, Korea, Laos, Malaysia,
Pakistan, Philippines, Taiwan, Thailand, Vietnam
Host plants other than rice: Cynodon dactylon (L.) Pers., Eleusine coracana (L.) Gaertn., Im-
perata cylindrica (L.) Raeuschel, Livistona chinensis R. Br., Miscanthus sinensis
Aderss., Paspalum scrobiculatum L., Sorghum halepense (L.) Pers., Zea mays L.
Pelopidas mathias is a minor pest of rice with an extensive distribution in the rice growing regions
of the world. It occurs in all rice environments but in Japan and Taiwan upland rice is the most
affected.
Fig. 7.6
Damage is caused by the larvae that defoliate the rice plants similar to that of P. guttata. The
newly hatched larvae feed on tender leaves. Then they roll up and stitch together two or more leaves
that are eaten from the margin inwards. Generally, only one larva is found in a fold (Teotia and
Siya Nand 1966).
The adult butterfly differs from P. guttata in usually having only 2 spots in the discal cell of the
wing and 2 groups of 3 white specks, one anterior and one marginal from the first group. The hind
wings are olive brown, usually without spots. Mating generally takes place during morning hours.
Eggs, averaging 90, are laid singly on the upper surface of leaves.
Eggs are smooth and creamy white in color. They are hemispherical in shape with a flat base.
The incubation period ranges from 3 to 6 days.
Larvae are pale green with a vertical red streak on either side of the head. There are five larval
instars. The full-grown caterpillar measures 30 mm in length and 3.5 mm across the middle of the
body. The larval stage lasts for 13 to 26 days.
Pupation takes place in a silken web spun inside the leaf fold. The pupa is at first green in color
with stripes clearly visible on the back, but changes to dark brown later. It is cylindrical and
gradually tapering towards the hind end. The cremaster is transversely flattened and bears on its
posterior margin a number of small spines. The pupal skin is thin and almost transparent. The
adult butterfly emerges in 7 to 12 days after pupation.
In India, the insect passes through four overlapping generations. The adults of the first
generation appear in the fields about the third week of August and those of the second generation,
the second week of September. The butterflies of the third and fourth generations appear towards
the third week of October and last week of November, respectively.
P. mathias is most active from August to November and with the advent of winter a high
percentage of the larvae hibernate thereafter. The overwintering pupae remain inside the cocoons
in folded leaves. The butterflies emerge from them in the spring when the weather becomes
warmer.
7.7 Rice Ear-cutting Caterpillar, Mythimna separata (Walker) (Lepidoptera: Noctuidae)

Synonyms: Leucania separata (Walker)
Pseudaletia separata (Walker)
Distribution: Afghanistan, Australia, Bangladesh, Bhutan, Burma, China, India, Indonesia,
Japan, Kampuchea, Korea, Laos, Malaysia, Nepal, New Zealand, Pakistan, Papua-
New Guinea, Philippines, Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Avena sativa L., Brachiaria mutica (Forssk.) Stapf, Brachiaria setigera
(Retz.) C. E. Hubb., Cynodon dactylon (L.) Pers., Cyperus rotundus L., Echinochloa
colona (L.) Link, Eleusine coracana (L.) Gaertn., Hordeum vulgare L., Nicotiana
tabacum L., Panicum antidotale Retz., Pennisetum purpureum Schumach.,
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Fig. 7.7
Saccharum ofjicinarum L., Setaria italica (L.) Beauv., Sorghum bicolor (L.)
Moench., Triticum aestivum L., Zea mays L.
Mythimna separata, a minor pest of rice until recently, has become a serious pest in several parts of
Bangladesh, China and India. Severe outbreaks of this insect have been reported by many workers
(Alam and Nurullah 1977, Chaudhary and Singh 1980, Barwal l983). The attacks may be sporadic
but if heavy, the entire crop may be lost. The pest occurs in all rice environments but is most
abundant in upland and rainfed wetland environments.
The early instar caterpillars first feed on dried leaf tissues and later move on to green leaves for
feeding. They also eat the lemma and palea of the developing grains as well as the anthers of flowers
(Papel et al. 1981). Mature larvae, from 4th to 6th instars, become gregarious and feed voraciously
on young leaves at night. The sixth and final instar larvae cut off rice panicles from the peduncle,
causing the most serious losses to the rice crop. In cases of severe infestation, the damage may go
up to 60% or even more. Katiyar and Patel (1969) reported grain losses up to 1,640 kg/ha in Madhya
Pradesh, India. Besides the direct loss, many farmers, anticipating the pest incidence, are tempted
to harvest the crop prematurely. This produces large quantities of chaffy grains which may reduce
the market price of the product. Moreover, a large number of grains and panicles are seen lying in
the fields as the result of feeding damage. Experiments with third instar caterpillars indicated that
for each increase of 1% leaf area consumed, there is a reduction of 0.058 cm in panicle length, 0.88
grain/panicle and 0.072 g yield/hill at the booting stage of the crop (Alam et al., 1980). But these
values are liable to change with the instar of larva and rice cultivar used for the study. Khamparia
et al. (1981), working with a different variety and fifth instar caterpillars, reported that there was a
yield reduction of 0.41 g/hill with an increase of every one percent of foliage loss by the larvae.
The adult is a stout, pale red brown moth which measures 20 mm long with a wing expanse of
40 mm. The forewings have two pale round spots, and the hind wings are dark on top and white
underneath. The moths feed on nectar from flowers and honeydew excreted by mealy bugs and
other Homoptera. The female moths start egg laying in three days. Eggs are laid in batches of
about 100 between the leaf sheaths and stems of rice plants or grasses. Males live for only 3 days
while the female moths live longer.
Eggs are spherical, green-white when fresh, gradually turning pale yellow and finally black. The
egg period varies from 5 to 7 days. Newly-hatched caterpillars are dull white, but later turn green.
The head is orange or brown. Four longitudinal light grey to black stripes run along the body. The
larvae are nocturnal in habit and during the day they hide in loose soil, under trash, in stubbles, in
leaf sheaths and in whorls. During the night, they become active and resume their feeding activity.
The entire larval stage lasts for about 28 days. The larvae pupate in the soil. Sometimes naked
pupae are also observed among the tillers. The pupal period is 8-11 days.
Plants with dense foliage coupled with heavy tillering are most commonly attacked by the pest
(Pate1 1979). Heavy application of nitrogenous fertilizers produces more succulent and green
plants in which the armyworm population is found to be higher (Koyama 1966). Survival rate and
fresh weight of larvae are both high on these plants. It is interesting to note that plants of the
aromatic rice variety 'Basmati' are seldom attacked by the pest.
Many conditions have been reported ideal for outbreaks of this pest in different countries. In
India, heavy late rains after a long drought often result in widespread oviposition and larval
development. In Bangladesh also, major outbreaks are always associated with a long drought
followed by wet weather (Alam and Nurullah 1977).
In China, the moths migrate to the north from southern China in the spring and fly back in the
autumn. The overwintering boundary is considered between 32 and 34°N latitude (Li Kuang-po and
Chen Rui-lo 1983). There are four important migrations annually. The first (March-April) and the
second (May-June) are from low latitude and low elevation to high elevations; the third and fourth,
vice versa.
Rice may not be the ideal plant for M. separata. On an overall basis of development, reproduc-
tion, survival and growth, Sorghum was found to be the most suitable host plant, followed by
Pennisetum (Alam and Khatri 1980). Larvae develop faster on maize than on rice (Tripathi et al.
1982).
In India, adult moths start coming to rice fields by early May. The population of the first
generation caterpillars is small and the damage is not conspicuous. The second generation larvae
are active from middle to late June. The damage of the third generation caterpillars is mostly
restricted to millets (Purohit et al. 1971). The rice ear-cutting caterpillar has 3-4 generations a year.
7.8 Common Armyworm, Pseudaletia unipuncta (Haworth) (Lepidoptera: Noctuidae)

Synonyms: Cirphis unipuncta (Haworth)
Pseudaletia unipuncta (Haworth)
Distribution: Southern Canada, USA, Central and some South American countries, Southern
Europe, West Africa
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Fig. 7.8
Host plants other than rice: Wheat, oats, barley, corn, forage grasses
Mythimna unipuncta is a cosmopolitan pest of grasses. Rice is not an ideal host for the armyworm.
However, it occurs sporadically in epidemic numbers on rice in California (Lange and Grigarick
1970).
The armyworm infestations occur relatively late in the growing season, 3 to 4 weeks before
heading. During the early instars, larvae skeletonize tender, young leaves or the inside of leaf
sheaths. Later, the larvae remove irregular portions of the entire leaf starting from the edge, but
leaving the midvein intact (Rice et al. 1982a). Feeding damage is generally found on the distal
portion of rice leaves. Rice et al. (1982b) have shown that defoliation averaging 25 to 30% by
armyworm larvae resulted in yield reductions up to 50%.
The adult moths are light red to pale brownish and have a hairy body with black spots and
patches. They have a wing span of about 35 to 45 mm. There is a distinctive white spot in the middle
of each forewing. Moths are nocturnal in habit, positively phototropic and remain inactive during
daytime. Mating and oviposition usually take place after sunset. Generally, the males die shortly
after mating. The female moths lay eggs for about 5 days.
The eggs are laid in several rows, each mass consisting of 90 - 230 eggs. They are round, pale
yellow and 0.6 - 0.7 mm in diameter. The incubation period is 7-9 days.
Larvae are nocturnal and hide under leaves and debris during the day. Freshly hatched larvae
are dull white with a brownish black head. There are 6 larval instars. Of the total amount of leaf
area consumed, more than 90% is eaten by the fifth and sixth instar caterpillars (Rice et al. 1982a).
The full grown larva is usually dark or greeenish grey in color and has a color yellow stripes just below a line
of a prominent black spiracles. The larval duration is on is 24-28 days
Pupation takes place in individual earthen cells in the soil. The pupae are light amber at the
beginning and turn dark brown just before adult emergence. The average pupal period lasts for 16
days. There are five generaations of the pest in a year; usually only one generation is spent on rice.
7.9 Rice Green Semiloopers, Naranga aenescens Moore, and Naranga diffusa Walker
(Lepidoptera: Noctuidae)
Distribution: China, India, Indonesia, Japan, Korea, Malaysia, Philippines, Taiwan, Vietnam (N.
aenescens); India, Indonesia, Malaysia (N. diffusa)
Host plants other than rice: Cyperus diffusa L., Echinochloa colona (L.) Link, Leersia hexandra
Sw., Leptochloa chinensis (L.) Nees, Paspalum conjugatum Berg., Sorghum bicolor
(L.) Moench.
Naranga aenescens (Fig. 7.9) is an important pest of rice in eastern Asia (Ando et al. 1980). But N.
diffusa is only a minor pest. The caterpillars feed on the leaves causing defoliation which often
results in a great loss in rice yields. N. aenescens is only found in wetland environments and is
abundant in the rainy season.
Fig. 7.9
The adult is a small moth with a wing expanse of 17 mm. The forewings of the female are golden
yellow each with two transverse brown bands; hind wings are dark brown. The male moth is dark
purple-red. The moths which emerge from overwintering pupae late in spring mate in the daytime.
Adults hide in rice fields or grassy areas at the base of the plants during the day and become active
at night. They are highly attracted to lights.
Each female lays 50-100 eggs during its lifetime of 3 to 5 days. Spherical eggs are laid on leaves
in groups of up to 15 each. They are yellow when newly laid and later develop purple markings.
The larva is a semilooper, light green with white stripes. The larvae prefer actively growing
plants, from the seedbed through the tillering stage. Young larvae scrape the tissue from the leaf
blades and older larvae eat large areas at the edge of leaves to form notches. Young plants can often
recover but yield loss occurs when N. aenescens occurs in combination with other non-defoliating
pests. The population density during the larval stage of N. aenescens is known to be one of the
factors responsible for inducing larval diapause (Kishino and Sato 1975). The larva forms a pupal
chamber by folding a rice leaf over and securing it with silk. The pupa is light brown and smooth
DALE 447
bodied. Development from egg to adult takes 18 to 21 days in the Philippines (Pantua and Litsinger
1984). There is often a short pupal diapause for the last generation.
Cloudy days accompanied by low temperature favor pest outbreaks. There are 4-5 generations
in a year except in northern Japan where there are only 3 - 4.
7.10 Green Hairy Caterpillars, Rivula atimeta (Swinhoe), and Rivula basalis Hamps,
(Lepidoptera: Noctuidae)
Distribution: Philippines (R. atimeta); India (R. basalis)
The hairy caterpillars defoliate rice at the vegetative stage. In addition to rice, R. atimeta (Fig. 7.10)
feed on grassy weeds. Damage is similar to that of the green semilooper. Damaged plants often
recover when defoliation is not severe but high populations can kill seedlings, especially when they
occur in combination with other insect pests. The description of the pest is based on the paper by
Sunio et al. (1983).
Fig. 7.10
Adult moths of both sexes are white-grey or light brown and are active at night. During the day,
they hide under vegetation with their heads pointed down.
Spherical, pale green eggs are laid singly in rows on both sides of a leaf. The number of eggs
laid per female moth averages 130. The incubation period is 3 - 5 days. Adult longevity of male and
female moths is 4 and 5 days respectively.
The larva has long thread-like hairs on its green body and the average larval period is 13.5 days.
When ready to pupate, larvae spin a cocoon of silk on a leaf blade. The pupal period is 5 days.
7.11 Fall Armyworm, Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae)

Synonym: Laphygma figiperdu (J. E. Smith)
Distribution: Central and South America, Guyana, Mexico, USA, West Indies
Host plants other than rice: Grasses, cereals, legumes, cotton, cabbage, tobacco
Fig. 7.11
The attack of Spodoptera frugiperda is perhaps the most spectacular because of its sudden ap-
pearance in the rice fields and the quick devastation of the crop. The insect is popularly called ‘fall
armyworm’ as it appears in the northern parts of USA in late summer or fall. It is a polyphagous
pest attacking many grasses and crop plants.
Newly-emerged caterpillars scrape leaf tips of young rice plants; as a result, the leaf tips appear
burnt. The older larvae defoliate the crop. Infestations appear as isolated, circular patches in the
rice field and seldom affect the whole field. All plants, except those eaten completely, normally
recover and produce grains. Generally, the pest appears on young rice plants and does not remain
in the same field for long. They march gregariously seeking fresh fields.
Adult moths are ash-grey in color with forewings having irregular white and pale grey markings.
The hind wings have a pinkish white luster bordered by a light brown band. The moths are 2 cm
long with a wing expanse of 4 cm. They are nocturnal, strongly attracted to light, and live for 2 - 3
weeks. The female moth flies into rice fields at night and deposits clusters of eggs on young rice
plants. There are 50 to 300 eggs per egg mass which is covered with hairs and scales from the moth‘s
body. Eggs are round, about 0.5 mm in diameter, dull white in color and with a ridged surface.
Freshly-emerged larvae are white with a shiny black head. They later turn green to black with
a white stripe down each side below which there is a yellow band. The caterpillars may crawl inside
the leaf sheaths or move from one plant to another suspended by a silken thread. They become
full-grown in 2 - 3 weeks.
Pupation occurs in the soil between leaves that are spun together. Most larvae that develop on
flooded rice never pupate (Bowling 1967). The pupa is yellowish brown and darkens as it ages. The
pupal period is 10 - 14 days.
In Guyana, the pest is more serious in spring (October to February) than on the autumn crop
(May to August) especially when the young crop passes through a dry period (Rai 1969). Poor
rainfall both in frequency and intensity during the spring crop is responsible for greater incidence
of S. frugiperda on this crop than on the autumn crop. Heavy rain kills this pest.
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Hibernation in the extreme south of the United States occurs as full-grown larvae. In succes-
sive generations the moths move northwards. As many as 6 generations are reported from the
southern States (Pathak 1975).
7.12 Common Cutworm, Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae)

Synonym: Prodenia litura (Fabricius)
Distribution: Australia, Bangladesh, Bhutan, Burma, China, India, Indonesia, Japan, Kam-
puchea, Korea, Laos, Malaysia, Nepal, Pakistan, Papua-New Guinea, Philippines,
Sri Lanka, Taiwan, Thailand, Vietnam
Host plants other than rice: Cabbage, castor, corn, jute, millets, sweet potato, tobacco and a wide
variety of other plants.
Spodoptera litura is a highly polyphagous pest infesting rice only sporadically. The young caterpil-
lars feed on leaf surfaces from the edge towards the midrib. As they grow, they become voracious.
Young rice plants are often cut at ground level while older plants are only defoliated. The pest is
generally a problem on upland rice because it needs dry soil for pupation. Lowland fields also
occasionally suffer damage when the larvae move from one field to another grazing over the young
rice plants.
The adult moth has dark purplish brown forewings with numerous spots and light colored lines.
The hind wings are whitish, narrowly banded along the outer margin. Moths are nocturnal in habit
and hide during the day at the base of rice plants and grassy weeds. Adults are strong flyers and can
migrate hundreds of kilometers. Female moths start oviposition 2 - 3 days after emergence.
Eggs are laid on leaves in clusters of 200 to 300 and they are covered with buff colored hairs.
The eggs are white, round and have a ridged surface. They hatch in 3-4 days.
Fig. 7.12
Newly-hatched caterpillars are small, and blackish green with a distinct black band on the first
abdominal segment. They feed gregariously for 3-5 days before they disperse. Larvae pass through
5 instars before they pupate. Mature larvae are stout, smooth, dull greyish in color, with a bright
yellow stripe down the back and along each side of the body. There are black crescent spots next
to the stripes. The head is black to dull brown with a yellow V-shaped marking. The larval period
is 20 - 26 days.
In upland conditions, pupation takes place in soil in individual earthen cells. In wetland fields,
larvae pupate in the rice plants or in grassy areas along field borders. The pupa in reddish brown
in color and the moth emerges in 6-7 days.
7.13 Rice Swarming Caterpillar, Spodoptera mauritia (Boisd.) (Lepidoptera: Noctuidae)

Distribution: Australia, Bangladesh, Bhutan, Burma, China, Fiji, Hawaii, India, Indonesia,
Japan, Kampuchea, Korea, Laos, Madagascar, Malaysia, Nepal, Pakistan, Papua-
New Guinea, Philippines, Sri Lanka, Taiwan, Tanzania, Thailand, Uganda, Viet-
nam
Host plants other than rice: Agropyron sp., Axonopus compressus (Sw.) Beauv., Brachiaria mutica
(Forssk.) Stapf, Fimbristylis acuminata Vahl., Isachne globosa (Thunb.) O. Kuntze,
Ischaemum indicum (Houtt.) Merr, Panicum repens L., Paspalum conjugatum
Berg., Saccharum sp., Zea mays L.
The rice swarming caterpillar is a sporadic pest occasionally causing up to 20% loss in rice yield. It
is also polyphagous and infests various graminaceous plants. Severe outbreaks of this pest have
been reported from Sarawak (Rothschild 1969) and India (Sathiyanandam et al. 1984). It occurs in
all rice environments but is least abundant in irrigated fields.
Fig. 7.13
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The caterpillar damages rice by feeding on the leaves at night. The damage by early instar
larvae is negligible and escapes notice. But the older caterpillars are voracious and can devastate a
whole field within a short time. They feed mainly on young rice plants in the nurseries or shortly
after planting. When a field has been stripped bare, they migrate in large numbers into adjoining
fields to continue feeding. Their migration is facilitated by the absence of standing water in the
fields. The plants are reduced to mere stumps which may die or even if they recover, do not bear
earheads uniformly. Nurseries situated near ill-drained marshy areas are attacked earlier than
those near dry ground. The infestation starts at the onset of the monsoon. Seedlings 4 to 20 days old
in wetbed nurseries as well as plants from direct seeded rice in poorly-drained fields suffer the most
serious attacks. Rice plants older than 6 to 7 weeks are usually not attacked.
The adult moth is stout with a body length of about 1.6 cm and wing expanse of 3.5 - 4.0 cm.
Forewings of the female moth are greyish brown with wavy lines and a dark spot subcentrally.
Wings of the male moth are more greyish. The hindwings are brownish white with thin black
margins. Moths are nocturnal in habit and hide in crevices in the soils or under vegetation during
the daytime.
Female moths generally mate on the night following emergence. Egg laying commences on the
third night and continues for a week. Peak oviposition is on the first night of egg-laying. Eggs are
laid on the underside of grass leaves and are covered with a thin layer of brownish grey felt.
Oviposition rarely occurs on rice plants and the first instar larvae usually migrate from grassy
vegetation to the rice field. But in the later stages of infestation, eggs may be laid on rice leaves. A
single female moth may lay 5 to 6 oblong egg masses each containing 150 to 200 spherical eggs. The
egg is cream colored when laid and acquires a brown tinge as development proceeds. The
incubation period ranges from 3 to 9 days.
Young larvae are light green with yellowish white lateral and dorsal stripes, later becoming
greyish brown, with a crescent shaped black spot on the side of each segment. The lateral stripes
have a reddish upper margin. The larvae are darker brown when populations are high.
First instar larvae prefer succulent young foliage for feeding. The weight of frass produced by
the last instar caterpillar is almost five times of that of the previous instar. This feeding pattern
explains the cause of sudden devastating damage by late instar armyworms in the field (Rothschild
1969).
The number of larval instars of S. mauritia has been reported from different countries; 4 in the
Philippines (Otanes and Sison 1952), 5-6 in India (Ananthanarayanan and Ramakrishna Ayyar
1937, Nair 1978), 7 in Sarawak (Rothschild 1969) and occasionally 8 in Hawaii (Tanada and
Beardsley 1958). The larval period is 15-24 days.
Pupation is in soil, within an earthen chamber. The pupa is dark brown in color and measures
16.5 mm long and 5 mm wide. The pupal stage lasts 7-14 days.
The adult moths are strong fliers and migrate long distances for oviposition. But there is little
evidence of migration in Sarawak (Rothschild 1969). It is suggested that the 1967-outbreak was due
to ecological factors. A prolonged period of dry weather followed by heavy rainfall led to the
abundant growth of young grasses which favored survival and development.
The pest infestation is governed by various climatic conditions. In south India, S. mauritia is
more prevalent in the ‘punja’ crop during October - December. Oviposition ceases by the end of
February when the weather becomes hot and dry. In Taiwan, 7 - 8 generations occur per year.
8. LEAF-FEEDING DIPTERA
8.1 Rice Leaf Miner, Hydrella griseola (Fallen) (Diptera: Ephydridae)

Distribution: China, Egypt, France, Germany, Italy, Japan, Korea, Malaysia, North Africa, USA
Host plants other than rice: Agropyron sp, Agrostis tenuis Sibth., Alisma plantago-aquatica L., Al-
lium cepa L., Avena sativa L., Bellis perennis L., Bromus inennis Leyss., Cynodon
dactylon (L.) Pers., Cyperus rotundus L., Dactylis glomerata L., Digitaria sanguinalis
(L.) Scop., Echinochloa colona (L.) Link, Festuca parvigluma Steud., Hordeum
vulgare L., Hydrocharis morsus-ranae L., Lamium album L., Lemna minor L.,
Lolium perenne L., Lychnis dioica L., Muhlenbergia mexicana (L.) Trin., Panicum
repens L., Paspalum scrobiculatum L., Phalaris arundinacea L., Phleum paniculatum
Huds., Phragmites australis (Cav.) Trin. ex Steud., Poa compressa L., Polygonum
lapathifolium L., Polypogon fugax Nees, Sagittaria latifolia Willd., Scirpus grossus L.,
Secale cereale L., Setaria glauca (L.) Beauv., Stellaria media (L.) Cyr., Siratiotes
aloides L., Trifolium pratense L., Triticum aestivum L., Typha latifolia L., Veronica
officinalis L., Zea mays L., Zizania aquatica L.
Fig. 8.1
DALE 453
The rice leaf miner is an early-season pest of rice in California and in Japan. Direct seeded rice is
more at risk than transplanted rice. The pest also attacks other crops such as barley, oats, timothy
and wheat.
The larvae mine the leaves and rarely leaf sheaths, and feed on the mesophyll tissues. The early
mines are usually linear which later widen and coalesce to form big white blotches. The leaves then
shrivel and lie on the surface of the water. Damage is more severe if the attack occurs at the seedling
stage. Deep water and cool temperatures during the early part of the growing season reduce plant
vigor (Kuwayama 1956, Grigarick 1963). These conditions cause the leaves to lie on the water
surface which results in increased oviposition and subsequent mining by the larvae. But if the
weather becomes warm and conditions become favorable for the growth of leaves in an upright
position, the infestation may be drastically reduced due to egg and larval mortality (Grigarick 1959).
Manandhar and Grigarick (1983) have shown that feeding effects on younger leaves are more
pronounced than on older leaves. The maggots are mobile and move on to new leaves after old ones
are completely mined.
The newly emerged adult is light grey, and looks like a small housefly. It walks briskly and stops
repeatedly to stroke and brush various parts of the body with its front and hind legs. Adults are
positively phototropic and are attracted to bright reflecting surfaces. Mating occurs 3 days after
emergence. Flies of both sexes are polygamous. Copulation is observed most often on the water
surface and floating leaves (Grigarick 1959). Adults are mobile and oviposit throughout the rice
field. They live for 3-4 months.
Leaves lying on or very close to the water surface are the preferred sites for oviposition. A
single fly may lay about 50-100 eggs singly on the upper surface of leaves. The average incubation
period is 5 days.
The newly hatched maggots are transparent to light cream in color. They feed using their
well-developed mouth hooks. Under conditions of limited food supply, crowding or submergence
of the infested leaves, the larvae may leave their first mines and migrate to newer leaves. The
maggots pass through 3 larval instars in a period of 7-10 days and pupate in larval mines.
The puparium is ovoid, tapering, and light to golden brown in color. It measures about 3.5 mm
long and 1.0 m wide. The pupae can easily be seen inside the transparent mines. Puparia are also
occasionally found on soil.
Kuwayama et al. (1955) reported the critical low temperature for the egg stage at 10°C, larval
stage at 6°C and pupal stage at 8°C. These temperatures are within 1.6 to 3.2°F of the threshold
temperatures recorded in California by Grigarick (1959). Total developmental time in California,
from hatched egg to adult varies from 13 days at 32°C to 94 days at 10°C. Larvae may be killed by
extremely high temperatures (44-46°C), especially when they mine upright leaves exposed to the sun
(Anon. 1983).
In Japan, high populations of adult flies are observed in the paddy fields from the middle of
June up to the end of July. But in the weedy fields, the adults begin to appear earlier (i.e. from April
onwards). The peak of fly population in Japan is reached in the months of May and July (Murai
1967). In California, the fly population is at its highest during March and April (Grigarick 1959).
H. griseola is a multivoltine species. All stages of the life cycle can generally be found after the
first generation of the year. It is possible to have 10 generations in a year under California
conditions.
8.2 Whorl Maggot, Hydrellia philippina Ferino (Diptera : Ephydridae)

Distribution: Bangladesh, India, Indonesia, Laos, Philippines, Taiwan, Thailand
Host plants other than rice: Cynodon dactylon (L.) Pers., Echinochloa colona (L.) Link,
Echinochloa crus-galli (L.) Beauv., Eleusine indica (L.) Gaertn., Fimbristylis
miliacea (L.) Vahl., Paspalum scrobiculatum L.
The insect was first reported as a pest of rice in Thailand (Patanakamjorn 1964). Ferino, then in
1968 studied the taxonomy and bionomics of the whorl maggot in the Philippines. At the Interna-
tional Rice Research Institute, the occurrence of the pest has been recorded throughout the year.
There are reports from many rice growing countries in Asia that H. philippina is becoming a serious
pest. This species is a pest of rice seedlings only under irrigated or rainfed conditions. Rice plants
grown with continuous standing water the first 3 or 4 weeks after transplanting have more insect
damage than plants in fields where the soil is water-saturated, but without standing water. The
gravid flies prefer flooded fields for oviposition (Viajante and Heinrichs 1985a). Ferino (1968)
reported an estimated yield loss of 1.4 tons per hectare due to the damage of this pest in the
Philippines. In south India, Thomas et al. (1971) reported that the whorl maggot could cause 20 to
30% yield loss on the first crop during April to September. But the infestation was less in the second
crop. However, Nurullah (1979) reported that whorl maggot leads to no adverse effect on tiller
production and it even increases the number of productive tillers to compensate for possible grain
Fig. 8.2
DALE 455
losses. Viajante and Heinrichs (1986) conducted six yield loss experiments and found that the whorl
maggot did not adversely affect yield of IR36 but did delay maturity. The experiments were
conducted by comparing yields of artificially infested and non-infested caged plots. The conflicting
reports regarding yield losses are likely due to different experimental conditions. When yield losses
are reported, insecticides that cause plant growth stimulation are sometimes used as an insecticidal
check to eliminate whorl maggots for comparison with infested plots. Also, cages are not used and
infested plots are a combination of the whorl maggot and several other pests at low levels and the
combined effect may be sufficient to cause yield loss. Another possible reason for reported yield
losses in some studies may be the inability of plants to fully recover from whorl maggot damage when
simultaneously stressed by soil mineral deficiencies or toxicities. Further studies should be con-
ducted to determine the importance of whorl maggot damage.
The maggots feed on unopened leaves, nibbling the inner margins. Conspicuous linear feeding
lesions are visible when the central leaf opens. Damaged leaves become distorted and may break
off in the wind (Ferino 1968). Infested plants are stunted. However, the pest can cause damage to
the boot leaf and developing panicles (Sain et al. 1983) which can lead to only partial filling of the
grains (Varadharajan et al. 1977). Small punctures appear in the middle of the flag leaf and its
margins get discolored (Basu 1979). The chlorotic effect coupled with disrupted sugar metabolism
and poor nutrient uptake are probably the reasons for the manifested effects on infested plants
(Ramamurthy et al. 1977).
The peak time of adult emergence is from 7 to 10 a.m. Adults are dull grey flies. Females are
about 1.8 to 2.3 mm long but the males are slightly smaller. The flies have a definite preference for
high moisture conditions and are not found in upland rice fields. They are often found on floating
foliage in calm water and prostrate vegetation near slow-flowing drainage and irrigation canals
(Karim 1969). The flies move by a combination of walking and hopping in a zigzag pattern. Adults
are saprophagous on dead aquatic insects. Mating occurs mostly during morning and evening hours
from the second day after emergence. Adults locate rice fields by reflected sunlight from the water
surface. Once the canopy closes they can no longer find the rice crop, thus direct-seeded fields or
seedbeds are not highly attractive to the adults.
The white elongate, cigar-shaped eggs are laid singly on either surface of the leaves. Females
lay about 100 eggs during the lifetime of 3-7 days. Flies prefer the basal portions of the leaf to the
upper half. The maggots emerge after 2-6 days. The larvae move down the leaf into the whorl on a
film of dew and feed within developing leaf whorls. The larva is transparent to very light cream
during the first instar but later becomes yellow. Larvae mostly remain outside the leaves, and feed
on the mesophyll tissues of the foliage. When the leaves emerge from the whorl, damage can be
seen as pinholes in the leaves and white and yellowish lesions at the leaf edges. Severely damaged
leaves break in the wind. The larva undergoes 3 instars; the larval period ranges from 8 to 17 days.
Pupation takes place between the leaf sheaths where the pupa is loosely attached to the stem. The
puparium is light to dark brown, ovoid and subcylindrical in shape. The pupal period is 5-9 days.
The fly population in the Philippines is at its lowest during the first six months of the year which
also coincides with times of high temperature and low rainfall. The fly population begins increasing
in July and continues to increase up to November after which it declines. But in India, the autumn
rice (April to mid-July) is reported to be most affected by whorl maggots, with a peak infestation in
the first week of June (Sasidharan et al. 1979).
The optimum temperature for normal development is between 29 and 33°C, at which the life
cycle is about 19 days (Karim 1969). Adult flies have a special preference for high moisture areas.
Under natural conditions, environments such as ponds, streams, lakes and irrigated rice fields
provide a favorable ecological niche for the breeding of the flies.
Close planting decreases oviposition and subsequent damage by the whorl maggots (Viajante
and Heinrichs, 1985b). Similarly, an azolla cover on the surface of rice fields reduces the egg
population and consequently leaf damage by the larvae.
Hydrellia philippina is a multivoltine species with overlapping generations under field condi-
tions. Temperature, humidity and availability of host plants influence the number of generations.
It is believed that the whorl maggot has 13-15 generations in a year.
8.3 Paddy Stem Maggot, Hydrellia sasakii Yausa et Isitani (Diptera: Ephydridae)
Distribution: Japan
Host plants other than rice: Leersia japonica Honda, Leersia oryzoides (L.) Sw. var japonica
Hack., Leptochloa chinensis (L.) Nees
The pest is particularly important on the late-planted rice crop in Japan. It is most damaging to
young rice plants at one month after transplanting. The insect closely resembles Chlorops oryzae in
its biology.
Newly hatched maggots feed on the unopened tender leaves which are marked with small spots
and stripes due to the insect attack. The older larvae enter the central whorl to feed on the inner
DALE 457
margin of unopened leaves causing large marginal spots on the leaves. Larvae occasionally infect
the panicles in the booting stage and damage the developing grains. A heavy attack may stunt the
crop and reduce yield. Heavy tillering varieties or plants transplanted with several seedlings per hill
are less damaged by the pest. The percentage yield reduction is 0.3 of the percentage of leaves
having large spots on the marginal part of the foliage.
The fly is blackish grey tinged with bronze. The halteres are vividly yellow and their bases are
orange in color. The females are usually larger and with a more swollen abdomen than the males.
The flies are often seen resting on the leaf tips of rice plants. Rice planted with wide spacing is
preferred for oviposition.
Eggs are laid singly on both sides of the leaf and hatch in about 2 days. The larvae are milky
yellow and 4.0-4.5 mm long. The larval stage lasts for 2-3 weeks. Pupation occurs between the leaf
sheath and stem. Fresh pupae are light brown and gradually turn to dark brown as they mature. The
average pupation period during summer is 5-8 days but it is about 17 days during spring.
During winter the insect hibernates as larvae on graminaceous weed plants. The maggots are
fairly tolerant to fluctuations in temperature of from 18 to 39°C (Okamoto and Koshihara 1962).
The insect has 5 overlapping generations in a year. The population is high during July to early
September. This period corresponds to the third and fourth generations of the pest.
9. LEAF-FEEDING COLEOPTERA
9.1 Rice Hispa, Dicladispa armigera (Oliver) (Coleoptera: Chrysomelidae)

Synonyms: Hispa armigera Oliver
Hispa aenescens
Distribution: Bangladesh, Bhutan, Burma, China, India, Indonesia, Kampuchea, Laos, Malaysia,
Nepal, Pakistan, Papua-New Guinea, Sri Lanka, Thailand, Vietnam
Host plants other than rice: Cynodon dactylon (L.) Pers., Echinochloa colona (L.) Link,
Echinochloa crus-galli (L.) Beauv., Panicum repens, L., Paspalum paspalodes
(Michx.) Scribn., Vetiveria zizanioides (L.) Nash, Zizania aquatica L.
The pest causes heavy crop losses in many Asian countries particularly in Bangladesh. It is most
common in wetland environments. Sporadic outbreaks have been reported from many parts of
India (Budhraja et al. 1980, Das 1980, Chand and Tomar 1984). Infestations of hispa have increased
in recent years. Introduction of high yielding varieties to more areas and improved agronomic
practices seem to be partly responsible for the increased infestation (Dhaliwal et al. 1980). The
extent of damage in Bangladesh reaches 65% (Alam 1967).
Both grubs and adult beetles feed on rice plants. Grubs mine the leaves by feeding on the
mesophyll between the veins and tunnelling the tissue in the direction of the main axis of the leaf
advancing towards the leaf sheath (Acharya 1967). Adults remove chlorophyll first between the
Fig. 9.1
veins of the lamina giving the appearance of white parallel streaks on the leaves. Later due to
feeding on the veins, white irregular blotches appear on the leaves. In severe cases, leaves are
brown and the field presents a dried up appearance. Even replanting may not be of much avail as
the pests persist in the field and infest the freshly planted rice seedlings.
Adult beetles emerge in early morning and rest on the lower parts of the plant during day time.
They are small, shiny black beetles about 5.5 mm long, with spines on their wing covers. The feeding
activity of beetles is maximum during morning hours. They begin feeding mostly from the apical
parts of leaves and proceed downward. The adult beetles prefer to feed on tender leaf tissues
mostly from the dorsal side of the leaf. A single adult beetle consumes about 25 mm2 of leaf area
per day (Budhraja et al. 1979). Females live for 20 days while the males’ longevity is only a fortnight.
The beetles mate 3-4 days after emergence. Eggs are laid singly within the epidermal layers of the
ventral surface of rice leaves. A single female lays on an average about 55 eggs. The minute eggs
are usually found towards the leaf tips. The incubation period ranges from 4 to 5 days.
The newly hatched larvae are pale yellow, dorso-ventrally flattened and about 2.4 mm long. They
start mining from the leaf tip towards the base of the leaf blade. Larval movement inside the leaf is easily
seen when it is placed against light. A single grub was found to consume, on an average 123.4 mm2 of
rice leaf area in its lifetime (Budhraja et al. 1979). The larval stage lasts for 7-12 days.
The pupae are flat, brown and exarate. The pupal stage is completed within the leaf mines and
takes 4-5 days. The adult beetles cut their way out of the rice leaf and become external feeders.
The number of generations of the pest seems to depend on the nature and number of crops
grown. In regions such as Bangladesh where three rice crops are grown in a year, 6 generations of
hispa occur. The sequence is: one generation during February in the winter rice (boro), one during
April-May in graminaceous weeds, one on upland rainfed rice (aus), and three others in the
monsoon crop (aman) from July to November in succession (Sen and Chakravorty 1970).
DALE 459
Adult beetles appear in rice fields in February and the population gradually increases until
June-July when the larvae as well as adults cause heavy damage to young rice plants. The popula-
tion declines after August. Adult beetles in small numbers can still be collected from rice fields up
to September-October (Pathak 1975).
Many factors have been reported to influence the incidence of hispa in rice fields. Prakasa Rao
et al. (1971) reported that top dressing with high levels of nitrogen during periods of pest abundance
resulted in greater susceptibility of the rice crop to the pest. Dhaliwal et al. (1980) found hispa
incidence to increase with an increase in nitrogen levels from 0 to 100 kg/ha. However, at 150 kg,
the infestation considerably decreased.
Adult beetles prefer to feed and oviposit on rice crops in the vegetative stage (i.e. plants from
25 to 60 days old). After this phase, the leaves become silicified and less succulent perhaps offering
mechanical resistance to feeding (Prakasa Rao et al. 1971). Data of studies conducted by Basu and
Banerjee (1957) showed that plants just after transplanting were more prone to damage than those
of other stages.
Date of plating influences the ultimate damage caused by hispa. It was shown by Prakasa Rao
et al. (1971) that though the July plantings suffered considerable infestation by the two early broods,
they soon recovered and reached a stage of least susceptibility for the two succeeding broods. But
plantings in August suffered severe damage with less chance of recovery.
Weather conditions prevailing in regions of infestation in south India indicate that high
humidity after rains and intermittent bright sunshine seem to favor hispa development. In north
eastern India, rainfall was reported to have a negative effect on the activity of hispa (Ghosh et al.
1960). Heavy rainfall in July followed by unusually low rain in August and September was charac-
teristic of epidemic years.
Dicladispa gestroi is a pest of rice in the coastal regions of Malagasy Republic. It is as important
as stem borers in this area. The nature of damage is similar to that of D. armigera.
Another species, Dicladispa vindicyanea, occurs in Burundi, Kenya and Zaire. The insect is
suspected to be a vector of yellow mottle virus in Kenya. The adult beetle is metallic blue-green,
about 5 mm long with five lateral spines on each side of the thorax. The female lays eggs on the
underside of young leaves and covers them with excreta. The grubs mine the leaves making
transparent patches and adults eat leaf tips. There are 3-6 generations in a year; the life cycle takes
24-40 days to complete.
9.2 African Rice Hispa, Trichispa sericea (Guerin) (Coleoptera: Chrysomelidae)

Distribution: Angola, Burundi, Cameroon, Ethiopia, Ivory Coast, Kenya, Madagascar, Mali,
Rwanda, Senegal, South Africa, Sudan, Swaziland, Tanzania, Togo, Uganda, Zaire
Host plants other than rice: Chloris virgata Sw., Echinochloa holubii (Stapf) Stapf, Eragrostis
aethiopica Chiov., Eragrostis heteromera Stapf, Digitaria zeyheri (Nees Hend.,
Diplachne fusca (L.) Beauv. ex Stapf
Fig. 9.2
Trichispa sericea is only recorded from Africa. It is an important pest in rice nurseries and only
sporadically serious on transplanted rice. It is reported to transmit rice yellow mottle virus in
Kenya.
Both adults and grubs feed on leaf tissues of young rice. The first attack is highly localized but
spreads rapidly. Attacked plants have irregular pale brown patches and narrow white streaks on
the leaves; the pale brown patches are the larval mines, and the whitish streaks are the feeding scars
by the adult beetles. Severe attacks kill the plant.
The adult is a dark grey beetle covered with spines. It is about 3-4 mm long. Adult females live
for about a fortnight and may lay more than 100 eggs during this period.
White, boat-shaped eggs (1 mm long) are laid singly in slits in the leaf made by the adult beetle,
the wound being covered by a spot of excreta. Hatching takes place after 3-4 days.
The grub is slender and yellowish which when fully grown is about 6 mm long. The larvae mine
inside the leaf, the damaged portions being visible externally as a pale brown blotch. The larval
period lasts about 10 days.
Pupation takes place within those portions of the leaf lamina that are not submerged. The
pupal period generally lasts about 6 days. Emerging adults migrate to alternate host plants.
9.3 Rice Blue Beetle, Leptispa pygmaea Baly (Coleoptera: Chrysomelidae)
Distribution: India
Host plants other than rice: Arundinella metzii Hochst., Ischaemum travancorense Stapf,
Paspalum scrobiculatum L., Pennisetum purpureotyphoides (Schum.), Saccharum
ofjicinarum L., Verivena zizanioides (L.) Nash
DALE 461
Fig. 9.3
Leptispa pygmaea, a minor pest of rice, assumed serious proportions in poorly drained rice fields of
central India in 1978. The pest appeared in an epidemic form for the first time during the second
crop (July to November) in Maharashtra.
The adult is a small bluish elongated beetle measuring about 5 mm in length and 1.5 mm in
width. Both adults and grubs feed on the surface tissues of rice leaves. Damage by the adult is
similar to that of the hispa but the feeding lesions are narrower and more regular in shape. The
attacked leaves become wafer thin, and dry up. This pest usually appears along with rice hispa
aggravating the damage done by it to the rice crop.
The pest survives the off-season (November-February in the adult stage on grasses, volunteer
rice plants, ratoon rice or on sugarcane. The activity of the pest is at its lowest during this period.
No egg laying has been observed on the alternate hosts during the pest’s inactive stage (Khanvikar
et al. 1983).
9.4 Rice Leaf Beetle, Oulema oryzae (Kuwayama) (Coleoptera: Chrysomelidae)

Synonyms: Lema oryzae Kuwayama
Lema suvorovi Jacobson
Distribution: China, Japan, Korea, Ryuku Islands, eastern Siberia, Taiwan
Host plants other than rice: Dactylis glomerata L., Glyceria tonglensis C. B. Clarke, Leersia
oryzoides (L.) Sw., Phragmites australis (Cav.) Trin. ex Steud., Zizania latifolia
(Griseb.) Turez. ex Stapf
Oulema oryzae occurs in upland and wetland environments and is one of the serious pests of rice in
northern Japan, Korea and China leading to crop losses of up to 30%. The percentage of yield
reduction in Japan is computed as 0.25 of the percentage of injured leaves. In Taiwan, it infests the
first crop in the mountain regions.
Both adults and larvae feed on the leaf surface causing a scorched appearance of the foliage.
Larvae skeletonize leaf blades in a linear fashion. In short duration rice varieties, plants often do not
recover from O. oryzae damage. Retardation of plant growth and decrease in the number of leaves
and tillers have been observed (Kojima and Emura 1979). Ripening of grains in infested plants is
considerably delayed leading to reduction in both quality and quantity of grain yield. In cases of
severe infestation, the plants die and the fields present an appearance of having been burned by fire.
In Japan, damage varies from 5% to 10% when slight and 20% to 30% when severe. The tolerable
injury level is set at 20% injured leaves in late June (Koyama 1978).
Adult beetles are metallic blue with the head and antennae black and the thorax yellowish
brown. The females deposit cylindrical eggs in masses on the upper surface of leaves. The
oviposition period lasts 15 days and incubation varies from 5 to 11 days. The life of the adult extends
for one year or more.
The brown larvae are globular in shape and rather heavily sclerotized and with dark brown
nodules on a yellow base. They cover their body with dark greenish excreta and appear as mud on
leaves. The larval period is 13-19 days. The full-grown larva pupates within an ellipsoidal whitish
cocoon mostly on the rice leaves, though in upland fields, pupation sometimes takes place on or
under the ground.
There is one generation per year and overwintering occurs in the adult stage. In Japan, the
increase in the number of leaf beetles often coincides with the transplanting of rice seedlings. The
beetles leave their hibernating sites by late May, mate and start laying eggs by early June. Oviposi-
tion continues until nearly the end of July. Then the adults bury themselves in the debris of mountain
bush and under the roots of grasses.
O. oryzae is highly adaptable to low temperatures. In the north temperate regions of Japan,
female beetles stop depositing eggs early and adult longevity is shortened if the atmospheric
temperature rises early in summer. On the other hand, when cooler days continue to prevail in
summer, the hibernating adults survive for a longer time in the field and oviposition continues
longer (Kuwayama 1966).
10. LEAF-, STEM-, AND PANICLE-FEEDING APHIDS

Many aphid species (Homoptera: Aphididae) infest the aerial parts of rice even though rice is not
the primary host plant for most of them. A list of aphids reported on rice with their geographical
distribution is provided in Table 6.
The rusty plum aphid Hysteroneura setariae feeds on the leaves and unripened grains of rice.
Moderately infested grains show brown necrotic spots. A heavy infestation turns all spikelets brown
and chaffy. Severe attacks by this aphid have been reported recently from countries such as Sierra
Leone (Akibo-Betts and Raymundo 1978), India (Garg and Sethi 1978) and Nigeria (Akinlosotu
1980).
The wingless adults of H. setariae are 3-4 mm long and are rusty to dark brown in color.
Nymphs are pink. The average nymphal period is 10 days. In Africa, the aphid appears in large
numbers with the May rains and remains active on the rice crop till December. But heavy rains
greatly reduce its population during August-October. In India, this aphid is found on rice
throughout the season.
DALE 463
TABLE 6
Distribution of aphid species feeding on the leaf. stem, and panicles of rice (Yano et al. 1983)
Aphid species Host plants Distribution
Aphis craccivora Koch Rice, many plants Many countries
Aphis gossypii Glover Rice, many plants Completely cosmopolitan, absent only from
the colder parts of Asia and Canada
Brachysiphoniella montana Rice India
(van der Goot)
Diuraphis noocia Rice, grasses North Africa
(Mordivilko ex Kurdjumov)
Hysteroneura setariae (Thomas) Rice India, Philippines, Thailand, West Africa
Melanaphis sacchari Rice, sugarcane, sorghum, Japan, Philippines
(Zehntner) maize, wheat, grasses
Metopolophium dirhodum Rice, grasses Central Asia
Myzus persicae (Sulzer) Rice, many crop plants, weeds Virtually cosmopolitan
Rhopalosiphum maidis (Fitch) Rice, many crop plants, some Throughout the tropics, subtropics and the
weeds warmer temperate regions
R. nymphaeae (L.) Rice, Monochoria vaginalis, Bangladesh, Fiji, North America, Philippines,
Saginaria, Trifolia, Prunus spp. Spain, Taiwan
R. padi (L.) Rice, many plants belonging to Africa, America, Asia, Australia, Europe
Gramineae and Cyperaceae
Rhopalosiphum sp. Rice Japan
Schizaphis graminum (Ron- Rice, wheat, barley, oats, Italian Africa, Asia, South America, USA
dani) millet
Sipha glyceriae (Kaltenbach) Rice, grasses Italy
Sitobion akebiae (Shinji) Rice, wheat, wild weeds Japan, Korea
S. avenue (F.) Rice, grasses Europe, North America, Taiwan
S. fragariae (Walker) Rice Spain
S. graminis (Takahashi) Rice, grasses Oriental and Ethiopian regions
S. miscanthi (Takahashi) Rice, graminaceous plants Australia, China, New Zealand, Philippines,
South Pacific regions, Taiwan.
Rhopalosiphum padi is variously known as the bird cherry oat aphid and the apple grain aphid.
It is a vector of the virus disease ‘giallume’ (yellow disease or rice yellows) of rice in Italy. The
primary host of this aphid species in Italy is the fruit tree Prunus padus. When P. padus is not
available, the aphid overwinters on secondary hosts such as rice and various grasses.
11. RICE STINK BUGS

The characteristic foul odor produced by scent glands on the abdomen of these insects has led them
to be commonly called ‘stink bugs’. About 60 stink bug species have been reported to infest rice
fields and at least half of them are considered pests causing yield loss or pecky rice.
‘Pecky rice’ is referred to as the condition of the grain after being sucked by stink bugs and
partially or wholly stained by bacteria or fungi. If a sample of rice contains some pecky grains, the
whole lot is considered contaminated and the market price is decreased in some countries. An
emphasis on rice has shifted from yield to quality. In Japan, the inspection for rice grain quality has
become stricter and the pecky rice problem has assumed increasing importance. The major bugs
causing pecky rice in Japan are listed in Table 7.
TABLE 7
Bugs that cause pecky rice in Japan (Kisimoto 1983)
Family Species
Alydidae Leptocorisa chinensis (Dallas)
Coreidae Cletus punctiger Dallas
Lygaeidae Nysius plebejus Distant
Togo hemipterus Scott
Miridae Stenodema sibiricum Bergroth
Trigonotylus coelestialium Kirkaldy
Pentatomidae Dolycoris baccanum L.
Eysarcoris lewisi Distant
E. parvus Uhler
E. ventralis Westwood
Lagynotomus elongatus Dallas
Nezara antennata Scott
N. viridula (L.)
Scotinophara lurida Burmeister
Ropalidae Aeschynteles maculatus Fieber
Stink bugs of the genera Leptocorisa, Nezara, and Oebalus are separately dealt with.
11.1 Leptocorisa spp. (Hemiptera: Alydidae)

Host plants other than rice: More than 65 plant species (Srivastava and Saxena 1967)
The rice bugs are major pests of rice wherever the crop is grown. The most important species in the
tropical and subtropical rice regions belong to the genus Leptocorisa. Another genus, Stenocoris
which was earlier a subgenus under Leptocorisa, also contains a few species which are very often
recorded from rice fields. Many species share a common distribution but very often their relative
abundance differs. For example, in the Philippines, L. oratorius is more abundant than L. acuta
(Fig.ll.1).
Rice bugs are found in all rice environments, but are more prevalent in rainfed wetland or
upland rice. Factors that favor high bug populations are adjacent woodlands, extensive weedy areas
near rice fields and staggered rice planting (Reissig et al. 1985).
The intensity and type of damage caused by rice bugs depends on the stage of the rice crop,
population density of the pest and ecological conditions. Both nymphs and adults are destructive
to the crop, even though the damage by nymphs is more severe (Pathak 1975). Nymphs prefer
grains at the milky stage for feeding. They feed by the insertion of the proboscis at points where the
glumes meet. During the process, the bug secretes a proteinaceous stylet sheath to form a feeding
canal for its sucking mouthparts (Gyawali 1981). Removal of stored assimilates from the developing
grains may result in either unfilled or partially filled grains with damage symptoms. Uichanco
DALE 465
Fig. 11.1
(1921) considered that the bugs inhibited kernel development either through mechanical injury at
feeding or injection of some toxin in addition to the removal of milky sap. Panicles in heavily
infested fields remain erect.
The quality of grains is also reduced by the insect attack. Pecky rice in Japan and the United
States is down graded and priced low. The smell of the infested grains lowers market value.
Damaged grains even after cooking retain the buggy smell. Sometimes even the straw has an
off-flavor which is unattractive to cattle.
In India, on an average, 5-10% of the grains are damaged due to infestations of the rice bug. In
years of severe attacks, damage reaches 40-60% (Kalode 1982). In the case of early paddy, maturing
in September, the loss is usually greater than in the late maturing varieties. Leptocorisa spp. cause
significant damage in Thailand every year. In the Malay Peninsula, the yield loss in certain areas
exceeds 25%.
The adult bug is long and slender, about 16-19 mm long. Rice bugs are diurnal but are most
active during early morning and evening. During the hotter parts of the day, the bugs are seen
taking shelter under the leaves of rice plants or weeds near rice fields. Rice bugs fly short distances
when disturbed. The females are stronger fliers than the males. The males are capable of mating
shortly after emergence but the females start only one or two weeks after becoming adults. Mating
generally takes place in the morning hours and rarely at night. Oviposition starts 3-4 days after
mating. The female lays 250-300 eggs in two or three straight rows along the midrib on the upper
surface of the leaf blade. Eggs are laid during the night. They are disc-shaped, the convex lower
surface being attached to the leaf by means of a white gummy substance. Eggs are dorsally flat and
elliptical, with the surface slightly granulate and shining. The incubation period is about a week.
Females feeding on rice lay more eggs than those feeding on grasses.
Rice bugs become active when monsoon rains start. They complete 1-2 generations on grassy
weeds before migrating to rice fields. A small number of adults usually appear in the rice fields as
soon as the plants start flowering. When about 80% of the grains have ripened, the adults migrate
to fields having rice plants of less maturity. Akbar (1958) noted that before flowering, the bugs feed
on the seedlings by sucking sap from leaves and tender shoots.
Rice bugs are often seen on flowering wild grasses near canals. In Papua-New Guinea,
numbers of Leptocorisa were observed (Sands 1977) to increase enormously when weeds in the
adjacent paddies were left unmanaged. The bugs are attracted to the rice crop when the plants start
flowering. Kainoh et al. (1980) have shown that adult bugs of L. chinensis can detect the odor of rice
plants when they are close to the rice field. But their aggregation on the panicles seems to be due
to the arresting effect of the flowers. The adults have been observed to live for up to 4 months.
The freshly hatched nymphs are green but turn brownish as they grow. They congregate around
the empty egg cases for some time before they disperse for feeding. Nymphs feed voraciously and
gregariously until the fourth instar. Feeding on the flowering heads continues throughout the day
especially in the early morning and at dusk when the sunlight is not intense. The nymphs pass
through 5 instars in a period of 25-30 days to become adults. The final molting takes place at night.
Nymphs stop feeding about 5 hours prior to selecting a place to molt on the leaves or flowers of the
host plant. The nymph then remains motionless in an extended position for about 2 hours before
ecdysis. The actual process of molting takes only 2 minutes.
The bug reacts favourably to high humidity and frequent drizzles prevailing from April to June,
the active season of the bugs in India (Singh and Chandra 1967). Srivastava and Saxena (1967)
reported that it was the light intermittent rain which favoured the build-up of bug populations while
heavy rains had an adverse effect on the pest. In the Philippines, rice bug activity was at its peak
during the rainy period in November-December, and dry season crops which attained the milky
stage in March-May were less severely attacked (Uichanco 1921).
In Papua-New Guinea, rice bug abundance depends on the availability of suitable host plants
rather than on seasonal weather fluctuations, except for periods when adult aestivation occurs
(Sands 1977).
In India, the bugs start laying eggs by the last week of June when the monsoon generally breaks.
In early July, many nymphs and adults are seen on rice nurseries and on grasses around fields
planted to rice or millet during the previous season. Maximum population of the pest occurs during
September. The population declines from the end of October and in December, only a few adult
bugs are seen on grasses growing in shady areas. Four overlapping generations have been recorded
in a year. In the Philippines, population build-up is greatest in September-October after which the
adult bugs undergo aestivation or diapause in the grasses (Pathak 1968).
In Japan, L. chinensis hibernates on ferns and weeds under humid and shady conditions (Ito
1978). In India, teeming populations of L. acuta have been found to hibernate on trees during the
relatively colder and drier months of December to March (Banerjee and Chatterjee 1965).
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11.2 Nezara viridula (L.) (Hemiptera: Pentatomidae)

Distribution: All the rice growing regions of the world
Host plants other than rice: Many, especially legumes, cotton, tomato.
Nezara viridula, the southern green stink bug, is a polyphagous pest. It occasionally causes serious
losses to rice. Nymphs and adults suck sap from stems and developing grains. At harvest, grains
may be shrivelled or speckled. An adult N. viridula can produce on an average 1.5 pecky rice grains
per day (Kisimoto 1983).
Fig. 11.2
The pest has assumed great economic significance in Japan since 1956 when the introduction
of early-transplanting resulted in the flowering of the rice crop at the same time as the emergence
of first generation adults. Moreover, at this time of the year, host plants other than rice are relatively
scarce. In Indonesia, N. viridula is considered as an important pest of rice. But in Malaysia,
although localized damage may be serious and spread over several hectares of rice fields, the overall
crop loss is less than 1% (Van Vreden and Ahmadzabidi 1985).
The adult bug is 12-18 mm long and green in color except for the membrane of the forewings
which is hyaline. Four distinct color morphs have been identified. Females start mating one week
after emergence and the preoviposition period varies from 2 to 3 weeks. The bugs are attracted to
light and heading rice. They are active fliers and are distributed rather randomly in the paddy field.
Egg are laid in parallel rows on the lower surface of leaves in masses of 70-130. Each female
usually lays 2-8 egg masses. Eggs are yellow but turn red just before hatching.
The newly-hatched nymphs congregate for some time around the empty egg shells before
they disperse for feeding. After the fourth instar, nymphs feed solitarily. The nymphs undergo
5 molts in 35-45 days to become adults. The nymphal population is usually heterogenous in
color patterns.
Rice fields surrounded by wild vegetation seem to be especially vulnerable to pest infestation.
Mass migrations of the bug from vegetable fields to the rice crop have also been observed. Adults
of N. viridula can fly over a distance of 1 km even with a hill in their flight path (Kiritani and Hokyo
1970).
In southern Japan, the insect has three overlapping generations with a partial fourth. In years
that are warm in March-June, the hibernating bugs become active earlier and complete the fourth
generation. Adults hibernate in shaded dry places. After hibernation the emerging adults feed on
grasses, orchards, and other spring and summer crops before they migrate to rice.
Among the external conditions influencing mortality of the green stink bug in the overwintering
season, weather conditions are the most important. In severe winters, mortality as high as 97.5%
has been reported in Japan (Kiritani and Hobo 1970). Intrinsic factors such as sex, body size and
color type of the adult influence the bug mortality in the overwintering period.
11.3 Oebalus spp. (Hemiptera: Pentatomidae)

Distribution: O. pugnax (F.): Cuba, Dominican Republic, USA
O. ornata (Sailer): Brazil, Dominican Republic, Haiti, Puerto Rico
O. poecila (Dallas): South American rice growing regions
Host plants other than rice: Avena sativa L., Cyperus iria L., Digitaria sanguinalis (L). Scop.,
Echinochloa crus-galli (L.) Beauv., Hordeum vulgare L., Panicum dichotomiforum
Michx., Paspalum urvillei Steud., Secale cereale L., Sorghum bicolor (L.) Moench.,
Triticum aestivum L., Zea mays L.
Several species of Oebalus cause damage to rice kernels in the Americas. Among them O. pugnax
is the most important and the one that has been studied in detail. The bionomics of other species
are similar to those of O. pugnax (Fig. 11.3).
Fig. 11.3
DALE 469
O. pugnax is commonly found in North America, east of the Rocky Mountains, and as far north
as Minnesota and New York (Sailer 1944). The insect is a pest of rice in the southern rice producing
areas.
Nymphs and adults suck rice grains at the milky stage. Florets fed upon during the early
endosperm formation are often wholly or largely drained of their content, which may result in either
empty or severely atrophied grains. Feeding is accompanied by subsequent formation of an area of
chalky discoloration surrounding the feeding site. This injury is evident on the mature hulled kernel
also. 'Pecky' grains are structurally weak at the region of insect injury and often break during
milling. This results in a reduction in market value. Pecky rice which escapes breakage during
milling is of inferior quality as a finished product owing to its discoloration.
Economic losses due to stink bugs vary. O. ornata causes crop losses as high as 50% in the
Dominican Republic. Early estimates in the USA put the loss caused by O. pugnax from a negligible
amount to 25% (Ingram 1927). Consistent and severe losses in rice yields were reported for
infestation levels of 7-8 bugs per 1,000 panicles (Swanson and Newsom 1962).
The adult bug is a straw colored pentatomid, approximately 1.2 cm long, with prominent lateral
pronotal spines. The antenna is pale red, with the first segment lighter in color than the others. The
legs are yellow with scattered black punctures. The hind wings are iridescent and frequently appear
green. The bugs give off a strong, disagreeable odor when disturbed.
The premating period for both males and females averages 2.4 days (Nilakhe 1976). The barrel
shaped eggs are laid in masses numbering 10-47. They are arranged in two rows on stems, leaves or
panicles. Eggs are bright green when laid but turn red before they hatch. The incubation period is
4-8 days. Females reared on barnyard grass and vasey grass laid significantly fewer eggs than those
reared on rice (Nilakhe 1976).
The newly hatched nymphs congregate on and around the empty egg shells for about a day
before they disperse to feed. When they hatch, they nymphs have a black thorax, head, legs and
antennae and two black spots on the red abdomen. But this coloration fades away with successive
nymphal molts. There are five nymphal instars and the nymphal period lasts from 16 to 20 days.
Bugs start hibernating during the first week of October. Wood trash and bunch grass are the
major hibernation sites (Odglen and Warren 1962). The insects start emerging out of hibernation
by the first week of April and complete their emergence by the last week. Males are the first to
come, about 10 clays before the females. Adult bugs are seen voraciously feeding on the flowers of
wild grasses. They reproduce on favourable host plants in or adjacent to rice fields or pasture lands
until rice begins to flower. Factors responsible for initiating movement of the adults from grasses
to rice are not fully understood. Douglas (1939) reported that the stink bugs forsake the grasses as
the rice panicles emerge. He also noted that when an early field of rice is harvested, the bug
population in the adjoining rice fields increases. Bowling (1967) suggested that the movement to
rice is initiated when other hosts become unfavourable as food.
Oebalus ornata is a serious pest of rice in the Dominican Republic. The bugs are very active on
the panicles during morning hours, but migrate to the base of plants when the sun becomes hotter.
On cloudy or light rainy days, the nymphs and adults remain and feed on the panicles all day. The
insect is active in the Dominican Republic throughout the year and has seven generations. The
population is usually at its peak during July, which coincides with the flowering of the first rice crop
and is comparatively lower in February when the second crop flowers. In May, the population is at
its lowest.
12. PANICLE THRIPS
12.1 Haplothrips ganglbaueri Schmutz (Thysanoptera: Phlaeothripidae)

Synonym: Haplothrips aculeatus Karny
Distribution: India, Malaysia, Pakistan, Sri Lanka
Host plants other than rice: Ageratum conyzoides L., Asytasia gangetica (L.) Aners., Blumia
wightiana DC, Bothriochloa pertusa (L.) A. Camms, Celosia argentia L., Chloris
barbata Sw., Cyperus rotundus L., Echinochloa crus-galli (L.) Beauv., Eragrostis nigra
Nees ex Steud., Hydrolea zeylanica Vahl., Imperata cylindrica (L.) Raeuschel,
Ischaemum indicum (Houtt.) Merr., Leucas aspera Sprng., Paspalum
scrobiculatum L., Pennisetum typhoides (Burm. f.), Polytrias amaura (Buse) O.
Kuntze, Solanum torvum Sw., Sorghum bicolor (L.) Moench., Triticum vulgare Vill.
Cereal thrips Haplothrips ganglbaueri, is a polyphagous species infesting the inflorescence of
various cereal crops such as rice, wheat, sorghum and millets. The pest also abounds in several
weeds common in and around rice fields. A positive correlation often exists between the thrips
populations in the weeds and in the rice plants (Ananthakrishnan and Thangavelu 1976). The
population build-up of the insect is very high and outbreaks have been reported from many parts of
India (Chaudhary and Ramzan 1971, Abraham et al. 1972).
Due to the laceration and feeding by larvae and adults of H. ganglbaueri, the rice inflorescence
is variously affected. The damage on the lemma and palea causes development of irregularly oval
and diffused brownish patches (Abraham et al. 1972). Vidyasagar and Kulshreshtha (1983)
reported three distinct types of damage: (1) light brown spots with a perforation at the proximal part
of the unopened spikelets, (2) sterile spikelets that retain their green color throughout their
development and (3) oozing of milk over the grain without any external feeding marks. The affected
panicles are erect due to the dropping of damaged grains. Continuous feeding causes chaffy grains.
The adult thrips are dark brown with a dominant reddish tinge. The average lengths of males
and females are 1.3 and 1.6 mm respectively. The preoviposition period lasts for 2-3 days. Eggs are
laid on the outer surface of the lemma and palea mostly in clusters of 4,8 and 12 (Ananthakrishnan
and Thangavelu 1976). Eggs hatch in 2-4 days. There are two larval instars which together last for
8-11 days. There is a short prepupal stage of 1 day which then passes to the pupal stadium. The
pupal period is 3-4 days. Female thrips live for 12-15 days while males live for only 3-5 days.
Incidence in South India is serious during the first fortnight of December. During this period,
the weather is humid with occasional drizzles and the sky is often overcast for most of the day.
DALE 471
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4
Rice Pests and Agricultural Environments
MICHAEL E. LOEVINSOHN
International Development Research Centre
Ottawa, Canada
1. INTRODUCTION
THIS CHAPTER FOCUSES ON the ways in which the fundamental forces affecting pest species, that
determine the presence and abundance of pests in a particular area and that govern their as-
semblage into communities, are shaped by the manner in which rice is grown. These forces,
generally considered as “natural” (the impact of weather and micro-climate, the limitations im-
posed by host plant availability and quality and the action of natural enemies), are profoundly
influenced by decisions at various levels, from the farmer’s to the policy-maker’s. Changes, for
example, in the time of planting, in irrigation practices, and in the spatial or temporal intensity of
rice cultivation relax or tighten the constraints on pest species by modifying the fundamental forces.
All too often, the effects are unintended and poorly understood by scientists and farmers. Effective
pest management demands, however, that we learn to predict the consequences of human action
and that we use this knowledge to better plan interventions.
Two themes dominate this chapter. The first is that the response of pests to alteration of their
environment is dynamic. Evidence of adaptation by pests can be found today in patterns of genetic
differentiation between populations. Evolutionary responses have been best studied with respect to
control measures, particularly the widespread use of insecticides and host plant resistance, but they
are evident as well in the face of changes in the timing, intensity and synchrony of cultivation.
Though there is little direct evidence of the process, it is quite possible that over the 10,000-odd
years that rice has been cultivated, human modification of natural forces has led to the emergence
of new pest species. The dynamic nature of pest response is also visible at the community level:
environmental change that acts to the detriment of one species may offer new opportunities to
another, leading to changes in species composition.
488 RICE PESTS AND AGRICULTURAL ENVIRONMENTS
The second theme that will be developed is that the key processes at work are scale-dependent.
The practice of rice entomology has been pre-occupied with the dynamics of individual species, for
the most part at the smallest of spatial and temporal scales. As one’s attention turns to forces acting
on populations and communities at scales greater than the dimensions of a field and longer than a
season, one is struck by the paucity of data and the abundance of speculation. The presence,
absence or relative importance of a species in an area has generally been treated as a given. This
orientation is perhaps understandable given the often pressing needs of control. But there has been
little basis on which to predict the impact of widespread agricultural change or to explain differen-
ces in the pest situation between rice environments. In the course of this chapter we point to new
methods and approaches that may provide a basis for a more forward-looking environmental
management, holding out the promise of avoiding rather than merely coping with pest problems.
2. THE ASSEMBLAGE OF PEST FAUNAE: MODIFICATION OF THE

PHYSICAL ENVIRONMENT
2.1 CLIMATE AND RANGE

Rice is grown on every continent, save Antarctica. It can be found from near 50ºN in China (IRRI
1978) to 35ºS in Australia (Moomaw and Vergara 1965), and from sea level to over 2600 m in Nepal
(Shahi and Heu 1979). The majority of rice production, however, is located within the tropics and
subtropics. Of the world total, some 90% is within Asia, 4% in Latin America and less than 2% in
sub-Saharan Africa (Palacpac 1982). Though few Asian countries are able to bring much new land
under rice cultivation, considerable scope for expansion remains in both Africa and Latin America.
Climatic factors vary widely across the range of rice cultivation. Temperature, humidity and
wind affect the rates of increase of pest populations directly, by limiting longevity, fecundity and
dispersal, and indirectly, through their effects on host plant vigor and natural enemy action. Pests
also make use of physical cues correlated with the availability of host plants, such as photoperiod
and humidity, to break the inactive states of quiescence and diapause. The literature on climatic
effects on rice pests has been reviewed recently by Mochida et al. (1986).
As man extended rice cultivation beyond its center of origin in Southeast Asia and southern
China, it was often to areas of greater climatic extremes. Individual insects able to make use of a
more widely available host were at a selective advantage within their populations. Where this ability
was due to a genetically determined tolerance or behavioral plasticity, natural selection resulted in
genetic differentiation. Though direct evidence of the process is impossible to obtain at this point,
perhaps centuries after the fact, the results of the process should be visible in differing tolerances
of populations inhabiting areas of contrasting climate.
Therc has been little research on this topic, particularly within the tropics. Many might consider
it of only academic interest that, say, yellow stem borers in the hot and arid Punjab show greater
resistance to high temperatures than those from Japan. But a similar process has occurred on
repeated occasions on a more local level and with more immediate consequences. I refer to the
LOEVINSOHN 489
response of pests to the new opportunities that have arisen across much of tropical Asia in the past
few decades with the shift from single to double rice cropping made possible by irrigation. In many
locations, there are marked differences between wet and dry seasons in temperature, relative
humidity and wind speed and direction.
2.2 ADAPTATION TO THE DRY SEASON

Few if any pests have been totally unable to colonize the dry season crop. Reliable contemporary
records are frustratingly rare, but in most cases the available evidence suggests that the lag between
the widespread adoption of dry season cropping and the appearance of damaging infestations has
been either nil or very brief.
The rice gall midge provides an interesting case study. Hidaka and his co-workers (Hidaka et
al., 1974) studied the seasonal dynamics of this pest in an area of northern Thailand where a single
crop of rice is grown each year from March to November. They found that after cultivated rice had
passed the vegetative stage and could no longer support infestation, females oviposited on nearby
grass hosts, particularly wild rice and, to a lesser extent, Paspalum distichum. Larvae were observed
there throughout the dry season, however, few large and readily visible galls could be found before
the wet season rice crop was established. Low relative humidity, the authors suggested, arrested the
insect’s development. Experimental evidence supported this conclusion: not one of 50 newly
hatched larvae placed on dry rice plants achieved penetration, while all of the larvae on plants
sprayed with water managed to do so (ibid., pages 54-62). More striking yet was their observation
that, in a 400 m2 field planted on-station during the dry season, not one gall was found in nearly
3,000 hills, and only 2 larvae upon dissection of the stems. The authors recommended planting a dry
season crop to avoid damage by the gall midge (ibid., pages 74-75).
These findings, however, might be looked at in a different light. As the authors noted, farmers
in that part of northern Thailand planted only during the wet season in the late 1960s and early
1970s. In such an environment, individual gall midges able to make use of a dry season crop, due to
their tolerance of drier conditions, responsiveness to different phenological cues and/or altered
dispersal behavior, had no particular selective advantage. Indeed, most likely the contrary. The
situation would be radically changed were rice to be grown in the dry season o n a regular basis and
over an appreciable area. Insects capable of colonizing the dry season would have an enormous
edge in fitness, especially initially, before natural enemies had themselves adapted to the new
conditions. How long a dry season crop need be cultivated and over how wide an area before natural
selection can be expected to produce a population fully able to exploit the new resource depends
on such factors as the numbers, likely very low, of “adapted” individuals in thc original population.
the rate of mutation of the necessary traits and the rate of immigration from populations where they
already exist. These factors would determine the durability of dry season cropping as a means of
escaping the gall midge.
There is evidence from several sources that the dry season in fact constitutes a fairly minor
evolutionary barrier to the gall midge. Four years after their studies in northern Thailand, Hidaka
and a different set of co-workers (Hidaka et al., 1978) reported on a serious infestation of gall midge
in the dry season in the Central Plain. There, farmers had planted the crop over nearly 500 ha and
were at the time in their second year of double cropping. The authors attributed the outbreak to the
susceptibility of the variety that had been used and, in particular, to the supposedly moister
micro-climate resulting from direct seeding and closer plant spacing. Yet serious dry season
infestations are now a common occurrence in areas of the Central Plain where double cropping is
regularly practiced (Hidaka and Yaklai 1980), as appears to be the case elsewhere in Asia
(Soeharjdan et al. 1973, Kalode 1974, Bhaskara Rat, 1981, Chand 1981). In the light of these
developments there is good reason to question the wisdom of Hidaka et als’ (1974) earlier recom-
mendation.
The purpose here has certainly not been to latch onto someone’s unfortunate turn of phrase,
secure in the wisdom of hindsight. This particular case illustrates as clearly as any could a principle
to which I will have occasion to refer again: the fundamental processes of population genetics and
population dynamics operate at characteristic scales. What one confidently documents at the scale
of the experimental plot of 100 m2 over a season or two is likely a poor guide to what will transpire
when the practice is implemented on hundreds or thousands of hectares and over a period of years.
2.3 ADAPTATION TO ALTERED TIMING OF CULTIVATION

In other situations, advancing or delaying the time of cultivation within a season has rendered
unreliable physical cues that pests had depended on to end quiescence or diapause. Such changes
have in some instances been introduced specifically as a control measure; elsewhere other con-
siderations have been determinant. Whatever the reason, the altered timing has presented pests
with an evolutionary challenge that has been met with variable success.
Two relatively well documented cases involving stem borer species may indicate the range of
responses. Research conducted in Java prior to independence (van der Goot 1925) showed that the
length of the larval diapause of white stem borer (Scirpophaga innotata) in rice stubble and the
timing of adult emergence was correlated with the date of the first rains of the wet season. The
relationship was sufficiently close that safe dates for transplanting, after which minimal infestation
might be expected, could be calculated once the first rain had fallen. From 1929 to 1941, planting
over an area of several thousand hectares was forbidden by regulation prior to the announced date.
The willingness of the colonial administrators to enforce the regulations strictly and the apparent
lack of variation in the white stem borer population with respect to diapause-breaking or emergence
cues combined to render the control measure remarkably effective over the 12 year period (van der
Goot 1948). Similar results have been obtained in northern Australia and Queensland (Halfpapp
1982).
In Japan after the Second World War, the development of cold-tolerant rice varieties allowed
farmers to plant earlier in the year. Although early planting had been suggested as a means of
avoiding damage due to the striped stem borer, it appears primarily to have been agronomic
concerns, particularly the possibility of increased double cropping that motivated most farmers to
adopt the new varieties. Kiritani and Iwao (1967) provide evidence that, as the area on which rice
was planted early increased, striped stem borer itself emerged earlier. Experimental evidence
LOEVINSOHN 491
confirmed that genetic differentiation was responsible for the phenomenon: insects taken from
areas where early rice was widely grown required fewer degree-days in spring to achieve emergence
from their over-wintering sites.
Aside from these two cases, there are few well-documented studies of the long-term effective-
ness of changed planting dates practiced over a wide area. The literature, however, is replete with
studies that have charted the seasonal pattern of pest abundance as determined by infestation in
experimental plots planted at different times. From this information, many authors have derived
recommendations regarding safe planting dates, either earlier or later than the prevailing practice
(This literature is reviewed in Chapter 6 in the context of cultural control). Here, as in the case of
adaptation to the dry season, scale-dependence is expected where evolutionary processes are
involved.
In many instances, a pest species can continue to exploit a rice crop that has been shifted in
planting date only by means of physiological or behavioral adaptations of a genetic nature. If the
area planted on the new schedule is small relative to the typical dispersal distance of the insect, the
selection pressure on the population will be low. That is to say, individuals (“phenotypes”, in genetic
terms) able to infest the new planting will not be better represented than other forms in succeeding
generations, as their offspring will disperse into areas following a different schedule where they will
be at a disadvantage. As the area planted on the new date increases, so does the advantage of those
phenotypes in terms of Darwinian fitness. The longer the practice continues, the greater the chances
that an adapted phenotype will emerge, through mutatation, recombination or immigration, if it
does not already exist within the population.
An important constraint to such adaptation may be the complex nature of the diapause
phenomenon in the species where it occurs. It is only phenotypes responsive to different cues or
with different responses to the original ones that will be able to exploit a rice crop shifted in time. It
appears to have been such a failure to adjust its diapause that prevented the white stem borer from
overcoming the delayed planting that was enforced in Java for 12 years. Yet pests of other crops are
often known in both diapause and diapause-free forms. Interestingly, the white stern borer in a
double cropped area in south Cotobato in the southern Philippines appears to have been able to
discard the diapause nature (J.A. Litsinger, personal communication).
The ability to exploit rice appearing at any time of year is characteristic of the major pests of
intensive tropical rice cultivation. In much of the tropics, neither temperature nor humidity prove
to be absolute constraints on pest activity. Lack of diapause and responsiveness to cues not linked
to a particular time of year appear to be the correlates in the insect of photoperiod-insensitivity in
the plant. Many authors however appear to assume that a particular seasonal pattern of abundance
represents a dependable species characteristic, rather than reflecting adjustment to the local
availability of the rice host (Fig. 1). In tropical areas, a recommendation to shift planting backwards
or forwards in order to escape pests that do not possess a diapause is likely to be of benefit to the
first people adopting it, but much more disappointing to the majority of farmers should they follow
suite.
Median Planting Date Within 1 km
Fig. 1. Yellow stem borer activity in relation to delay in planting. Due to damage to the main canal that limited
irrigation flows, planting along Lateral C of District III, Upper Pampanga River Integrated Irrigation
System, Philippines, was unusually drawn out during the 1981 dry season. At each of 7 sites along a 13 km
stretch of the canal, two kerosene light traps were installed. Median flight dates are the day by which 50%
of the seasonal total in those traps had been caught. Median planting dates were similarly calculated for a
circle of radius 1 km centered on the traps, making use of records of the National Irrigation Administration.
The equation for the regression is : Y = 159 + 0.520 X (r = 0.86, P < .02). where X and Y are measured
in days from January 1. Loevinsohn (unpubl.).
3. THE ASSEMBLAGE OF PEST FAUNAE: THE INFLUENCE OF THE

CROPPING SYSTEM
We have been concerned to this point with the effect of the physical environment on the occurrence
of insect pests at one location, particularly as cultivation is extended into the dry season or shifted
in time. We consider now spatial patterns of species occurrence in relation to the form of rice
culture that is practiced and the distribution of rice in the landscape.
3.1 CULTURAL TYPES

Three principal cultural types may be recognized, distinguished by the degree of flooding (De Datta
1981). In upland cultivation, flooding is no more than transient, in lowland culture standing water is
present more or less continuously but does not exceed 1 m, while in deepwater areas flooding
regularly exceeds this depth at some point in the crop’s growth. As Table 1 indicates, irrigated or
rainfed lowland rice is dominant in tropical Asia, while a significant fraction of the rice area in
Africa and Latin America is devoted to upland culture.
LOEVINSOHN 493
TABLE 1
Harvested rice area (percentage) in four cultural types by region, 1985. Adapted from IRRI (1989) a
Region Irrigated Rainfed lowland upland Deepwater Total (× 1000 ha )

South Asia 32 40 13 15 60,270
Southeast Asia 38 40 11 12 31,150
East Asia 92 6 2 0 34,890
Africa 22 28 46 3 4,510
Latin America 32 6 62 0 7,180
aAreas of mangrove swamp cultivation in Africa are included in the rainfed lowland total.
Environmental conditions are often sufficiently different as to effectively restrict Certain pests
to one or another of these cultural types. White grubs (Scarabaeidae), for example, widely dis-
tributed pests that attack the roots of rice in upland cultivation, cannot survive in the saturated soils
of lowland or deepwater culture. For other species, the differences between cultural types may be
reflected in varying rates of population increase and hence in abundance and pest status. This is the
case for example with the brown planthopper: a major pest of lowland cultivation, it is of generally
minor importance in deepwater rice (Catling et al. 1987a). Recent descriptions of cultural types as
habitats for insect pests and of the fauna encountered are provided by Litsinger et al. (1987) for
upland cultivation and by Catling et al. (1987b) for deepwater rice.
However, the form of rice culture practiced in an area is only one determinant of the composi-
tion of its pest community and of the abundance of individual species. The spatial scale over which
rice is cultivated and the proximity of alternative host plants, as well as the seasonal pattern of
cultivation within the area will affect the success of different life-history strategies, that is, the
characteristic ways in which pest species exploit crucial resources. Certain trends can be recognized
in the data available.
3.2 SPECIALIZATION AND THE GEOGRAPHY OF CROPPING SYSTEMS

One of the dimensions along which insect lie-histories can be classified is the degree of specializa-
tion in the use of host plants. To the extent that specialization is associated with greater efficiency
in the exploitation of the plant resource (Roughgarden 1972) monophagous and oligophagous pests
are expected to be relatively more abundant where their host plant is concentrated than
polyphagous insects of otherwise similar biology (ie. within the same ecological guild). In other
agricultural contexts, specialist herbivores are indeed found to be more abundant in fields where
monoculture rather than mixed cropping (polyculture) is practiced (Risch et al., 1983, Altieri and
Letourneau 1984). Rice being seldom intercropped with other plants (except in certain cases of
upland cultivation), our primary concern is with “resource concentration” at a larger scale, beyond
the field. Yet similar trends are observed at this level as well. A number of studies have examined
the distributions in different environments of stem boring insects that are either monophagous on
rice or that can also infest other grasses and crop plants. In Kenya, Ho and Kibuka (1983) found that
in upland areas, where rice was often grown in proximity to other cereals, the principal stem borers
were Chilo partellus and Sesamia calamistis which also attack sorghum and maize. In the lowlands
where large tracts of irrigated rice are found, the monophagous Maliarpha separatella was
dominant. A similar pattern is apparent in Madagascar (pers. observation, M. Loevinsohn).
Surveying the composition of the stem boring guild in the Philippines, Cendaña and Calora
(1967) found the polyphagous Chilo suppressalis dominant on rice farms in Laguna province, while
some 100 km away in Central Luzon, the monophagous Scirpophaga incertulas was almost exclusive-
ly present. The authors proposed climatic factors in explanation, but though Laguna is somewhat
moister and has a less extreme range of temperatures, the difference is far less than either species
has successfully accommodated elsewhere in its geographic range. A more significant difference
may be the fact that in Laguna, rice cultivation is concentrated in a narrow strip surrounding Laguna
de Bay, and upland crops, such as maize (an alternative host of C. suppressalis) are planted nearby
on higher land. In contrast, over much of the broad Central Luzon plain, rice is the only crop grown.
This hypothesis gains credence when one examines trends in species composition at one
location over time. In Laguna, major changes have occurred in farming systems since the late 1960s,
changes that are often collectively referred to as the Green Revolution. One element in this process
has been an increase in the extent of dry season cropping: in the irrigation systems surrounding the
IRRI farm at Los Baños, the proportion of farm area double-cropped rose from 39% in 1969 to 99%
two years later (Loevinsohn 1984). Three fluorescent light traps have been in operation at IRRI
continuously since 1965 and provide a record of insect abundance in the vicinity. As Table 2
indicates, mean annual catches of both S. incertulas and C. suppressalis have increased, though the
former to a significantly greater extent than the latter (176% vs 65%).
TABLE2
Contrasting effects of agricultural change on specialist and generalist pestsa
Mean annual catch Ratio

Years Scirpophaga incertulas Chilo suppresalis Scirpophagal/Chilo
1965-70 4370 670 6.49
1971-79 12042 1107 10.9
a Mean annual catches and their ratio for two stem borer species, one monophagus on rice (S. incertulas), the other
(C. suppressalis) polyphagous, in periods of lesser (1965-70) and greater (1971-79) incidence of double cropping. The
change in the ratio of the species is significant ( c 2 = 688, P < .001). Annual totals are for 3 light traps on the farm of the
International Rice Research Institute at Los Baños in Laguna Province, Philippines.
A similar trend is observed in N. Krian, Malaysia, where agricultural change appears to have
included the same key elements as in Laguna (Loevinsohn 1984, Loevinsohn et al., 1988). In
particular, the completion of a major irrigation project made dry season cropping possible for most
farmers for the first time. Light traps were used to monitor the occurrence of several pests,
including S. incertulas and the dark-headed stem borer Chilo polychrysus, which, like C. suppressalis,
attacks cereals other than rice. S. incertulas increased in abundance relative to its polyphagous guild
mate coincidentally with the spread of double cropping. Explanations for why the Chilo species
increased in the one case and declined in the other must be sought in the distribution of alternative
hosts and in the detailed biology of the two insects. We will return to the light trap records of Laguna
LOEVINSOHN 495
and N. Krian in a later section when we consider more closely the population dynamics of rice
specialist pests under agricultural change.
3.3 SPECIALISTS vs GENERALISTS: DISTRIBUTION AND IMPORTANCE

Although comparative studies appear to have concerned themselves most often with stem borers,
the dominance of specialist life-histories in areas of intensive rice cultivation can be observed for
the pest fauna as a whole. This pattern is likely to become increasingly generalized as further
investment in irrigation enlarges the area under intensive cultivation. Two related trends are worthy
of particular note.
Firstly, the specialist pests appear also to have been the most successful in following rice as its
cultivation spread to new areas and thus today are the most widely distributed. This trend is most
apparent within Asia, a point to which I return below. Monophagous and oligophagous pests such
as brown planthopper, green leafhopper, yellow stem borer, gall midge and caseworm are common
across tropical Asia and several of these are common in the semi-tropical and sub-temperate
regions, while, with a few notable exceptions, generalists tend to have a more restricted distribution.
The similarity of lowland rice culture across the continent and the moderating effect on
microclimate of irrigation likely facilitate the spread of specialist pests.
Secondly, specialists appear to be in general the most damaging to the crop. Having coevolved
with rice as cultural practices developed, these insects are efficient at locating and tracking their
host, overcoming its chemical and physical defenses and surviving the fallow between crops. Large
populations, in prolonged contact with their host, would be expected to extract substantial amounts
of photosynthate from the plant, and from the yield at harvest. Disease organisms that cause
significant mortality to their host in a natural population may, over time, evolve reduced virulence,
for their own persistence is dependent on the survival of at least some hosts (Lenski 1988). In
agricultural systems similar selective pressures are unlikely to be acting on specialist pests as
farmers replant the crop each season.
3.4 FALLOW SURVIVAL STRATEGIES

Specialization is certain to prove a more complex phenomenon than the simple categorization
above suggests. While a species as a whole may be found to have a broad host range, individual
populations may display considerably greater discrimination (Cates 1981). As well, polyphagy may
have a seasonal character: a species effectively restricted to a preferred host at one time of the year
is obliged to shift when it is no longer available. A second important dimension therefore along
which pest life histories can be assessed is their strategy for surviving the fallow period between rice
crops. Four principal types may be identified:
(1) “Promiscuous” pests are those that abandon rice, and often the semi-aquatic paddy
environment, after the harvest. These are generally species that show no marked
preference for rice or adaptation to efficiently exploiting it and are polyphagous at an
individual level. Examples are the armyworm Spodoptera spp., the oriental locust Locusta
migratoria and, in Africa, the African pink borer Sesamia calamistis and the stalk-eyed
borer Diopsis apicalis.
(2) “Companion” pests on the other hand prefer rice and are adapted to aquatic conditions.
Following the harvest they move to aquatic weeds and grasses, on which their rate of
increase is found (e.g. for the caseworm) (IRRI 1981) to be less than on rice and indeed
often negative. Other Asian examples of this strategy are the rice green semilooper
Naranga aenescens and the green leafhopper Nephotettix virescens.
(3) “Stick-it-out” strategists are oligophagous or monophagous and generally remain with
the rice plant through the fallow. Some form of quiescence or diapause is usually found.
The most striking examples are among the specialized stem borers: yellow stem borer and
white stem borer in Asia and Maliarpha separatella in Africa.
(4) “Vagabond” species are also rice specialists but escape the fallow by dispersing to areas
following different cropping schedules. The best known example is the monophagous
brown planthopper.
3.4.1 Pest Strategies and the Availability of Rice

Each of these strategies has different habitat requirements and is affected differently by increases
in the extent and timing of rice cultivation (Fig. 2). A promiscuous life history may be the only viable
one when rice is introduced to an area and when it is a minor element in a diverse floral landscape.
Such habitats, and pest faunae dominated by promiscuous species, can be found in the more
isolated rice growing regions, including many parts of Africa. For example, rice was introduced to
Rwanda some 25 years ago and is today cultivated in widely separated valleys, nowhere more than
600 ha in extent. The only pest commonly encountered is a Diopsis sp., clearly polyphagous for it is
already abundant several days after transplanting in valleys where rice has never before been grown
(M. Loevinsohn, pers. obs.).
Where rice is cultivated more extensively, and over a longer period of time, a companion
strategist that can more efficiently exploit the crop will be able to build up large populations and
hence selective pressures will emerge for specialization. As has been observed for other crops and
plants, this may involve adaptation by a polyphagous species or the recruitment to rice of a specialist
on related grasses or possibly on plants in a different family (Turnipseed and Kogan 1976, Strong et
al., 1984). Polyphagous pests that do not adapt to rice and the paddy environment may suffer from
the extension of rice culture, particularly if they depend on alternate hosts that are displaced.
With the increased area under cultivation, a stick-it-out strategist may be able to maintain a
viable population. A strictly monophagous pest that exploits an annual crop like rice requires a
minimum degree of asynchrony in cultivation between fields if it is to persist and such variation in
the timing of farm operations is typically found to increase with area (Loevinsohn 1984). To see
more clearly the importance of asynchrony, consider an insect such as the yellow stem borer. It
passes the fallow period as a quiescent late instar larva in the base of rice stubble (Bannerjee and
Pramanik 1967), and must pupate and emerge before its host is destroyed by the plowing, harrowing
LOEVINSOHN 497
Fig. 2. The relationship between pest life history strategy and the availability of rice. Fallow survival strategies are
defined in the text. Solid lines suggest optimum environments for a given strategy, broken lines more
marginal conditions.
and puddling of the field. Even if it puts off emergence to the last moment, the adult cannot possibly
oviposit in the field in which it developed or others planted at the same time: it lives only a few days,
while land preparation may take several weeks and the new crop cannot support an infestation until
perhaps 20 days after transplanting. Successful oviposition thus depends on the adult locating a field
sufficiently delayed or advanced in the cultivation sequence within its flight range.
The requirement for asynchrony is even greater in the case of a vagabond strategist such as the
brown planthopper: without diapause or quiescence, it depends on a succession of rice plants at a
stage suitable for infestation. Where a single wet season crop is grown each year, dispersal on the
scale of 10s or 100s of kilometers may be needed to find its hosts. Numbers reaching a point are
typically found to decline exponentially with distance and thus the proportion surviving to infest
another crop may be very low. In a northern semi-temperate area such as Japan, where “environ-
mental resistance” (including in particular natural enemy action) is relatively weak, the low density
of brown planthopper immigrants that appears responsible for reestablishing the population each
year is able to multiply to economically significant levels (Kuno and Hokyo, 1970). In tropical areas,
the more effective and consistently density-dependent response of natural enemies to the brown
planthopper appears sufficient to contain population increase where only one crop a year is grown.
Twenty five years ago, the brown planthopper was considered of no more than occasional impor-
tance — at a 1964 symposium on the “Major Insect Pests of the Rice Plant” (IRRI 1967) it was paid
only cursory attention — and today its abundance in areas of single cropping appears often to be
related to the distance from more intensive cultivation (Perfect and Cook 1987).
The most recent stage of habitat evolution in Asia has been that associated with the Green
Revolution. Although, as will be discussed below in greater detail, the process has been complex,
common features have been the marked expansion in double cropping and increased asynchrony in
the timing of cultivation that has often followed in its wake. In intensive rice cultivation, promis-
cuous pests find it difficult to locate non-aquatic host plants and appear seldom to be of economic
importance. Companion and stick-it-out strategists, on the other hand, are favored by the reduced
fallow and the shorter time they must remain on less preferred alternative hosts or in quiescence or
diapause. We have already seen evidence of a shift in the relative importance of promiscuous and
stick-it-out pests in the case of stem borers. But it is vagabond species that appear to have been the
most favored by the increased availability of rice. The rise of the brown planthopper and other
Homoptera to the status of major pests is among the most important and widespread environmental
impact of the Green Revolution.
3.4.2 Agricultural Development and Species Establishment

It is certainly a crude simplification to reduce the diversity of pest life histories to only four types.
Similarly, the evolution of rice agroecosystems has not everywhere followed the same path toward
greater host availability. Changes in the various components of host availability (areal extent and
time within and between seasons) are likely to have quite different effects on pest populations and
communities (see further below). Yet even in its present form, we believe the approach is of value
in emphasizing that the dominant species in a pest community are far from a random selection of
those present in the regional pool. Species have particular requirements, dictated by their life
histories, for rice; whether these can be met depends on the pest being able to locate suitable stages
of its host within a distance it can disperse. This perspective implies that it is a combination of the
area over which rice is cultivated, the number of crops grown per year (cropping intensity), the
asynchrony with which they are established and the isolation from other rice growing regions that
determine the success of a given species and of the life history strategy it embodies.
We explore below some ways in which these habitat parameters may be combined in quantita-
tive form and employed to explain variation in the abundance of pests. Qualitatively however, the
framework permits some broad predictions about the evolution of the pest fauna as rice cultivation
develops. Returning to the example of Rwanda, we find that the production of rice rose almost
5-fold between 1970 and 1985, faster than any other food crop (Government of Rwanda 1987). The
area cultivated, in many valleys now irrigated, is expanding rapidly and double cropping is becoming
the norm. On the basis of what has been said above, one might expect that companion and
LOEVINSOHN 499
stick-it-out strategists could now maintain viable populations in some areas. Similarly, cultivation in
certain coastal areas of Madagascar is now sufficiently extensive and asynchronous that a vagabond
pest might be able to establish itself.
It is impossible to predict, in either case, whether the pest will be a long distance migrant or
whether it will emerge from the indigenous fauna as a result of an expansion or shift in host range.
The probability of invasion might be thought higher in the case of Rwanda: Maliarpha separatella,
the stick-it-out stem borer, is found in neighboring Uganda and Tanzania, whereas Madagascar is
isolated by several thousand kilometers from Asian sources of vagabonds. However, factors other
than distance affect the likelihood of such long-range dispersal, notably wind direction and speed.
Human agency, “assisted transport” by plane or surface means, has also played a role in past
invasions and may have been responsible for the introduction of the rice water weevil Lissorhoptrus
oryzophilus Kuschel, of American origin, to Japan. But such a chance event, represented perhaps
by the arrival of a single gravid female, is a necessary but not sufficient condition for successful
establishment: cultivation must have evolved to a stage where rice is sufficiently available to support
the pest’s life history.
4. PEST POPULATION DYNAMICS AND THE CULTURAL ENVIRONMENT
4.1 THE IMPACT OF AGRICULTURAL CHANGE

The shifts we have been considering in the composition of the pest fauna as agriculture evolves can
be followed as well in the rising and falling fortunes of populations of particular species. It is in
periods of rapid agricultural change that one might expect the effects of the cultural environment
on pest populations to be most clearly highlighted. In practice, however, such change is rarely a
simple process: many aspects of the environment typically vary together and their separate effects
may be difficult to unravel.
The Green Revolution that transformed rice cultivation across much of tropical Asia in the
1960s and 1970s is a case in point. The revolution entailed widespread adoption of modern varieties,
greater use of chemical inputs and expansion of irrigation that brought in its wake increased
cropping intensity and asynchrony of planting. Each of these elements has been at some point
implicated in the marked increases in the abundance of specialist pests that typically followed. The
division of opinions and resulting confusion is nowhere better illustrated than in the proceedings of
the 1977 symposium that sought an explanation for the widespread outbreaks of the brown plan-
thopper (IRRI 1979). The focus of research on individual factors has made it difficult to put their
relative contribution into perspective or to support a comprehensive understanding of rice pest
dynamics to guide management.
A recent attempt to come to grips with this challenge (Loevinsohn and Litsinger, in press)
draws on series of light trap catches spanning the period of agricultural change. At the IRRI farm
in Laguna, Philippines, annual totals for three of the principal pest species increased markedly
Fig. 3. Population of three rice specialist pests over a period of agricultural change. Annual total catches are for
three fluorescent light traps on the IRRI farm at Los Banos, Laguna, Philippines.
beginning in the late 1960s (Fig. 3). Similar changes at about the same time occurred for yellow stem
borer and the green leafhopper in North Krian, Malaysia.
In all 5 cases, the rise and fall of catches between months reflect the local pattern of cultivation,
making it possible to identify seasonal minima, ny , maxima, Ny, and totals and to examine the forces
regulating pest populations. Two statistics may also be defined the within-season rate of change,
Rwy = ln Ny – ln ny (1)
and the (negative) between-season rate of change,
RBxy = In nx – ln Ny (2)
LOEVINSOHN 501
where season x precedes y. The changes that have occurred in the agricultural environment can be
divided into those that affect RW, such as increased use of agrochemicals, and those that affect R B,
notably increased dry season cultivation with irrigation. Asynchrony of cultivation between fields,
often exacerbated by irrigation, acts on both rates, extending the time available to mobile insects for
increase and reducing the length of the fallow they need endure.
4.1.1 Population Change Within and Between Seasons

One means of assessing the relative importance of factors acting within and between season is to
plot log maximum numbers during a season against log minimum numbers at the beginning of that
season. As is illustrated for the brown planthopper in Figure 4, significant positive correlations were
obtained in 4 of the 5 cases in the wet season, with the coefficient of determination (the proportion
of variation in maximum numbers accounted for by variation in minimum numbers) in the range of
47% to 78%. The residual variation reflects the contribution of changes in factors acting within the
season, in addition to measurement error and imprecise model formulation. In certain cases, it is
possible to implicate particular developments in these deviations: for example, the first and third
largest positive residuals from the regression for brown planthopper occurred in 1973 and 1976
when varieties incorporating the Bph 1 and bph 2 genes respectively succumbed to evolved virulence
Fig. 4. Dependence of pest population on initial numbers. Maximum 4-weekly catches of brown planthopper
during the wet season in relation to minimum catches at the beginning of the season; IRRI farm Laguna.
1967-1986. The best fitting equation using Bartlett’s technique (Sokal and Rohlf 1981), which provides
unbiased estimates of regression parameters when both variables are measured with error is: In Y = 6.90
+ 0.594 In X (P < .01).
in the insect (Loevinsohn et al., 1988). Continuous records of within-season factors that would make
it possible to put such an analysis on a firmer footing are unfortunately not available. Nevertheless,
the results suggest that factors acting within a season are no more and are often less important in
determining maximum numbers that season than are those that affect initial levels.
What then determines these initial levels? Two variables prove to be significant: maximum
numbers during the preceding season (4 of 5 cases) and the number of months between the seasons
(again, 4 of 5 cases). In multiple regressions, the two variables together account for from 4.6% to
95% of the variation in log minimum numbers in the wet season.
The size of the preceding season’s population is determined both by within-season factors and,
when it is a dry season, by the proportion of the farm area on which double cropping is practiced.
However, only the between-season factors are implicated in the inverse correlation between the wet
season minimum and time between seasons. The data are inadequate to calculate an index of
asynchrony, but the effect of cropping intensity can be clearly seen. In N. Krian, there were, on an
average, 6.0 months between seasons for yellow stem borer in the years when double-cropping was
practiced on less than 10% of farms in the surrounding irrigation system, compared to 2.9 months
thereafter (t = 2.53, P < 0.05). The overall impact of cropping intensity on the numbers surviving
Fig. 5. Effect of cropping intensity on population dynamics between seasons: yellow stem borer in N. Krian,
Malaysia, 1965-75. The rate of population change between successive wet seasons was calculated as the
difference between the in minimum 4-weekly catch in the light traps at the beginning of one wet season and
the In maximum catch the wet season previous. The rice cropping index, the mean number of crops per year
on land devoted to rice cultivation, is calculated for the adjacent Krian/Sungeri Manik irrigation scheme.
From data in Lim and Heong (1977). The least squares regression is: Y = -7.25 + 2.53 X; t = 4.71,
P < .005.
LOEVINSOHN 503
to infest a crop is illustrated in Figure 5: RBww , the rate of change between successive wet seasons,
increased with the spread of dry season cropping. Significant relationships of this type are found in
3 of the 5 cases.
4.1.2 Population Regulation and Cropping Intensity

A double logarithmic regression as in Figure 4 provides important information about the extent of
density-dependence in the population's rate of change. A slope that, as in this case, lies between 0
and 1, implies that the within-season rate of change is imperfectly density-dependent, that is, it
declines as the seasonal minimum rises, but to an extent insufficient to compensate for the variation
in these initial levels. The same is found to be true for the 3 other populations for which significant
regressions were obtained. The pervasive and effective control exerted on rice pests in the tropics
by their natural enemies, described in Chapter 7, is likely responsible in large measure for this
density-dependence.
In contrast, the rate of change between seasons is found in each case to be independent of
density, that is, minimum numbers at the beginning of a wet season are a more or less constant
proportion of the preceding maximum when the time between the seasons is held constant. This
finding should not be surprising, as survival through the fallow of a stick-it-out pest like yellow stem
borer or a vagabond like brown planthopper appear in large measure to be a function of physical
factors (desiccation and flooding in the first case, distance and meteorological conditions in the
second) that do not respond to density.
This pattern of imperfect density-dependence within seasons and density-independence be-
tween is found once again when one examines the determinants of pest abundance in the dry season.
These findings can be combined in a model of population dynamics over the whole year, one
prediction of which is that increases in the area planted to rice in the dry season will lead to larger
populations not just in that season but in the wet season as well. This was indeed found to be the
case when pest abundance was measured in areas of the Philippines differing in cropping intensity
(Fig. 6). Crop losses appear to parallel pest numbers: the proportion of the wet season harvest lost
to insects averaged 13% in the sites where a single rice crop was planted each year, compared to
33% where double cropping was practiced (Loevinsohn et al., 1988).
The above results suggest an underlying similarity in the annual dynamics of rice specialist pests
and point to limitation of cropping intensity as a fundamental element in their management. Further
reduction in the length of the fallow as a consequence of triple cropping would be expected to result
in yet higher seasonal populations and reports do indicate that infestations are more damaging in
such situations than under double cropping (Li 1982). Similarly, retrenching from two crops per
year to one would be expected to result in lower densities of specialist pests and introducing an
upland crop before or after rice would augment the effect by breaking the life cycles of species like
yellow stem borer.
In general, all other conditions being equal, a given rice hectarage should result in the lowest
abundance of specialist pests when production is spread over the widest area possible: from the
perspective of crop production, the goal should be to "extensify" rather than intensify cultivation.
Fig. 6. The response of specialist pest populations to cropping intensity. Mean wet season maxima in kerosene
light traps installed at sites of IRRI’s Cropping Systems Program in the Philippines. A single rice crop per
year was grown in Cagayan and Pangasinan provinces under rainfed wetland conditions (n = 5 sites,
1979-80); two crops were grown under irrigated conditions in Nueva Ecija province (n = 6, 1981). The
means differ significantly (P < .05), save in the case of the green leafhopper.
In the major Asian rice-growing areas, the scope for drawing on these insights is constrained by a
host of social, economic and agronomic factors and the overall effects of any such change must be
carefully considered. The prospects may be brighter in upland or rainfed areas with a history of
diversified production and in those parts of Africa and Latin America where rice is expanding but
is not yet a dominant crop.
4.2 ASYNCHRONY AND ABUNDANCE

But what of the effect of asynchrony of cultivation on rice pest populations? For modern varieties
whose flowering is largely insensitive to photoperiod and, in the lowland tropics, to temperature,
variation in the time of planting results in equivalent variation in maturity. As noted earlier, the
consequent asynchrony of plant host availability both extends the period of increase for pests able
to disperse between fields and shortens the fallow they must endure. The evidence available
(Loevinsohn 1984; Loevinsohn and Litsinger, in press) bears on the former within-season effect.
Considering the dispersive (adult) phase of a species, increased populations in one season are
expected where the range of planting dates exceeds a generation length and where this variation is
LOEVINSOHN 505
encountered over a distance that adults can readily move. Beyond the threshold, the population
response to asynchrony will be related to the species’ intrinsic powers of increase and to the extent
of density-dependent constraints.
These hypotheses were tested in a 20,000 ha irrigated area in Nueva Ecija province,
Philippines, of almost uniform double cropping, planted to modern varieties of similar growth
duration. Significant relationships between seasonal abundance in light traps and asynchrony,
as measured by the standard deviation of planting dates within a given radius of the traps, were
obtained for 8 of the 9 species considered. This is illustrated for brown planthopper (Fig. 7),
whose numbers varied 28-fold among the sites, none of which were separated by more than 13
km. Results for the other species are presented in Table 3. Density-dependence in the response
to asynchrony is apparent for 5 of the 8 species, and the effect of variation in the time to maturity
of varieties in the vicinity in a further 2 cases. The one pest whose numbers appeared unrelated
to asynchrony, in either of the two seasons considered, was Cnaphalacrocis medinalis (Guenée),
a pyralid leaffolder.
Fig. 7. Seasonal abundance of the brown planthopper in relation to the asynchrony of planting within 2 km. Nueva
Ecija. Philippines; wet season 1981. At each site, 2 simple kerosene light traps were established in rice fields
and lit nightly by farmer cooperators. Standard deviations of planting date around each of the sites were
calculated within radii of 0.2 - 2.0 km, using maps and data furnished by the irrigation administration. For
the brown planthopper, the correlation between In abundance and asynchrony was maximized when the
latter was calculated within 2 km. In Y = 4.31 + .223 X (t = 3.13, n = 14 sites, P < .01).
TABLE 3
The response of rice pests to asynchrony and the ecological characteristics
conditioning that response over 14 sites, wet season, 1981
Best fit Response to Maxiumum increase c Best fitting Median dispersal

Species
ting arynchrony b (rate/mo.) radius d distance e
model a (rate/day) (km)
N. nigropictus 2 1.17* 2.37 1.4 na
(green leafhopper)
N. virescens 3 0.79* 2.10 0.6 na
(green leafhopper)
N. malayanus 1 0.32*** 1.86 2.0 na
(green leafhopper)
N. depunctalis 2 0.87 ** 1.92 0.4 na
(caseworm)
N. aenescens 3 0.64* 2.22 0.4 0.3
(green semilooper)
P. fluctuosalis 2 0.51* 1.78 0.4 0.6
N. lugens 1 0.22** 1.46 2.0 2.5
(brown planthopper)
S. incertuas 1 0.08* 1.13 0.4 0.6
(yellow stem borer)
a Determined by maximum R (correlated for df) in regressions of In season total on different independent variables: 1 -
standard deviation of planting date (SD); 2 - SD and SD; 3 - SQ, SD and mean maturity of varieties planted.
b
Coefficient of SD in best fitting model. * P < .05; ** P < .01; *** P <. 001 by one-tailed test.
c Exponential rate calculated from mean greatest proportional change between consecutive monthly light trap totals.
d
Radius within which SD calculated in best fitting model.
e Calculated from catches over several weeks in paired light traps placed upwind of isolated sources of naturally dispersing
insects: a 2 ha field planted out of season (species 5, 6,8; n = 16 traps) and an extensive area of wild and volunteer rice
cultivated for the first time in several years (species 7; n = 32 traps). These experiments took place during a fallow period
(January-February 1982) when there were no other significant areas of rice in the vicinity. Median dispersal distances were
computed from the integration and rotation of the best-fitting equations through 360 degrees, assuming, in other words,
that movement was similar in all directions. na - not assessed.
The details of the relationship between asynchrony and abundance agree with expectation in
three regards:
(1) The species' rate of response to asynchrony were significantly related to a measure of
their maximum realized rates of increase (r = 0.87, P < .01). Both parameters were
lowest in the case of yellow stem borer, whose generation length, 40-50 days, is substan-
tially greater than those of the other pests.
(2) The radius within which asynchrony is calculated that provides the best fit to seasonal
abundance differs among the species and appears to reflect relative validity. This radius
was consistent between seasons (r = 0.94, n = 5, P < .02) and was significantly
correlated (r = 0.98, P < .02) with the median dispersal distance of four species,
estimated independently.
LOEVINSOHN 507
(3) There was evidence of the expected threshold in population response to asynchrony. In a
dry season in which planting date variation was unusually low and at some sites insuffi-
cient for any of the species to complete an additional generation, the relationship between
asynchrony and seasonal abundance was initially flat and then curved upward for 4 of the
5 species considered. These results suggest that an ecologically significant degree of
synchronization is attainable in some areas under existing conditions without special
effort.
The results of this and the previous section demonstrate that changes in the duration of rice
cropping, reflected in reduced time between and increased time within seasons, affect rice specialist
pests in a broadly similar fashion. This suggests that efforts aimed at restricting the availability of
rice will have multiple benefits, a decided advantage in comparison with many control measures,
cultural, biological, or genetic, that are species-specific in their effect.
Dispersal range, generation length and rate of increase provide the spatial and temporal
criteria with which to predict the impact on pest populations of variation in the time of planting. In
intensive rice growing areas, problems in delivering water to or draining it from fields appear often
to be key in determining the magnitude and distribution of asynchrony. In Nueva Ecija, it was found
that most farmers in the normal course of affairs attempt to compensate for delays imposed by the
centrally-managed irrigation system by hastening farming operations under their own control, such
as land preparation and transplanting (Loevinsohn 1984). There, as elsewhere, farmers’ efforts to
limit asynchrony can be supported by appropriate irrigation scheduling and correction of system
faults. The above results provide a basis for setting realistic targets: there is no justification in terms
of pest suppression for attempting to complete planting within much less than a pest’s generation
length over an area much larger than its dispersal range. Imposition of such standards could harm
both farmers and agricultural laborers by, for example, driving up tractor hire rates and encourag-
ing the shift from transplanting to direct seeding.
While there have been scattered attempts at synchronizing cultivation, the contrary, pur-
poseful staggering of planting within a farmer’s holding, has also been promoted. The ad-
vantages are that seasonal peaks in input demand and slumps in harvest price are avoided, while
the risks of catastrophic loss are minimized, for example, due to typhoons. One such approach,
known as the “rice garden” (Morooka et al., 1979), involves the division of a farm into plots, one
of which typically is planted and another harvested each week. In this way, up to four harvests
a year are obtained and asynchrony is maximized. Adoption of the rice garden has been limited
as few farmers have access to or can afford the continuous irrigation that it demands. Concern
has also been raised about the potential for pest build-up, to which the above results lend
considerable support.
Pantua (1979) however, found no consistent difference in infestations between a 2 ha rice

garden and an adjacent double cropped field. While a number of points can be made about the
manner in which the comparison was made, the most important one to consider is the question of
scale. The dispersal estimates reported above indicate that the majority of insects landing in a 2 ha
field would have originated from outside it and thus density would be affected by factors of host
plant abundance over a much wider area. The experience of an isolated, early- adopting farmer
would be very different from one who takes up the practice well after his neighbors. (Recall that
similar advantages accrue to early adopters of altered planting dates and double cropping.) Where
the rice garden has been implemented over a wider area, as on some corporate farms on the island
of Mindanao, Philippines, serious losses have been reported due to brown planthopper, and green
leafhopper and the viruses they vector.
4.3 ADAPTING TO THE AVAILABILITY OF RICE

The characteristics that determine species’ responses to factors of rice availability such as
asynchrony and cropping intensity, generation length, fertility, survival and dispersal range, are not
constants but vary among individuals. Efforts to manage the environment so as to limit the popula-
tion that can develop, whether by synchronizing cultivation or by restricting the intensity of crop-
ping, can be expected to create counteracting selection pressures favoring individuals less affected
by these measures. As variation in the above characteristics is likely to have at least some heritable
component, evolution would with time give rise to populations with at least partial resistance to such
cultural controls.
Evidence for processes of this sort can be found in populations inhabiting neighboring areas
differing naturally in synchrony or cropping intensity (Loevinsohn 1984). In Mapalad, Nueva Ecija,
Philippines, at the base of the Sierra Madre Mountains, a single crop of rainfed rice is grown in a
relatively synchronous fashion. The area lies about 10 km from the irrigated Central Luzon Plain
where double cropping predominates (it was here that the study on pest abundance in relation to
asynchrony was conducted). In the vicinity of Zaragoza, in the center of the plain, planting is highly
asynchronous due to irrigation and drainage faults.
Egg masses were collected from yellow stem borer moths at both locations and the newly
hatched larvae reared on caged plants at Zaragoza under conditions of constant density. The
duration of the larval and pupal stages, survival over this period and maximum fecundity of these
offspring are presented in Figure 8 and Table 4. Significant differences in all three parameters are
found insects from Mapalad emerged sooner, were more fecund but had poorer survival than those
from Zaragoza. Calculations suggest that under conditions of relative synchrony, where the length
of a season is most reduced, a papulation with the Mapalad characteristics would have the greater
rate of increase and attain a larger size by harvest. The Zaragoza parameters, on the other hand,
would prove advantageous where the season is more drawn out and where the pressure exerted by
natural enemies is high. Predators, parasites and pathogens are expected to be more abundant in
asynchronous, double cropped environments, like Zaragoza, where, as we have already seen, their
insect hosts are more readily available. The differentiation that is observed between the two sites in
bionomic parameters of the yellow stem borer thus appears to be in the direction that natural
selection would be expected to work.
LOEVINSOHN 509
DAYS AFTER INFESTATION
Fig. 8. Differentiation of generation length at sites of contrasting availability of rice. Time to adult emergence of
newly hatched yellow stem borer on caged rice plants. A synchronously planted single rice crop is grown at
Mapalad (n = 32); Zaragoza (n = 49) is an area of asynchronous double cropping. See also Table 4.
TABLE 4
Adaptation of yellow stem borer to the availability of rice a
Site Time to emergence Total oocytes in ovary Survival rate

(days) (number) (%)
Single crop, synchronous 38.5 a 269 a 11.3 a
(Mapalad)
Double crop, asynchronous 42.5 b 234 b 18.6 b
(Zaragoza)
a Life history parameters were determined by infesting newly hatched larvae on caged rice plants (IR36) at the beginning
of the 1981 wet season. Means in a column followed by different letter differ at P < .05.
Our confidence in this interpretation is increased by the results of a second experiment, similar
to the first, but that involved stem borers collected from three sites. Here, insects originating from
Mapalad and Zaragoza were compared to those from Ibabaw bana, lying within the area of double
cropping but benefiting from relatively problem-free irrigation and experiencing asynchrony com-
parable to Mapalad’s. Conditions of rice availability there are thus intermediate between those of
the original sites. As Table 5 indicates, parameters of the Ibabaw bana population are also
intermediate. The shorter mean times to emergence of both Mapalad and Ibabaw bana are an
effective adaptation to a short rice season (determined by asynchrony). Survival rates are similar at
the two sites that are double cropped, though if these do in fact reflect adaptations to natural enemy
pressure, a somewhat lower value might have been expected at Ibabaw bana where asynchrony is
less than at Zaragoza. Unfortunately, due to equipment failure, it was not possible to measure
fecundity in this experiment and to ascertain whether Ibabaw bana’s relatively short generation
length and high survival are purchased at the expense of reduced egg production.
TABLE 5
Adaptation of yellow stem borer to the availability of rice a
Site Time to emergence (days) Survival rate (%)

Single crop, synchronous 44.4 a 11.8 a
(Mapalad)
Double crop, synchronous 45.1a 17.6b
(Ibabaw bana)
Double crop, asynchronous 49.1b 17.2b
(Zaragoza)
a The experiment was similar to that reported in Table 4 but employed the variety IR42 and was carried out toward the end
of the 1982 wet season. Means in a column followed by different letter differ at P < .05.
The conditions of cropping intensity and asynchrony that distinguish the three sites date from
the introduction of widespread irrigation in the mid-1970’s. Thus, within a period of 6 years (at the
time of the study), significant genetic differentiation had already taken place in those parameters
and in those directions which would tend to counteract the change. Yet this evolution has not been
sufficient to eliminate the differences in abundance due to rice availability: catches of yellow stem
borer in light traps were 3.2 times greater at Zaragoza than at Ibabaw bana in the 1981 wet season.
Whether further evolutionary change is possible that would reduce this difference or whether a
“design limit” has been reached can only be determined by follow-up studies in these same areas.
Answers to such questions are key to an assessment of the long term durability of cultural control
measures based on restricting the availability of rice to its pests.
5. CONCLUSIONS
In considering the environments that rice pests inhabit, we have focused on a limited number of
characteristics, in essence, those that determine the spatio-temporal availability of the rice host. The
variability of these factors over space and their change over time is mirrored in a hierarchy of
responses at the level of pest populations, in their dynamics and genetics, and in the rising and
declining dominance of species. The processes at work operate at characteristic scales, often
greater than those considered in most entomological experiments. Taking the measure of effects
LOEVINSOHN 511
acting over kilometers and over seasons will in many cases require rethinking and refining accepted
methods. It is vital that we accustom ourselves to thinking in terms of such processes, including pest
evolution, that too often appear to take us by surprise. For the most part, variation and change in
agricultural systems has tended to act on pests and diseases; return flows, in the form of design and
management options based on an understanding of the effects, have been very weak. It is in
developing these options and elaborating a preventive pest management, that we encounter one of
our greatest challenges.
ACKNOWLEDGEMENTS
The comments and assistance of J.A. Litsinger, D. Catling, M. Austria-Alvarez and A. Barrion are
gratefully acknowledged.
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Landbouw. 20: 465-94.
III. CONTROL TACTICS
AND STRATEGIES
5
Host Plant Resistance
E. A. H EINRICHS
Baton Rouge, Louisiana
1. INTRODUCTION
RICE IS THE WORLD'S MOST important food crop as based on cultivated area. Rice serves as a major
source of calories for 40% of the world's population (De Datta 1981). Most of the rice is produced
in tropical Asia where the human population is high and where per capita income and rice yields are
low. Among the numerous abiotic and biotic stresses that affect rice yields, insect pests and the
plant pathogens that they vector, are major factors. Estimates in Asia by Cramer (1967) and Pathak
and Dhaliwal (1981) indicate that insects are responsible for more than 30% yield losses. A 10%
increase or decrease in food grain production on a global scale can make the difference between a
glut and acute scarcity (Swaminathan 1983). Because of the high yield losses national rice research
programs throughout the world are in various phases of developing and implementing management
strategies to minimize the losses caused by insects. Management strategies consist of various mixes
of chemical control with insecticides, cultural controls, biological agents (parasites, predators and
diseases) and planting of cultivars that have genetic resistance to insects. Resistant cultivars are
sought as the major tactic in an integrated approach to rice insect control. The incorporation of
insect resistance into modern cultivars is a major objective of most rice breeding programs in Asia
(Hargrove and Cabanilla 1985) (Fig. 1) for several reasons (Heinrichs et al., 1985a):
Income per hectare in rice production is relatively low and money spent for controls such as
insecticide significantly lowers profits. Resistance in rice cultivars is essentially free to the
farmer.
Insecticide prices are increasing faster than the rice price in most countries.
518 HOST PLANT RESISTANCE
Fig. 1. Rice breeding objectives in ten Asian countries in 1984

(modified from Hargrove and Cabanilla 1985).
• Land holdings of most rice farmers are small. Maintaining insecticide application equipment
is not economical on many small farms.
• Prophylactic or calendar based applications of insecticides are often wasted because
farmers do not know how to apply insecticides according to economic thresholds.
• Insecticides cause accidental poisoning incidents.
• Insecticides may pollute the environment killing fish, and livestock.
• Varietal resistance is generally compatible with other control tactics: chemical control,
cultural control, and biological control.
Because of its unique advantages host plant resistance is often utilized as a key tactic in the
integrated control of rice insect pests. Insect resistant rice cultivars are of special value under the
conditions existing in most developing countries and as such have become a component of rice
breeding programs. Teams of entomologists working with plant breeders and other problem area
scientists throughout the rice growing regions of the world have been, for the most part, successful
in developing rice production systems with an increased level of resistance/tolerance to abiotic and
biotic stresses. This is of extreme necessity as stresses are severe constraints to scientists’ attempts
to increase the profitability of agricultural production (Heinrichs 1988a). The progress achieved in
the identification of insect resistant rice germplasm and its use in the breeding of high yielding
cultivars with resistance to multiple stresses has been an integral part of the success of the green
revolution and has increased the profitability of rice production and minimized safety risks to
farmers, and contributed to a more healthful environment.
In the last three decades there has been a dramatic shift away from the planting of traditional
rice cultivars and the use of traditional cultural methods for insect control to the growing of modern,
high-yielding rice cultivars with an emphasis on insecticides and plant resistance. As a result, yields
HEINRICHS 519
have in many cases increased but yield stability, with the unilateral use of insecticides, or resistant
cultivars, has not been achieved (Heinrichs and Mochida 1984, Heinrichs 1988b). Because of the
short life of control programs based on one control tactic, there has been a significant interest
among rice scientists in the development and implementation of programs that integrate a multiple
tactic approach to rice insect control (see Chapters 10, 11, 12 and 13).
2. ROLE OF HOST PLANT RESISTANCE IN RICE IPM

Professor Chiu Shin-Foon of the South China College of Agriculture in a meeting of rice scientists
in Guangzhou in 1980 reported that “the breeding of resistant varieties has been the missing link’ in
the integrated pest control in China for some time but resistance breeding has recently been
initiated” (Chiu Shin-Foon 1980). The development of insect resistant cultivars has proceeded at a
rapid pace in China and throughout the world and the planting of insect resistant cultivars is being
integrated with well established cultural, chemical and biological control methods.
Resistant cultivars alter the physiology and behavior of rice insects which in turn affects the
insects’ susceptibility to chemical and biological controls. The adverse effect on the physiology of
the insect is evident in the work of Padgham (1983) where he reported a decrease in the survival and
migration potential of brown planthoppers Nilaparvata lugens (Stål) on resistant cultivar IR36 as
compared to those on susceptible IR20. Because of the decreased level of lipid synthesis when
feeding on IR36 only 27% of the insects had a flight capacity of greater than 10h whereas 74% of
those reared on IR20 had a comparable flight potential (Fig. 2).
Fig. 2. Distribution of potential flight durations of the brown planthopper Nilaparvata lugens developing on
mature plants of susceptible IR20 or resistant IR36 rice cultivars (after Padgham 1983).
2.1 HOST PLANT RESISTANCE AND CULTURAL CONTROL

Agronomic practices that are used in rice production systems to evade insect populations are the
synchronous planting of fields in a given area and the planting of early maturity cultivars (Heinrichs
1988c). Synchronous planting of rice within an area related to the effective dispersal of the brown
planthopper and yellow stem borer Scirpophaga incertulas (Walker) decreases the pest load on the
crop (Loevinsohn et al., 1988). In Indonesia, brown planthopper populations in areas synchronously
planted were one fourth that of areas where planting was staggered (Oka 1979). The combined
effect of resistant cultivars and synchronous planting significantly decreases pest populations.
Early maturing rice cultivars evade the later generations of multiple generation pests. They
become unsuitable for feeding before the brown planthopper reaches the damaging third genera-
tion (Heinrichs et al., 1986a). Early maturity complements host plant resistance and other cultural,
chemical and biological controls.
2.2 HOST PLANT RESISTANCE AND BIOLOGICAL CONTROL

Resistant cultivars have an indirect adverse effect on natural enemies because of lower prey density.
They have also been shown to adversely affect the development of parasites and predators by
decreasing the suitability of the prey (insect pest) as a food source (Orr and Boethel 1986).
However, resistant cultivars and biological controls are generally considered to be compatible.
Resistant cultivars have a major impact on conserving natural enemies by decreasing the insecticide
load.
Fig. 3. Brown planthopper (Nilaparvata lugens) /spider (Lycosa pseudoannulata) ratios at 40 days after transplant-
ing of rice cultivars with varying levels of N. lugens resistance. IR36, IR42, and ASD7 are resistant, Triveni
is moderately resistant, and IR8 and TNl are susceptible (modified from Kartohardjono and Heinrichs
1984).
HEINRICHS 521
The activity of certain natural enemies is enhanced when preying on insects that are feeding on
resistant cultivars. Resistance increases the predation rate of the spider Lycosa pseudoannulata
(Boesenberg et Strand) on the brown planthopper (Kartohardjono and Heinrichs 1984). Apparent-
ly, the restlessness and subsequent movement of the brown planthopper on the resistant cultivar
causes it to be detected by the spider. Brown planthopper: spider ratios in the field are most
favorable for biological control activity on resistant cultivars (Fig. 3). Also, the combined effect of
predation by a mirid bug Cyrtorhinus lividipennis Reuter and cultivar resistance significantly
reduces brown planthopper populations more than either the predators or cultivar resistance alone
(Kartohardjono and Heinrichs 1984).
The combination of varietal resistance and predation by C. lividipennis has a cumulative effect
on green leafhopper Nephotettix virescens (Distant) control (Myint et al., 1986). N. virescens mor-
tality when feeding on the resistant cultivar IR29 was 66% but increased to 92% with the addition
of C. lividipennis. On susceptible TN 1, N. virescens mortality was only 41% with C. lividipennis
predation.
2.3 HOST PLANT RESISTANCE AND CHEMICAL CONTROL
Insecticides provide more effective control when insects are feeding on resistant plants. White-
backed planthoppers Sogatella furcifera (Horvath) and brown planthoppers feeding on resistant or
Fig. 4. LD50 of insecticide applied topically to the whitebacked planthopper Sogatella furcifera reared on suscep-
tible TN 1 and moderately resistant N22 rice cultivars (modified from Heinrichs et al., 1984).
Fig. 5. Contact toxicity of insecticides applied with a Potter’s spray power against the brown planthopper Nilapar-
vata lugens biotype 2 when reared on susceptible (IR26) and moderately resistant (ASD7) rice cultivars
(modified from Heinrichs et al., 1984).
moderately resistant rice cultivars are killed with lower insecticide rates than those feeding on
susceptible cultivars (Heinrichs et al., 1984). LD50 rates of insecticides applied topically to the
whitebacked planthopper were two to three times as high on susceptible TN 1 than moderately
resistant N22 (Fig. 4). Biotype 2 brown planthoppers reared on moderately resistant ASD7 had
mortality rates 2-12 times that of brown planthoppers reared on susceptible IR26 when sprayed with
various insecticides (Fig. 5). Thus insecticide rates required for adequate planthopper control are
lower on a moderately resistant cultivar than that on a susceptible cultivar.
Host plant resistance and insecticides have a cumulative effect in the control of the green
leafhopper and the tungro virus that it vectors (Heinrichs et al., 1986b). Green leafhopper resistant
cultivars have a low incidence of tungro and susceptible cultivars have a high incidence regardless
as to the number of insecticide applications (Fig. 6). However a cultivar such as IR36 that has
moderate resistance to the vector shows a response to insecticides in that treated field plots have a
decreased tungro infection percentage and untreated plots have a high incidence of tungro infected
plants.
Host plant resistance can mitigate the degree of brown planthopper resurgence when rice fields
are treated with resurgence-inducing insecticides. Brown planthopper populations on resistant
plants treated with a resurgence-inducing insecticide were only 10 per hill whereas there were 1,100
hoppers per hill on treated susceptible cultivars (Aquino and Heinrichs 1979). In fields where the
leaffolder or other insects regularly reach the economic injury level and require insecticide treat-
ment, brown planthopper resurgence can be minimized by planting a brown planthopper resistant
cultivar.
HEINRICHS 523
- -
No of diozinon applications
Fig. 6. Green leafhopper Nephotettix virescens mortality (A), and field populations (B), rice tungro virus infec-
tion (C), and rice grain yields (D) as affected by number of diazinon applications at 1.5 kg (AI)/ha on
N. virescens susceptible rice cultivar IR22, moderately resistant IR36, and resistant IR28 (modified from
Heinrichs et al., 1986b).
Decreasing the amount of insecticide applied to rice through the planting of insect resistant
cultivars conserves natural enemy populations, preserves the quality of the environment, slows the
rate of selection for insecticide resistant insect strains and increases the profitability of rice
production. The integration of host plant resistance with insecticides, cultural controls and biologi-
cal control agents in a crop production system is more effective, environmentally safer, economical-
ly more profitable, and more stable than complete reliance on insecticides to control insects on
susceptible cultivars.
2.4 INSECT RESISTANT CULTIVARS AND CONTROL OF NON-INSECT PESTS

There are a myriad of interactions among the various pests (insects, diseases, nematodes, rodents,
and weeds) that attack rice plants. The various pests, abiotic environmental factors, and the rice
plant are interdependent and linked. Thus the control of a target pest may cause population
increases, or decreases, or may not affect another pest (Reissig et al., 1986).
Weeds and plant diseases may be adversely affected by the planting of insect resistant cultivars.
Insects that feed on susceptible seedlings and plants in the vegetative stage slow plant growth and
reduce the plant foliage area. This causes the closing of the plant canopy to be delayed and reduces
the competition for light and nutrients which in turn allows greater weed growth. Insect feeding
predisposes rice plants to attack of bacterial and fungal pathogens that invade the insect-damaged
plant tissue. Some insect species, especially the leaf- and planthoppers, are rice virus vectors.
Decrease in the feeding activity on non-preferred rice cultivars decreases the incidence of virus
infection.
3. COLLECTION, EVALUATION, AND UTILIZATION OF

INSECT RESISTANT GERMPLASM
3.1 PRESERVING RICE GERMPLASM

Since antiquity rice has been cultivated under a multitude of diverse growing conditions in respect
to elevation, climate, and soil types. The ecogenetic differentiation of rice occurred primarily
through natural and human selection in these different environments, following hybridization,
segregation or mutation (Swaminathan 1984). Germplasm in Oryza is derived from three sources:
(1) wild species, natural hybrids between cultivated varieties and their wild relatives and primitive
cultivars, (2) commercial, obsolete, and special purpose cultivars, and (3) cultivars arising from
human intervention, whether unwittingly, or intentional.
It has been estimated that the number of rice cultivars in collection may reach 120,000. Starting
in the 1930s every major rice-producing country in Asia began collecting germplasm from in-
digenous rices (Chang 1984). The size of national collections varies from several hundred cultivars
in Nepal to about 40,000 in China (Table 1). India has about 35,000 cultivars and the U.S. more than
18,000. Although the commercial cultivars are well represented in these collections few wild rices
have been collected and few primitive cultivars and breeding lines have been maintained. Because
of a lack of refrigeration facilities many of the tropically stored collections have been eroded.
HEINRICHS 525
TABLE 1
Estimates of rice germplasm holdings in major centers and their storage conditions (From Chang 1985)
Number of samples in national center

Country/Center Total Indigenous Duplicate samples in Storage condition 2
IRGC1
– 5,285 5,285 MT
Bangladesh
Burma – 2,080 1,724 MT
China (mainland) 40,0003 33,000 3,767 ST, MT, LT
China (Taiwan) 6,5203 1,662 1,185 MT
India 35,0004 33,000 14,585 ST
Indonesia 7,500 – 7,840 MT
Japan 42,4555 16,200 978 MT, LT
– – 906 –
Kampuchea
Korea (S.) 4,227 1,027 1,023 LT
Laos – – 1,431 –
Liberia 409 – 1,159 MT
Malaysia 4,550 3,130 2,395 MT
Malagasy 2,000 2,000 218 ST
Nepal 780 – 1,247 ST
Pakistan 1,224 1,404 936 MT
– 1,187 3,954 ST
Philippines
Sri Lanka 2,145 2,745 2,069 MT
Thailand 6,000 – 2,906 MT, LT
U.S. 18,0656 92 4,511 MT, LT
U.S.S.R. 3,514 – 372 LT, MT
Vietnam – – 1,634 –
IITA 7,633 n/a 1,024 MT
IRAT 3,842 n/a 1,577 MT
1 IRGC = International Rice Germplasm Center (IRRI)
2 ST = Short term, MT = medium term, LT = long term
3 Including many duplicates; collections scattered over more than 10 institutions.
4 Including many duplicates; collections scattered over 70 institutions; 15,000 accessions at CRRI; 3,464 accessions
deposited at IRRI.
5 Including many duplicates; collections scattered over the national center (NIAR), more than 30 national and lesser
experiment stations, 12 universities, and the Institute of Genetics; 18,000 accessions at NIAR.
6 Including deposits by IRRI; more than 5,000 accessions are held at BARC, Beltsville under MT storage as a working
collection.
The International Rice Research Institute (IRRI) established a germplasm center in 1961. It is
a conglomerate of all of the national collections. According to its Director, Dr. T. T. Chang (pers.
communication), at the end of 1989 the IRRI International Rice Germplasm Center held 78,420
accessions of O. sativa, 2,408 strains of O. glaberrima, 2,214 wild species and species-hybrids, and
206 genetic testers and mutants. This collection makes IRRI the world’s largest repository for a
single crop. To provide extra security for the IRRI collection the institute deposits a duplicate set
of seed in the U.S. National Seed Storage Laboratory, Fort Collins. Colorado, USA. In cooperation
with the International Board for Plant Genetics Resources, IRRI is collecting and conserving the
remaining 30,000-40,000 cultivars and when this task is completed, rice may be the first major food
crop to be preserved in as complete a collection of naturally occurring genetic variability as can be
achieved (Swaminathan 1984).
3.2 EVALUATION OF RICE GERMPLASM FOR INSECT RESISTANCE

Collection of germplasm is a prerequisite for any crop improvement program. Because of the
abundance and diversity of rice germplasm the collections provide a rich source of insect resistant
material. The importance of interdisciplinary cooperation in a typical insect resistance breeding
program is illustrated in Figure 7. The role of the entomologist is to (1) develop screening methods,
(2) identify donor parents with insect resistance for use in the breeding program, (3) evaluate the
level of resistance in breeding lines produced, (4) identify the mechanisms involved in resistance,
and (5) collaborate with the breeder in identifying the genes imparting insect resistance.
Breeding for insect resistance has only been a focus of rice development programs since the
early 1960s. The discovery of high levels of resistance to leafhoppers and planthoppers in traditional
cultivars sparked a widespread interest in the development of high yielding, insect resistant rice
cultivars. As a result, screening methods for more than 30 rice insect species have been developed
(Table 2) (Heinrichs et al., 1985a).
TABLE 2
Status of screening and breeding for varietal resistance to rice insect pests (modified from Heinrichs et al., 1985a)
Insect Status
Common name Scientific name Screening Resistance Resistant Resistant Genes for
methods sources breeding lines varieties resistance
developed identified available released identified
Brown planthop- Nilaparvata † † † † †

Per lugens
Whitebacked Sogatella furicifera † † † † †
planthopper
Smaller brown Laodelphax † † † † –
planthopper striatellus
Rice delphacid Tapsodes † † † † –
orizicolus
Green leafhopper Nephotettix † † † † †
virescens
Zigzag leafhopper Recilia dorsalis † † † † †
White leafhopper Cofana spectra † † – – –
Blue leafhopper Empoascanara † † – – –
maculifrons
Rice striped borer Chilo suppressalis † † † † –
Yellow stem borer Scirpophaga † † † † –
incertulas
(Contd. )
HEINRICHS 527
Insect Status
Common name Scientific name Screening Resistance Resistant Resistant Genes for
methods sources breeding lines varieties resistance
developed identified available released identified
Sugarcane borer Diatraea † † † – –

saccharalis
African striped Chilozacconius † † – – –
stem borer
African white † † – – –
Maliarpha
stem borer separatella
Lesser cornstalk † † – – –
Elasmopalpus
borer lignosellus
– – –
African pink borer Sesamia † †
calamistis
S.A. white borer Rupelaalbinella † † – – –
Stalk-eyedfly Diopsis † † † – –
macrophthalma
Rice stem maggot Atherigonaoryzae † † – – –
Whorl maggot Hydrellia † † † † –
philippina
Gall midge Orseolia oryzae † † † † †
Rice seedling fly Atherigona exigua † † – – –
Armyworm Mythimna † – – – –
separata
Thrips Stenchaetothrips † † † † –
biformis
Rice bug Leptocorisa † † – – –
oratorius
Black bug Scotinophara † † † † –
latiuscula
Caseworm Nymphula † † – – –
depunctalis
Leaffolder Cnaphalocrocis † † † – –
medinalis
Rice water weevil Lissorhoptrus † † † – –
oryzophilus
Hispa Dicladispa † † – – –
armigera
Bloodworm Chironomus † † – – –
tepperi
Rice weevil Sitophilusoryzae † † – – –
Maizeweevil Sitophiluszeamais † † – – –
Lesser grain borer Rhyzopenha † † – – –
dominica
Angoumois grain Sitotroga † † – – –
moth cerealella
Fig. 7. Steps in the breeding for insect resistant rice cultivars at IRRI (after Heinrichs et al., 1985a).
HEINRICHS 529
Screening for rice insect resistance is being conducted throughout the world. Major emphasis
has been on the leafhoppers in Asia and stalk-eyed fly in Africa, rice delphacid in South America,
and rice water weevil in the U.S.A. Minor emphasis has been given to the rice storage insects.
Screening methods employed for the various rice insects throughout the world are described in
detail and sources of resistance identified are listed in Heinrichs et al. (1985a).
3.3 BREEDING INSECT RESISTANT RICE CULTIVARS

Successful rice breeding programs consist of an interdisciplinary approach where scientists from
several key disciplines work together as a team. This is because the degree to which a new cultivar
is a success depends on a number of factors involving many disciplines. Thus, entomologists, plant
pathologists, physiologists, agronomists, cereal chemists, soil scientists and biotechnologists must
work closely with the plant breeders. Such rice breeding programs have resulted in the development
of insect-resistant cultivars with high yield potential that are being utilized as a major tactic of
integrated pest management in rice production systems in many countries.
3.3.1 Green Leafhoppers, Nephotettix spp.

Nephotettix virescens (Distant) occurs in South and Southeast Asia, southern Japan and Taiwan. It
is of major importance because it is an efficient vector of rice tungro virus. Because of the lack of
virus resistance in rice, insect resistance has been employed as a means of controlling tungro virus.
All of the IR cultivars except IR22 have moderate to high levels of resistance to N. virescens and all
have varying levels of resistance to N. nigropictus and N. malayanus (Heinrichs et al., 1985b). Many
countries are growing rice cultivars with resistance to the Nephotettix species. Genetic analysis has
identified eight genes (Glh 1, Glh 2, Glh 3, glh 4, Glh 5, Glh 6, Glh 7, Glh 8) that impart resistance
to N. virescens (Athwal et al., 1971, Siwi and Khush 1977, Karim and Pathak 1982, Ghani and Khush
1988). Many of these genes provide cross resistance to N. nigropictus and N. malayanus.
3.3.2 Zigzag Leafhopper, Recilia dorsalis

The zigzag leafhopper is a minor pest but occasionally reaches populations of sufficient magnitude
to directly damage the rice crop through the removal of plant sap. Breeding for resistance to this
insect has been limited. IR56 among the IR cultivars is resistant to this insect (Heinrichs et al.,
1985b). Three genes have been identified for zigzag leafhopper resistance: Zlh 1, Zlh 2, and Zlh 3
(Angeles et al., 1986).
333 Brown Planthopper, Nilaparvata lugens

Because of the importance that the brown planthopper has assumed as a pest in the last three
decades, all of the major rice producing countries in South and Southeast Asia have breeding
programs to control this pest. Distinct differences in levels of resistance to the brown planthopper
under greenhouse conditions were first observed at IRRI in 1963 (Pathak et al., 1969). More than
50,000 accessions have been screened for resistance to this pest at IRRI and more than 400 resistant
accessions have been identified (Rapusas and Heinrichs 1987a). Biotype development has compli-
cated breeding for resistance to the brown planthopper. Sources resistant in one region may be
susceptible in another (Heinrichs 1988b). The latest IR cultivars are resistant to the three biotypes
in the Philippines (Heinrichs et al., 1985b)
Seven genes have been identified that impart resistance to the brown planthopper; Bph 1, bph
2, Bph 3, bph 4, bph 5, Bph 6 and bph 7 (Athwal et al. 1971, Lakshminarayana and Khush 1977, Khush
et al. 1985, Kabir and Khush 1988). IR26, the first IRRI cultivar with brown planthopper resistance
resulted from a cross made in 1969, of IR24 and TKM6, the latter having the Bph 1 gene for
resistance. IR26 has multiple resistance to several major diseases, and insects, good grain quality,
and high yield potential (Khush 1977). Breeding programs for brown planthopper resistance have
been established in most of the Asian countries and numerous cultivars have been released since
that of IR26 in 1973. Brown planthopper resistance has been incorporated into the hybrid cultivars
being grown in China. IR36, the most widely grown rice cultivar has brown planthopper resistance
from the recessive gene, bph 2 (Khush 1977).
3.3.4 Whitebacked Planthopper, Sogatella furcifera

The whitebacked planthopper is an important pest in the tropics and subtropics in Asia. Removal
of plant sap by the insect causes 'hopperburn', a drying of the plant. Of about 50,000 accessions
screened at IRRI, more than 400 have resistance to the whitebacked planthopper (Romena et al.,
1986). Genetic analyses of selected accessions have resulted in the identification of five genes for
resistance; Wbph 1, Wbph 2, Wbph 3, Wbph 4, and Wbph 5 (Sidhu et al., 1979, Angeles et al., 1981,
Hernandez and Khush 1981, Wu and Khush 1985). Breeding for resistance is being conducted at
IRRI and has resulted in the release of moderately resistant cultivars IR48, IR52, IR60, and IR62
(Heinrichs et al., 1985b).
3.3.5 Rice Delphacid, Tagosodes orizicolus

The rice delphacid is a serious pest of rice in South and Central America, and in the Caribbean. In
addition to causing direct damage, by feeding, it vectors the hoja blanca virus, the most serious rice
disease in the Western Hemisphere (Khush 1977). In 1967 scientists at the Centro International de
Agricultural Tropical (CIAT) in Colombia observed that plants in field plots of the Asian cul-
tivar,IR8, were resistant to T. orizicolus. Further evaluation of 5,000 cultivars under greenhouse
testing at CIAT identified 100 from Asia, where the insect does not occur, as resistant (Jennings and
Pineda 1970). The cultivars CICA 4 and CICA 6 released jointly by CIAT and the Institute
Colombiano Agropecuaria (ICA) were resistant to T. orizicolus. Resistance to T. orizicolus is due
to tolerance and no major genes have been identified. Resistant cultivars are playing a major role in
the control of T. orizicolus and hoja blanca.
3.3.6 Smaller Brown Planthopper, Laodelphax striatellus

The smaller brown planthopper occurs in Japan, Korea and China and is a vector of rice stripe, a
serious rice disease in East Asia, and a vector of black-streaked dwarf virus. Commercial cultivars
with resistance to this pest have been released. ASD7 has been reported as highly resistant in Korea,
HEINRICHS 531
Murunaga 137 and Pannetti are resistant in Taiwan and Tadukan is resistant under field conditions
in Japan (Khush 1977).
3.3.7 Rice Striped Stem Borer, Chilo suppressalis

Of about 20 rice stem borer species, the rice striped borer and the yellow stem borer Scirpophaga
incertulas occur most commonly in Asia and cause serious yield losses (Chaudhary et al., 1984). The
rice striped borer is the most widely studied rice stem borer species and occurs in East Asia,
Southeast Asia, South Asia, Egypt and Japan. It occurs in both the tropics and subtropics while the
yellow stem borer is primarily a tropical species.
Extensive screening for resistance to the striped borer under natural field infestations has been
conducted in India, Japan, and at IRRI (Khush 1984). High levels of resistance have not been found
and resistance is of a polygenic nature. Breeding for resistance at IRRI started in 1965, using as a
donor TKM6, the best source of resistance at the time. In 1968, a breeding line IR532E576 was
identified as resistant and it was released as IR20 in 1969 (Chaudhary et al., 1984). Since then,
several striped stem borer resistant cultivars with better agronomic qualities than IR20 have been
released by IRRI (Heinrichs et al., 1985b). Of the IR cultivars, IR36 has the highest level of
resistance. Cultivars such as Ratna have been developed in India from TKM6 crosses.
3.3.8 Yellow Stem Borer, Scirpophaga incertulas

The yellow stem borer has become the dominant rice borer species in several regions. The breeding
program for yellow stem borer resistance began in India in 1964 when TKM6, CB1 and CB2 were
used as donors and crosses have been made with high yielding semi-dwarf cultivars and local
improved lines (Roy et al., 1969, 1971). None of the lines developed had combined high levels of
resistance to the borer and high yields.
Evaluation of the germplasm collection for resistance to the yellow stem borer at IRRI started
in 1962 (Rapusas and Heinrichs 1987a). About 17,000 accessions have been screened. Screening is
conducted in a screenhouse and in field plots where stem borer populations are high (‘hot spots’)
(Heinrichs 1980, Heinrichs et al., 1985a). Only about 40 O. sativa accessions (0.1 % of the accessions
tested) and 80 wild rice accessions (18% of the accessions tested) have been identified as having
good levels of resistance to the yellow stem borer (Rapusas and Heinrichs 1987a). Several O. sativa
accessions from India such as CO 7, CO 15, CO 18, CO 21, Ratna, TKM6 and WC 1263 were
reported as resistant (Chaudhary et al., 1984). A breeding line, IR 1820-52-2, was found to have a
higher level of resistance than previously identified donors. Wild rice species with resistance
included O. eichingeri, O. punctata, and O. latifolia from Africa, O. minuta from the Philippines, and
O. officinalis from India.
Breeding for yellow stem borer resistance at IRRI began in 1972. IR20 was the first moderately
resistant cultivar released. IR36 and IR40 were released as moderately resistant cultivars in 1976.
Later releases having moderate resistance were IR50 and IR54 (Heinrichs et al., 1985b). Because
of a lack of high levels of resistance in commercial cultivars, a male-sterile facilitated recurrent
selection approach was utilized to breed for resistance (Chaudhary et al., 1981). Genetic male-
sterile IR36 as a female parent was crossed with 26 donor parents including CO 18, CO 21, IR
1820-52-2, TKM6, and WC 1263. Twenty lines with a high level of resistance were selected and these
are being used as sources of resistance in the breeding program.
3.3.9 Gall Midge, Orseolia oryzae
The Asian gall midge, Orseolia oryzae Wood-Mason is a serious pest in India, Indonesia, Sri Lanka
and Thailand. The larval stage damages the plant by causing the leaf sheath to produce a gall called
a ‘silver shoot’ that prevents panicle production. An early observation on differences in cultivar
reactions to the gall midge was reported by Nguyên (1922) in Vietnam and Hegdekatti (1927) in
India. Rice improvement programs in Bangladesh, India, Indonesia, Sri Lanka, and Thailand screen
for resistance to the gall midge and about 200 resistant cultivars have been identified (Heinrichs and
Pathak 1981).
Most of the resistant cultivars identified in screening programs are from India and a few
originate in Thailand. Field screening of the 3,600 cultivars of the National Collection of Rice at the
Central Rice Research Institute (CRRI) in India began in 1948. The Indian cultivars Ptb 18 and
Ptb 21, and the Thai cultivar Leuang 152 were selected. Later screening at Warangal, India, resulted
in the selection of Eswarakora, HR42 and HR63, and the Thai cultivar Siam 29. Between 1968 and
1970 the All India Coordinated Rice Improvement Project (AICRIP) coordinated the screening of
accessions from India and IRRI. Two cultivars from the Jeypore Botanical Survey and 44 Assam
Rice Collection (ARC) cultivars were selected. Of the 3,804 accessions from IRRI, Ptb 10, AC 1423,
DNJ45, and DV 12, originally from India, were resistant. Cultivars identified as resistant in India
have been evaluated in several national programs through the coordination of the IRRI Internation-
al Rice Testing Program. The development of means of mass rearing the gall midge has accelerated
the screening program.
Studies on the inheritance of resistance in Indian cultivars indicated dominant genes GM-1 in
Usha, Samredhi, and BdG-1, and GM- 2 in Surekha and IET 6286 (Chaudhary et al., 1985). Breeding
for resistance is being conducted in India, Indonesia, Philippines (IRRI), Sri Lanka, and Thailand
(Heinrichs and Pathak 1981). Breeding was first conducted in India and the Indian program is the
most active. In the 1960s, crosses between Ptb 18 and Ptb 21 as resistant donors, and GEB24, then
the most popular local cultivar, produced several CR55 and CR56 selections with resistance to the
gall midge (Khush 1984). With the advent of the short-statured, high yielding cultivars such as IR8,
hybridization programs at CRRI and AICRIP used CR55, CR56, Eswarakora, Ptb 18, Ptb21, and
Siam 29 as donor parents. Selections from these crosses resulted in the release of the gall midge
resistant cultivars Shakti in Orissa State, Vikram in Karnataka, and Surekha in Andhra Pradesh.
Many of the gall midge resistant cultivars have multiple resistance to insects and diseases. The
Warangal selections, WC 1252, WC 1253, WC 1257 and WC 1263 are resistant to the gall midge,
thrips, stem borers and leafhoppers (Shastry et al., 1972). Utilizing Warangal cultivars as gall midge
resistant donors, RD4 and RD9 were released in Thailand. BG400-1 and BG276-5 from the
Batalagoda program have been released in Sri Lanka (Heinrichs and Pathak 1981).
HEINRICHS 533
At IRRI, crosses between multiple-resistant, improved-plant type breeding lines and Ptb 18,
Cr 94-13 (Ptb 18 x Ptb21), Eswarakora and Ob677 have been made. Commercial gall midge
resistant cultivars with multiple resistance to insects and diseases include IR36 released in 1976 and
the subsequent IR cultivars IR38, IR40, and IR42 (Khush 1984). Utilization of resistant cultivars in
combination with other control tactics in pest management programs is providing effective and
economic gall midge control.
4. MECHANISMS OF RESISTANCE
Breeding for resistance to rice insects has resulted in the release of commercial cultivars that have
successfully been integrated into crop management systems and have had a significant economic
impact on rice production. Plant resistance has been successful in spite of the fact that knowledge
of the nature of resistance in rice is limited. However, a more in-depth understanding of the factors
that cause resistance is needed as the information can be used to develop more efficient screening
programs and it provides guidance for the breeding program.
Resistance in plants is the result of a series of interactions between plants and insects that
influences the degree of establishment of insects on plants (Saxena 1986). Insect responses are
influenced by physical factors that interfere with insect vision, locomotion, feeding, mating, and
oviposition and chemical factors that alter the behavior or physiology of the insect.
Information on the physical and chemical factors that impart resistance to rice insects is
limited. Research has concentrated primarily on the green leafhoppers, brown planthopper, rice
striped stem borer, and the yellow stem borer.
4.1 GREEN LEAFHOPPERS

Kawabe (1985) studied thc mechanisms involved in the resistance of Japanese cultivars to Nephotet-
tix cincticeps. Mechanisms consisted of the three components of resistance; antibiosis, antixenosis,
(nonpreference) and tolerance (Fig. 8). Electronic monitoring of feeding behavior indicated that
the insect withdraws its stylets from the phloem sap soon after obtaining a taste. He suggested that
the phloem contained a biological or physiological mechanism such as a phytoalexin or possibly a
mechanical barrier that inhibits phloem sap ingestion.
Heinrichs and Rapusas (1990) used an electronic monitor and a video camera to study the
feeding behavior of N. virescens on resistant and susceptible rice cultivars. When on susceptible TN
1 most of the feeding time was spent ingesting phloem sap. On resistant cultivars xylem ingestion
was the main feeding activity although the stylets were able to reach the phloem. On moderately
resistant IR36 a small amount of phloem ingestion was observed but most of the time was spent
feeding on the xylem. Similar to Kawabe's study, there appeared to be some unidentified
mechanism in the phloem seive elements that prevents N. virescens from ingesting phloem sap. The
shift from phloem to xylem feeding on resistant cultivars is not understood. Auclair et a1. (1982)
speculated that the presence of a feeding deterrent adjacent to or within the phloem seive elements
or the formation of callose or slime plugs in the phloem prevents feeding. Cheng and Pathak (1972)
RICE PLANT FEEDING BEHAVIOR RICE GREEN LEAFHOPPER
Fig. 8. Mechanism of resistance to the rice green leafhopper Nephotettix cincticeps

(after Kawabe 1985).
stated that resistance to N. virescens is not due to mechanical factors that interfere with feeding but
that resistant plants lacked phagostimulants, possessed feeding deterrents or toxic materials or
lacked nutrients.
Application of the steam distillate extract of a resistant rice cultivar to susceptible plants
disrupted the normal feeding behavior of N. virescens. Reduced phloem feeding on extract-treated
plants was associated with a significant increase in probing frequency and an increase in the
duration of salivation and xylem drinking (Khan and Saxena 1985). Because phloem feeding is
considered necessary for N. virescens to transmit tungro virus (Heinrichs and Rapusas 1984a) N.
virescens resistant plants have low levels of tungro infection.
Plant age has a distinct effect on the levels of N. virescens resistance in rice (Rapusas and
Heinrichs 1982a, 1987b). Plants of the moderately resistant cultivar IR36 at 10 days after sowing
(DAS) were most preferred as compared to older plants. N. virescens survival, growth index and
adult weight were higher on 20 DAS plants than on plants at 40, 60, or 80 DAS. Phloem feeding was
highest on young plants and tungro virus transmission was related to extent of phloem feeding
(Fig. 9).
4.2 BROWN PLANTHOPPER

Resistance of rice cultivars to the brown planthopper has not been attributed to morphological or
anatomical characteristics of plants. Reduced feeding on resistant plants, however, has been
attributed to either a lack of phagostimulants or to the presence of antifeedants (Saxena 1986).
HEINRICHS 535
Fig. 9. Percentage of moderately resistant rice cultivar IR36 plants infected with tungro virus when two
viruliferous green leafhoppers Nephotettix virescens were released per plant of four plant ages and suscep-
tible TN 1 infected at 20 days after sowing. Columns with the same letter are not signifiantly different
(P<0.05; Duncan’s multiple range test). Bars indicate ± 1 Sd (after Rapusas and Heinrichs 1987b).
Brown planthopper resistance in the cultivar Mudgo was considered to be due to a low
concentration of amino acids, especially the sucking stimulant, asparagine (Sogawa and Pathak
1970). Later, Yoshihara et al. (1979, 1980) reported that soluble silicic acid and oxalic acid in
resistant rice plants served as sucking inhibitors. Using a brown planthopper resistant, Japanese
breeding line, Shigematsu et al. (1982) identified b sitosterol and other sterols in the phloem sap.
They reported that b sitosterol is a strong brown planthopper sucking inhibitor and Kaneda (1982)
reported that low asparagine content intensifies the inhibitory effect of b sitosterol.
Volatile chemicals extracted as steam distillates, have been shown to affect the behavior and
biology of the brown planthopper (Saxena and Okech 1985). The insect preferred susceptible plants
sprayed with its own extract over the same cultivar sprayed with the extract of a resistant cultivar.
Feeding activity was greater when feeding on a sucrose solution mixed with the extract of a
susceptible cultivar than that on a sucrose solution mixed with extracts of resistant cultivars. The
identity of the allelochemics was not determined but it is believed that the steam distillation method
employed extracts of essential oils, terpenoids, aldehydes, fatty acids, esters, waxes, etc.
Cook et al. (1987) suggested that the surface of the rice plant plays a role in food plant selection
by the brown planthopper. Surface waxes have been shown to affect brown planthopper behavior
(Woodhead and Padjham 1988). Reduced settling and probing of the plant surface after explora-
tion, and movement off of the stem on to the leaves results from chemical cues in the wax. Sensory
structures on the labial tip of the brown planthopper receive chemical stimuli during the exploration
of the plant surface that occurs before probing and this affects host choice. Chemical cues received
by the brown planthopper from the plant surface originate from the alkanes or carbonyl compounds
of the epicuticular wax and vary among cultivars. Analysis indicates that the chemical differences
between waxes are due to a higher ratio of long to short carbon chain components in resistant
cultivars. The enhanced surface activity and brown planthopper dispersal from IR46 as reported by
Woodhead and Padjham (1988) may account for the field resistance of this cultivar as reported by
Heinrichs (1986). This resistance mechanism is believed to be associated with minor genes which
could be exploited in breeding for polygenic resistance to the brown planthopper.
4.3 STEM BORERS

Various plant morphological, anatomical, biochemical, and physiological factors have been sug-
gested as imparting resistance to the rice striped borer and the yellow stem borer (Chaudhary et al.,
1984). Although, some of the reported factors are correlative with resistance, they may not actually
be resistance mechanisms.
Several plant morphological factors have been reported. Plant height, stem diameter and length
and width of the flag leaf were positively correlated with number of eggs laid by stem borer moths
(Patanakamjorn and Pathak 1967). In addition, resistant cultivars possessed tight leaf sheaths that
totally covered the internodes whereas susceptible cultivars had loose leaf sheaths that partially
covered the internodes. Tight leaf sheaths were believed to prevent newly hatched larvae from
feeding on the inner part of the leaf sheath before boring into the stem. A positive correlation
between stem size and borer infestation was reported (Jodon and Ingram 1948). Tall plants with
wide and long leaves and a large number of tillers appeared most susceptible (Israel 1967). Awnless
cultivars were more susceptible than those with awns (Shiraki 1917).
Anatomical studies of stem borer resistant cultivars indicated that the stems had four to five
layers of sclerenchymatous tissue that apparently offer resistance to larvae boring inside the stem
(Van and Guan 1959). Also, cultivars with a narrow stem lumen were less susceptible to borers.
Non-preference of stem borers for oviposition is considered to be at least partially of a
biochemical nature. A stem borer attractant, oryzanone (p-methylacetophenone) attracts oviposit-
ing moths and larvae by its odor (Munakata et al., 1959). Non-preference of the rice striped borer
for TKM6 is due to the presence of allomones which inhibit oviposition and adversely affect egg
hatching, larval survival, and larval development (IRRI 1978). The chemical has been identified as
a pentadecanal in a collaborative effort between IRRI and the former Tropical Development
Research Institute, London.
HEINRICHS 537
Silica content has been associated with resistance to rice stem borers. High amounts of silica in
rice plants adversely affected larval survival and reduced ’dead heart’ formation (Pathak et al.,
1971). Mandibles of larvae feeding on the cultivar Yabani Montakhab, with a high silica content,
were worn down, whereas those feeding on Sapan Kwai, with a low silica content, were normal.
Larvae with damaged mandibles have a low feeding efficiency and suffer high mortality.
Studies on the chemical bases of resistance to stem borers and to other rice pests, have received
relatively little attention and much is left to be discovered. A better understanding of the
mechanisms of resistance would allow a more directed program for the breeding of rice insect
resistance and may lead to the development of rice cultivars with more stable resistance to insect
pests.
5. VARIABLE RESISTANCE TO INSECTS IN RICE CULTIVARS

Insect populations have a wide range of genetic variability that maximizes their fitness in the
presence of genetic diversity of host plants. The widespread planting of one rice cultivar that has
been commonplace since the release of the ‘green revolution’ cultivars has significantly decreased
the genetic diversity of rice plants. As a result, certain rice insects have developed virulence and this
causes severe damage to cultivars that previously were resistant. Such apparently new forms of pests
have been termed ‘biotypes’ (Claridge and den Hollander 1983). The term ‘biotype’ refers to a
population of insects that is capable of damaging plant cultivars that are resistant to other popula-
tions of the same species. It is distinguished from other populations of the same species by parasite
ability (Heinrichs 1988b). Biotype selection enables an insect species to keep pace with the changes
in the defenses of its host plant that occur through natural selection and plant breeding programs
(Saxena and Barrion 1987). Biotypes of the rice leaf miner Agromyza oryzae (Munakata), gall midge
Orseolia oryzae (Wood-Mason), rice stem maggot Chlorops oryzae (Mats.), green leafhopper N.
virescens, and the brown planthopper N. lugens have been reported on rice (Saxena and Barrion
1987). Green leafhopper, brown planthopper, and gall midge biotypes have been a major constraint
to the breeding of resistant rice cultivars and have been extensively studied.
5.1 GREEN LEAFHOPPER

Allopatric populations of N. virescens with variable levels of virulence to differential cultivars were
reported by Karim and Pathak in 1979 when comparing reactions of seedlings to a Philippine N.
virescens population maintained in culture at IRRI, and a Bangladesh population maintained at the
Bangladesh Rice Research Institute. Cultivars with the Glh 1, Glh 2, Glh 3 or Glh 5 gene were
resistant at IRRI but susceptible at BRRI. Results of an international collaborative study conducted
a few years later indicated that Pankhari 203 (G1h 1 gene) was susceptible in Bangladesh, India and
Malaysia but was resistant in the Philippines, Thailand, and Vietnam (Heinrichs et al., 1986c).
Although N. virescens resistant cultivars with the Glh 3 gene were sequentially grown in the
Philippines for 20 years, evidence of an increase in virulence was not discovered until 1982 (Rapusas
and Heinrichs 1982b). IR36 was released in the Philippines in 1976 and subsequently became the
most widely grown cultivar. Results of a survey in the Philippines in 1979 indicated no evidence of
an increase of N. virescens virulence on the cultivar. However, in a survey conducted in 1984 N.
virescens populations of Luzon Island had levels of virulence significantly greater than those of the
greenhouse colony that had not been exposed to selection pressure (Rapusas and Heinrichs 1986).
Laboratory selection for virulence of N. virescens on resistant cultivars indicates an increase in
survival, decrease in the nymphal duration and a shift from xylem feeding to increased feeding on
the nutrient-rich phloem (Heinrichs and Rapusas 1984a). Increased phloem feeding results in
increased rates of tungro virus transmission (Heinrichs and Rapusas 1984b).
5.2 BROWN PLANTHOPPER

Variability in the levels of brown planthopper virulence on resistant cultivars has been a very severe
problem in South and Southeast Asia. IR26 with the Bph 1 gene was the first N. lugens resistant
cultivar released by IRRI in 1974. Although it was resistant in the Philippines it was susceptible to
the Indian N. lugens population indicating variable levels of virulence in allopatric populations.
Within the Philippines, N. lugens outbreaks were observed in IR26 after 2-3 years of commercial
cultivation (approximately 6 crops). Laboratory studies showed that high levels of virulence of N.
Iugens populations to resistant cultivars could be selected in 7 to 10 generations (Pathak and
Heinrichs 1982). N. lugens resistant cultivars released subsequently to IR26 have also succumbed to
biotype selection within a few years after release (Heinrichs 1988b).
Confirmation of the extreme variability of virulence patterns in the field was evident in the
result of the studies of Claridge et al. (1982). When six Sri Lankan N. lugens populations collected
from a wild rice species, traditional cultivars, and modern improved cultivars were tested for
virulence, they were shown to be most closely adapted to the cultivar, from which they were
collected, despite the fact that all populations were collected within a 200 km area. Grouping of
brown planthopper biotypes as based on the response of differential cultivars has indicated the
presence of several different biotypes in South and Southeast Asia (Heinrichs 1988b). Virulence has
been shown to be inherited as a polygenic or quantitative trait (den Hollander and Pathak 1981).
5.3 GALL MIDGE

The first evidence of differential reactions of rice cultivars to gall midge populations was reported
in comparing reactions in Nizamabad, South India with those of the same cultivars exposed to a gall
midge population at Cuttack, India, 1,200 km to the northeast (Israel and Vedamoorthy 1953).
Further studies at 11 sites in India confirmed the existence of gall midge populations with varying
levels of virulence on specific cultivars.
International collaborative studies among rice scientists in China, India, Indonesia, Sri Lanka
and Thailand have resulted in the classification of gall midge biotypes in South and Southeast Asia
(IRRI 1981). Testing of six cultivar groups against the various gall midge populations resulted in the
identification of eight different reactions. The populations were designated as the China, Indonesia,
Thailand, India-Raipur, India-Andhra Pradesh, India-Orissa, India-Bihar, and Sri Lanka biotype.
In spite of the presence of allopatric biotype populations increased virulence of gall midge popula-
HEINRICHS 539
tions through selection on commercially released, gall midge resistant cultivars, has not been
documented as it has for the brown planthopper.
6. RICE BREEDING AND GENE DEPLOYMENT STRATEGIES

The successful management of rice insect populations requires an integrated apprach in which
insect resistant cultivars are combined with other control tactics and other crop production prac-
tices. In spite of the development of virulent biotypes on previously resistant rice cultivars, plant
resistance to insects will remain a key tactic in rice insect management programs. To increase the
value of plant resistance several strategies have been developed to increase the stability of insect
resistant cultivars.
There is a great deal of genetic diversity in the world collection of rice. Because of a concerted
effort to collect and maintain the worlds rices at IRRI and in national program gene banks, this
diversity is available for use in rice breeding programs. Because of the tremendous amount of
diversity, several strategies are available to cope with the problems caused by biotype selection.
6.1 SEQUENTIAL RELEASE

Sequential release refers to a gene deployment strategy where a cultivar with a single major
resistance (R) gene replaces a cultivar with an R gene that has been overcome by the selection for
a virulent biotype. Replacement of IR26, a Bph 1 gene cultivar, with IR36, a bph 2 gene cultivar, is
such an example. This strategy requires a continuous search for new R genes. Genetic analysis of
the world collection of rices has been limited to a few insects and a limited number of cultivars and
there is much scope for identifying additional R genes, especially for leafhopper and planthopper
resistance. A prerequisite to the success of this strategy is an active and productive breeding
program to keep ahead of the biotype selection process.
6.2 GENE PYRAMIDING

Incorporation of two or more major R genes into the same cultivar using the pedigree method of
breeding is a strategy that is believed to be a means of increasing the stability of insect resistant
cultivars. Although the success of this strategy has not yet been proven in rice, crosses have been
made to combine two major genes for grown planthopper resistance and future studies will indicate
the value of this strategy in slowing the development of biotype selection and increasing the
commercial life of brown planthopper resistant cultivars.
6.3 HORIZONTAL RESISTANCE

Horizontal resistance is a type of resistance that is expressed equally against all biotypes. Because
it is generally considered to be of a polygenic nature (several minor genes), which is difficult for an
insect species to overcome, it is durable. In contrast to antibiosis, tolerance is of a polygenic nature
and is durable because it exerts no selection pressure on the insect population. Tolerant plants
possess the ability to grow and reproduce and to repair injury in spite of supporting a population
approximately equal to that damaging a susceptible host (Painter 1951).
Greenhouse and field methods have been developed to evaluate rice cultivars for tolerance to
the brown planthopper (Heinrichs et al., 1985a, Panda and Heinrichs 1983). Field evaluation of
moderately resistant cultivars identified cultivar Utri Rajapan from India as having a high level of
tolerance. Genetic analyses indicated that the field resistance (level of resistance increases with
plant age) in Utri Rajapan and Triveni is governed by the same R gene (Velusamy et al., 1987).
Triveni, also an Indian cultivar, possesses both antibiosis and tolerance.
6.4 MULTILINES
This strategy involves the incorporation of different R genes into isolines and planting the lines
together in the same field as a mixture. Although a feasibility study was conducted under field
conditions at IRRI the lack of isogenic lines and the difficulty for farmers to maintain seed mixtures
has dampened the enthusiasm for this approach.
6.5 GENE ROTATION AND GEOGRAPHICAL DEPLOYMENT

Gene rotation, also referred to as temporal deployment, is a strategy where cultivars with different
R genes are used in different cropping seasons. In geographical deployment different R genes are
used in adjacent cropping areas.
Gene rotation is used in certain regions in Indonesia where the green leafhopper-vectored
tungro virus has been a severe threat to production. A set of cultivars with certain genes for green
leafhopper resistance is planted in the dry season and another set of cultivars with a different gene
is planted in the wet season
A type of geographical deployment is utilized in selected villages of northern Vietnam. Fields
within a village are planted to several different cultivars having different maturities and grain types
and possessing different R genes for brown planthopper resistance. This is a type of diversity that
insures against several abiotic and biotic stresses and minimizes the need for labor at harvest time
because the harvest season is spread over a long period.
6.6 BIOTECHNOLOGY
Recent advances in biotechnology provide the possibility of solving some of the constraints that
have limited the practical use of genetic resistance to insects in pest management programs.
Biotechnology involves several techniques that provide new possibilities of manipulating
germplasm.
6.6.1 Wide Hybridization

Wild rice species are a rich source of R genes for use in breeding programs for insect resistance.
This is of extreme importance in breeding for resistance to those insects for which R genes have not
been found in Oryza sativa and in cases where biotype selection may overcome all of the O. sativa R
HEINRICHS 541
genes. In this context, wide hybridization is a plant breeding tool for the incorporation of alien
genetic variation from wild species of Oryza into commercially useful cultivars.
Most of the wild rice accessions in the IRRI germplasm collection have been screened for
resistance to planthoppers, leafhoppers, stem borers, leaffolder Cnaphalocrocis medinalis, whorl
maggot Hydrellia philippina and the caseworm Nymphula depunctalis (Heinrichs et al., l985a. WU.
Jung-Tsung et al., 1986, Romena and Heinrichs 1989). Wild rices are a rich source of insect
resistant germplasm and may provide new genes which can be used in the various gene deployment
strategies (Table 3). The embryo rescue technique has been utilized in the crossing of O. sativa and
wild rice species in breeding for brown planthopper resistance (Khush and Jena 1986).
TABLE 3
Agronomically important characteristics identified among the wild Oryza species (from Toenniessen and Herdt 1989)
Species 2n Genome Characteristics

O. nivara 24 AA Grassy stunt virus resistance
O. rufipogon 24 AA Source of cytoplasmic male sterility, tolerance to stagnant flooding
O. glaberrima 24 AA GLH resistance, early vegetative vigor
O. barthii 24 AA Bacterial blight resistance
O. longistaminata 24 AA Floral characteristics for outcrossing
O. punctata 24,48 BB,BBCC BPH, WBPH, GLH resistance
O. officinalis 24 CC BPH, WBPH, GLH resistance
O. eichingeri 24 CC BPH, WBPH, GLH resistance
O. minuta 48 BBCC BPH, WBPH, GLH, blast, and bacterial blight resistance
O. australiensis 24 EE BPH resistance, drought tolerance
O. brachyantha 24 FF Whorl maggot and stern borer resistance
O. ridleyi 48 — Whorl maggot resistance
Wild rice species are serving as useful donors in wide hybridization projects for rice improve-
ment. The IRRI breeding program has utilized the wild rice species. O. brachyanthta in crosses with
0. sativa in the development of brown planthopper, whitebacked planthopper and stem borer
resistance. Some progeny are resistant to all known biotypes of the brown planthopper as well as
resistant to the whitebacked planthopper.
6.6.2 Molecular Genetics

The Rockefeller Foundation International Program on Rice Biotechnology was established in 1984
and has significantly enhanced the scientific input into the development of molecular biology
techniques for rice (Toenniessen and Herdt 1989). The program consists of a coordinated effort
among research institutions in the developed world that are at the forefront of plant molecular and
cellular biology and the International Rice Research Institute.
Based on a set of criteria a list of priority traits, for use in the Rockefeller Foundation Rice
Biotechnology Program, was established. Insect pests include the yellow stem borer, gall midge,
brown planthopper, leaffolder, storage pests, striped stem borer, and whitebacked planthopper
(Table 4).
TABLE 4
Rank order of priority traits for the Rockefeller Foundation’s Rice Biotechnology Program
(from Toenniessen and Herdt 1989)
Resistance to tungro virus

Resistance to yellow stem borer
Resistance to gall midge
Cytoplasmic male sterility
Drought tolerance
Resistance to brown planthopper
Submergence tolerance
Greater lodging resistance
Seedling vigor
Resistance to ragged stunt virus
Tolerance to drought at anthesis
Tolerance to waterlogging
Resistance to leaffolder
Resistance to sheath blight
Cold tolerance at seedling stage
Apomixis
Tolerance to coastal saline/acid sulfate conditions
Resistance to bird damage
Resistance to storage pests
Resistance to bacterial blight
Resistance to blast
Resistance to striped stem borer
Resistance to whitebacked planthopper
The Rockefeller Program includes the (1) development of genetic maps and markers,
(2) protoplast techniques, (3) genetic transformations, and (4) cloning and characterizing rice
genes. The genetic transformation of rice through the introduction of alien or modified genes is a
principal goal of the program. Genetic engineering should make it possible to incorporate any gene
from any source into rice. This will make it possible to further utilize the insect resistant wild rice
accessions and other much more distant relatives of rice. It has been demonstrated that transgenic
plants other than rice express toxin genes from the insect pathogen, Bacillus thuringiensis and are
resistant to insect feeding. If this can be done in rice there are hundreds of B. t. strains that can be
utilized in the development of transgenic plants that express resistance to the various lepidopteran
species such as the leaffolder, stem borers, and caseworm.
Because of the high yields of modern rice cultivars, the amount of rice in storage has increased.
Storage insects cause severe losses and there is a need to develop cultivars with resistance to insects
attacking grain in storage.
Genetic engineering provides the potential of providing a novel means of rice storage insect
control. Current projects supported by the Rockefeller Foundation are seeking to isolate genes
from wheat which code for inhibitors of digestive enzymes of the rice weevil Sitophilus oryzae, a
HEINRICHS 543
major pest of stored rice. The goal is to isolate wheat genes that provide a good source of genetic
resistance to weevils and transfer them to rice which lacks resistance to weevils (Toenniessen and
Herdt 1989).
7. CONCLUSIONS
Rice is an outstanding example of the successful utilization of host plant resistance as a tactic in the
integrated management of insects. The large number of cultivars in the world collection of rice is a
wealth of genetic diversity. Much of the world collection has been evaluated for insect resistance
and many cultivars have been used as donors in breeding programs throughout the world. These
breeding programs have successfully provided rice farmers with insect resistant commercial cul-
tivars that are playing a significant role in the management of rice insects.
In spite of the significant progress in the breeding and commercial exploitation of insect
resistant rice cultivars, much remains to be done. There are still numerous insects for which suitable
levels of resistance have not been found in O. sativa. Molecular genetics holds promise as a means
of transferring insect resistance from wild rice species and other species into O. sativa. In certain
cases insect resistant commercial rice cultivars have succumbed to biotype selection. Innovative
conventional plant breeding techniques and molecular genetics may provide means of developing
commercial cultivars with stability to variable insect populations.
It must again be stressed that no one control tactic is a panacea for rice insect control. Host
plant resistance integrated with cultural, biological and chemical control tactics has been shown to
be an effective strategy for rice insect control and must be pursued by rice entomologists working
in concert with scientists in other disciplines.
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Shiraki, T. 1917. Paddy borer, Schoenobius incertulas Walk. Taihoku Agric. Exp. Sta.
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Sogawa, K., and M. D. Pathak. 1970. Mechanisms of brown planthopper resistance in Mudgo variety of rice (Hemiptera:
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6
Cultural, Mechanical, and Physical

Control of Rice Insects
JAMES A. LITSINGER
Intentational Rice Research Institute
Los Baños, Philippines
1. INTRODUCTION
Cultural methods to control insects involve crop husbandry practices that have a dual purpose of
crop production and insect pest suppression. Farmers have developed these practices through
observation and trial and error. The insect control component of these practices, handed down
through generations, is often forgotten by present day farmers. Thus there are two types of cultural
control practices, primary and secondary. Primary cultural control practices are those done specifi-
cally to control insects such as draining a field when the brown planthopper is abundant, planting a
trap crop for stem borers, or transplanting older seedlings to minimize whorl maggot damage.
Secondary practices are those that are specifically done for crop husbandry such as land prepara-
tion, weeding, and fertilization but also happen to minimize insect buildup. Farmers often adopt
cultural practices that embody several beneficial effects. Plowing prepares the soil for planting but
also turns under stubble harboring stem borers. Weeding grasses removes host plants of the rice
bug. Splitting application reduces fertilizer losses as well as minimizing brown planthopper buildup.
Mechanical and physical control practices use principles of physics — mass, energy, and force.
Mechanical methods involve motion and force such as trapping or crushing insects by hand, tool, or
machine. The energy to produce the force can be chemical or gravitational. Man and machines
chemically convert food and fuel into mechanical energy. Gravity often augments this energy.
Physical methods involve mass such as an obstruction (covering a seedbed), electromagnetic energy
550 CULTURAL, MECHANICAL AND PHYSICAL CONTROL OF RICE INSECTS
such as heat (spreading straw in the sun to kill resident stem borer larvae), or light (light traps) to
arrest nocturnal flying insects.
Cultural, mechanical, and physical methods of insect control are often complementary. Tillage
by plow and harrow is a cultural control practice, but insects die from mechanical crushing and
physical exposure to heat. Flooding is a cultural practice to control weeds and minimize drought but
the timing and depth can be manipulated to physically drown insects such as armyworms attacking
a seedbed. Dense plantings provide physical obstructions to flying insects. Older seedlings are
physically harder for stem borer larvae to tunnel into. Burning straw is normally done as a cultural
practice to remove a waste product because straw does not biodegrade quickly, but insects are
physically killed by the heat.
Magical and superstitious practices are, in the broad sense, forms of cultural control that
employ the anthropological definition of culture as patterns of behavior handed between genera-
tions. Rice culture has a rich source of folklore regarding indigenous practices. Unfortunately few
of these methods have been objectively evaluated as to efficacy. Most practices that offer potential
fall in the categories of chemical, cultural, mechanical, and physical control.
TABLE1
Cultural, Mechanical, and Physical Insect Control Practices for Rice
Cultural Mechanical Physical

Field geometry Handpicking from plants Light trap
Mixed cropping Tools to capture or remove insects Shaving bunds to drown or expose insects
Planting method Dislodging insects by beating foliage or Plastering bunds to trap insects in their
passing a rope tunnels
Age of seedlings Digging out insects Barriers (trenches or nursery boxes)
Pruning Roguing Burning stubble or straw
Plant density Pounding or rolling the soil to crush insects Spreading straw in the sun
Crop background
Water management
Fertilizer
Crop rotation
Crop area
Number of rice crops
Plant maturity
Planting time
Synchronous planting
Trap crop
Tillage
Weeding
Harvest method
Use of straw as fodder,
mulch, fuel or mush-
room culture
Ratooning
Grazing
LITSINGER 551
Mechanical, physical, and cultural rice insect control methods are listed in Table 1. I have
divided the discussion of mechanical and physical control methods in this chapter into two
categories – (1) those that require mainly labor and (2) those that require mainly capital. Cultural
control methods are divided into single field and community-wide methods.
2. LABOR INTENSIVE MECHANICAL AND PHYSICAL METHODS

Manual methods of controlling rice insects are among the oldest and most labor intensive. These
methods decline in usage as labor costs rise and less expensive alternative methods become
available. Socialist countries that can organize large gangs of cheap labor have a greater tendency
to practice them. Manual methods of insect control are perhaps underutilized as they are no more
tedious than manual weed control practices (Maxwell-Lefroy 1906). Manual methods of insect
control are most attractive for high value crops where labor is cheap, insect pests are aggregated,
and the collected insects can be used as food or have an economic value (Isely 1951). Collected
insect egg masses can be held in containers to allow parasites to escape, increasing this method's
utility.
2.1 HAND PICKING

The most direct manual method is removal of insects by hand. Fast moving or camouflaged insects
are too difficult to collect. Slow moving and aggregated large armyworm, cutworm, and butterfly
larvae can be handpicked from foliage (Maxwell-Lefroy 1906, Hutson 1920, Kuwana 1930, Liu
1935). Rice bug nymphs fall when disturbed and can be tapped into hand held buckets filled with
kerosene and water (Austin 1922). Grasshopper adults are sluggish at night and can be picked
directly from foliage and either eaten or destroyed by burning, crushing, burial, or placing in oil or
kerosene (Jones and Mackie 1913). Immobile caseworm larvae are skimmed from the water surface
after the foliage is beaten (Ghosh 1924a, Otanes 1947). Dispersing stem borer larvae cling to
floating leaves that can be removed (Kondo 1917). But insects floating in water are more efficiently
removed by nets than by hand.
Eggs are immobile and are thus attractive stages for handpicking. Species which lay eggs in
masses are preferred. Rice bug (Otanes 1937), stink bug (Wallace 1957), and black bug (Corbett
and Yusope 1924) eggs can be removed by hand and placed in special containers to allow parasites
to escape. In Japan, China, and India, stem borer and armyworm egg masses as well as moths were
traditionally removed by hand from seedbeds (Ramakrishna Ayyar 1933). Moths are difficult to
capture by hand but are relatively easy to capture by net. Skill is required to locate Chilo and
Sesamia egg masses because they are deposited behind leaf sheaths. Best results occur with
Scirpophaga, Mythimna and Spodoptera which lay egg masses openly on foliage.
Seedbeds laid out in 1 m wide beds between alleyways facilitate inspection by children (Dam-
merman 1929). Egg masses are removed every 5 days and placed in special containers to allow
parasitic wasps to escape while trapping the emerging larvae in a kerosene moat (Kiritani 1979).
The necessity of removing egg masses from seedbeds is questionable. Seedlings have narrow solid
stems that cannot be successfully tunnelled by first instar stem borer larvae which then succumb to
natural enemies and desiccation. The stem walls of seedlings are also densely packed with silica
making larval penetration difficult. Consequently, removing stem borer egg masses from the
seedbed is considered to be of doubtful value because damaged plants usually compensate from
early stage damage (Jepson 1954). Those that believe removing egg masses is highly useful cite up
to 40% yield loss if not done (Ghose et al., 1960). Such studies probably refer to situations where
older more susceptible seedlings undergoing elongation were transplanted.
Removal of stem borer egg masses from the field is more efficiently done if timed as based on
light trap collections (10 days after a peak moth flight) or timed to coincide with a susceptible crop
growth stage. Light trap peak catches in tropical environments, however, have proved to be poor
indicators of stem borer field populations. Better prediction is possible in temperate areas. In
northern China, egg collections are timed for three generations of yellow stem borer and two
generations of striped stem borer (Asa and Fey 1924). Timing of stem borer egg mass removal
should coincide with susceptible growth stages – (1) stem elongation in the vegetative stage (to
prevent deadhearts) and (2) panicle exsertion (to prevent whiteheads) (Koyama and Tsuchiyama
1964, Tian 1981). Hand removal of egg masses 1 week before panicle exsertion reduced whiteheads
by 45% and increased yield by 18% in China (Shiraki 1917) and reduced whiteheads by 50% in
Japan (Jepson 1954). It takes approximately 4 days for one skilled person to remove yellow stem
borer egg masses from 1 ha (Grist and Lever 1969).
With the onset of monsoon rains, scarab beetles in dryland rice areas fly to host trees to mate.
Their larvae, called white grubs, feed on rice roots. White grub beetles can be shaken from trees and
collected by children (Otanes 1947, Raodeo and Deshpande 1987). Community efforts are success-
ful because the beetles, with one or two year life cycles, aggregate to preferred host trees and are
abundant only over a period of several months each year. Collecting adults prevents egg laying. But
some report that the beetles in the trees have already laid their eggs thus nullifying the effect
(Kalshoven 1981). Adults can also be boiled and deep fat fried as food.
Swarming locusts can be herded into pits. The technique is to form a human gauntlet made by
two rows of workers holding banana leaf or galvanized iron sheet barriers (Jones and Mackie 1913).
Others drive the nymphs slowly without alarming them, otherwise they scatter. Rather than digging
pits, the nymphs can be driven into portable bag traps (Pratt and South 1913).
Tools have been helpful in removing insects by hand. In Japan striped stem borer larvae,
emerging in mass in the spring from stored straw, were raked up and destroyed (Japan 1953). In the
field, large combs were designed to run through rice foliage to remove caterpillars (Kuwana 1930).
Sweeping rice plants with nets, bags, or baskets – some coated with sticky materials (jackfruit,
latex, castor oil, grease) – have been used to remove rice bugs (Barnes and South 1925), black bugs
(Corbett and Yusope 1924), stink bugs (Wallace 1957), leafhoppers (Abalos 1939), planthoppers
(Miller and Pagden 1930), armyworm larvae (Nanta 1933), caseworm larvae (Shroff 1920), leaf
beetles and hispa (Ghosh 1924b, Kuwayama and Onozaki 1936), grasshoppers (Main 1912) and leaf
worms (Korea 1973). Early morning is the best time to net hispa and Oulema larvae that are
transfering to new leaves (Taiwan 1936). One person can net 2 liters of hispa per day. Armyworms
LITSINGER 553
are best netted at night or, if the field is flooded, swept into baskets (Ramakrishna Ayyar 1933).
Large canvas bags (2 x 1 m) on bamboo frames or sectioned oil drums coated on the inside with
grease are dragged through rice foliage by two men to capture insects. Walking into the wind keeps
canvas bags open and blows the insects inside. Wind inflation of the bag also minimizes crop
damage. Some farmers object, however, to passing a net through a crop at the ripening stage, but
allow it in grassy borders (Corbett 1930). Triangular aquatic nets are used to collect armyworms,
caseworms, and migrating stem borer larvae in cases (Banerjee 1954). Often the water level is raised
ahead of time so the insects can be driven to the top portions of the plants for easy removal. Ropes
are dragged across the plants or the plants are beaten to dislodge the insects (Ballard 1922).
Sometimes ropes are coated with grease to snare rice bugs and other insects (Hutson 1930). A man
can rotate a rope across the foliage by tying one end to a stake and walking in a circle holding the
other end taut.
Digging up mole cricket (Novelli 1914) and grasshopper (Pratt 1915) eggs from the soil is most
efficient in breeding grounds (Gonzales 1932). Plowing is preferred to digging with a shovel or hoe.
Mole crickets, their egg pods (Korea 1973) and white grubs (Otanes 1947) are collected by hand
from fields during plowing. Mole cricket tunnels are destroyed by shovel (Korea 1973) and adults
eaten (Nanta 1935). Logs are dug out of slash-and-burn dryland fields to remove termites (Dam-
merman 1929), but others (Raymundo 1986) state that logs are beneficial and act to divert termites
from the crop.
Techniques have been developed to attract insect pests for more efficient manual destruction.
The methods have been most commonly used in seedbeds or small fields. Rafts made of bamboo or
banana leaves attract armyworm larvae after the crop is submerged (Otanes 1947). Weeds can be
heaped to attract armyworm larvae (Nanta 1935), mole crickets (Kureha et al., 1974), or field
crickets (A.T. Barrion, IRRI, personal communication). These insects should be regularly
destroyed otherwise they will build up to severely damage the crop. Echinocnemus root weevil
adults are attracted to erect sticks placed around the seedbed or to sweet potato sections
(Kuwayama 1963). Bundles of straw dipped in a mixture of sugar, wine, and vinegar attracts
Echinocnemus adults. Male white grub beetles are attracted to crushed chillis (Dammerman 1929).
Rice bugs are attracted to putrefied meat (Dresner 1958) or decaying aquatic plants (van der Goot
1949) bagged and suspended from poles around a field. Such mixtures are laced with insecticide.
2.2 ROGUING
Roguing is removal of pest infested plants or plant parts by hand or with hand tools. The damage
has been done but the effort is exerted to prevent the pest population from increasing. Stem borers
(Maxwell-Lefroy 1907, Okazaki 1938), mealy bug (Cherian et al., 1935), root aphids (Tanaka 1961),
gall midge (Nguyen 19221, and leaffolders (Korea 1973) have been claimed to be controlled by
roguing. Removing deadhearts and whiteheads was first popularized on sugarcane (Jepson 1954).
For rice stem borer control roguing should be done thoroughly over a large area and over a number
of years (DuPort 1919). However, pulling deadhearts or whiteheads often fails to extract all the stem
borer larvae (Ramakrishna Ayyar and Anantanarayanan 1937). Others practice cutting off dead-
hearts with a knife (Kondo 1917).
A small hand tool was developed in Japan to remove whiteheads but a farmer could only kill
40-80 larvae per day as only half to two-thirds of those whiteheads contained larvae. A further step,
said to be only efficient on young rice plants, is to uproot the whole plant upon seeing deadhearts
(Usman 1954). Such a practice is counterproductive as plants are destroyed. Roguing silver shoots
for gall midge control is advocated but some feel this hinders the plants’ compensatory ability
(Barnes 1956).
2.3 BUND SHAVING

Mowing and shaving rice bunds is a common rodent and weed control practice that secondarily
affects a number of insect pests that spend at least a part of their life in the soil. Armyworms pupate
(McKay 1917), grasshoppers oviposit (Ramakrishna Ayyar 1933) and mole crickets tunnel (Kureha
et al., 1974) in rice bunds. Soil trimmed from the bunds can be thrown into the flood water to drown
eggs and pupae. The sides of bunds can be plastered with fresh mud to trap mole crickets (Hutson
1920).
2.4 CRUSHING, BEATING, AND DRAGGING

Locust breeding grounds can be pounded with malls to crush eggs (Jones and Mackie 1913).
Plowing is an easier method. Rice bugs are driven from a crop by frequent beating with sticks
(Maxwell-Lefroy 1906). Farmers can beat the foliage during the day but rice bugs are also active at
night. A rope soaked in kerosene and held by two men can be brushed across the foliage to kill
thrips, hispa, and mealybug (Banerjee 1954, Ghose et al., 1960) by contact. The degree of control,
however, is minimal.
2.5 BARRIERS
Aggregated nonflying armyworm larvae, ants, and chinch bug and locust nymphs can be prevented
from invading a field or a seedbed by constructing a moat, trench, or petroleum barrier along the
field perimeter. Trenches with back-sloping front walls have a greater effect (Hutson 1920). A water
moat can be made more impervious with a kerosene film. Barriers work only if the direction of pest
movement can be predicted and if the movement occurs over a short period (Isely 1951).
3. CAPITAL INTENSIVE MECHANICAL AND PHYSICAL METHODS

Using mesh or cloth to protect the seedbed or rice crop is a relatively new technology whereas the
use of kerosene light traps for insect control is a traditional practice. The former is used because
other modern methods failed and the latter was developed before the era of highly effective
petroleum-based insecticides.
3.1 BARRIERS
In temperate areas, nursery boxes are used to incubate seedlings for machine transplanting during
early spring and this practice prevents early infestation of insect vectors (Kiritani 1983) and root
LITSINGER 555
weevils (Kuwayama 1963). In warmer climates, covering seedbeds with mesh cloth prevents virus
transmission by leafhoppers. But covered seedbeds developed high stem maggot infestations
(Okamoto 1957). Farmers in Japan have gone to the extreme of erecting vinyl sheeting wall barriers
to prevent Nephotettix from entering fields (Hashizume 1964).
3.2 ROLLERS
Animal drawn rollers can be used to crush armyworms and locusts in fallow areas before they invade
rice fields (Hutson 1920, Jones and Mackie 1913). Rollers are used to kill overwintering stem borers
in stubble (Wouters et al., 1961) and work best if combined with flooding (Douglas and Ingram
1942). In Australia, crushing the stubble with tractor drawn rollers killed 88% of Scirpophaga
innotata compared to 58% mortality from burning (Halfpapp 1979).
3.3 OIL FILM ON PADDY WATER

A traditional method to control many insect pests is to pour 30-40 liters of kerosene per ha onto a
flooded field (Korea 1973). A kerosene soaked rag can also be placed at the water inlet (Ghosh
1924a). Kerosene works best against more mobile insect pests — leaf beetles (Frappa 1938), black
bugs (de Alwis 1941), leafhoppers (Japan 1913), planthoppers (Suenaga 1966), seed bugs (Maxwell-
Lefroy 1908), root weevils (Kuwayama 1963), armyworm larvae (MacKenna 1918), caseworm larvae
(McSwiney 1919), caterpillars (Chu 1936), and grasshoppers (Korea 1973).
In a young crop, two men can drag a rope across the foliage bending it into the kerosene film
on the water (Misra 1920). Older rice plants are beaten with bamboo poles or branches or brushed
with a rope soaked in kerosene to dislodge insects into the oil. More insects are killed if the
operation is done early in the morning when insects are lethargic and after raising the water level as
high as possible (Sakai 1934). This technology dates from the 1600s in Japan where whale oil was
used before kerosene became available (Korea 1973). A kerosene film is made more toxic if mixed
with pyrethrum (Iso 1954). Fields were drained after 6 hours to prevent phytotoxicity. Oil films on
the paddy water are highly toxic to many predators and may have caused resurgence of brown
planthopper (Kenmore et al., 1984).
3.4 LIGHT TRAPS

Impressed with the many insect pests attracted to light, farmers long ago used fire and established
grids of light traps as a communitywide control measure. Bonfires were lit to attract rice bugs
(Hutson 1930) and moths (Ramakrishna Ayyar and Anantanarayanan 1937) which then died in the
flames. Light traps are said to control stem borers (Yusope 1920), armyworms (Jhaveri 1920),
leafhoppers (Fletcher 1913), seed bugs (Sen and Chaudhuri 1959) mole crickets (Otanes 1947),
leaffolders (Nanta 1935), gall midges (Murthy 1957), planthoppers (Ghose et al., 1960), root weevils
(Newel1 1913), caseworms (Ghose et al., 1960) and caterpillars (Korea 1973). Greater numbers of
leafhoppers, planthoppers, and black bugs are collected during a full moon whereas more moths are
attracted during a new moon. At peak adoption of this method in the 1930s, over one million traps
were set out 1-1.5 m above the crop canopy in China (1 trap per 2.3 ha) (USA 1977) and in Japan
556 CULTURAL, MECHANICAL AND PHYSICAL CONTROL, OF RICE INSECTS
(20 traps per ha) (Kaburagi 1938). A trial in India placed 7 petromax powered light traps per 200
ha and claimed 33% yield increase from gall midge control (Khan and Murthy 1955). As further
proof of the value of light traps, pest outbreaks have been reported to occur in India after power
blackouts because many insects are normally killed at street lights (Israel and Seshagiri Rao 1954).
Electric powered traps attract high insect numbers but are expensive to operate. Inexpensive
low voltage electric traps were developed in China to lower costs (Wittwer 1987). To further
minimize expenses, light traps can be set out using seedbeds as trap crops or during critical periods
such as right after transplanting for stem borers (Ballard 1923, Puttarudriah 1945) and gall midge
(Murthy 1957) and at booting for stem borers (Ramakrishna Ayyar and Anantanarayanan 1937).
Also costs can be reduced by operating the traps only during the first few hours after sunset
(Kaburaki and Kamito 1929).
Greater catches of leafhoppers are claimed if a person walks through the field at night carrying
a lantern held over a basin with soapy water while a second person beats the rice foliage with a stick
(Abalos 1939). Better results are achieved if done early in the crop season before most leafhoppers
have oviposited.
The use of light traps to control insects has been criticized as farmers in Japan and China noted
the same number of deadhearts or whiteheads with or without light traps (Shiraki 1917). Dissatis-
faction with light traps also came because of their high cost (Li 1982), theft (Ballard 1923), erratic
attraction (Pang 1932), and killing of beneficials (Kondo 1917). Not all pest species are attracted
and trapped. The sex ratio of collected insects often favors males (Squire 1933). Many attracted
females also had laid most of their eggs (Stewart 1934).
The recent revelation that captures from pheromone traps occur at different times than those
in light traps points out that light traps attract insects which are dispersing and thus are not
reproductively active. A high mortality normally is associated with dispersing insects so the efficien-
cy of the trap is less than if it attracted virgin females.
Cultural control methods employing crop husbandry practices which require little additional
labor or capital may be divided into two spatial dimensions: (1) single field or (2) communitywide.
Some practices are effective on either scale.
4. SINGLE FIELD CULTURAL METHODS

4.1 FIELD GEOMETRY
Larger and more square fields have a lower perimeter to area ratio and therefore suffer relatively
less damage from water weevils (Grigarick 1984) or grasshoppers (Main 1912) that accumulate
along field borders. Small isolated fields concentrate highly mobile rice seed bugs which seek out
flowering stage rice crops (Lim 1970a).
4.2 MIXED CROPPING

One of the first decisions in crop husbandry is selecting the type of crop to be grown. Dryland rice
has greater opportunities than puddled rice for crop diversification. Few studies have been con-
LITSINGER 557
ducted to show how insect abundance is affected by intercropping. Dryland rice intercropped with
cotton or pigeon pea in one study (Satpathy et al., 1977) resulted in low green leafhopper and
whitebacked planthopper numbers. Mixing dryland rice cultivars of various maturities and harvest-
ing individual panicles at maturity reduces storage time and consequently storage pest problems
(Raymundo 1986). Wetland rice intercropped with sweet flag, an insect repellant plant, did not
affect gall midge (CRRI 1964). In Java the labor intensive sorjan system of alternating ditches and
raised beds is practiced. Wetland rice is transplanted in the ditches, and vegetables, field crops,
and/or fruit trees are cultivated on the beds. The nonrice crop may offer protection to adjacent
intercropped rice. But two studies comparing the incidence of gall midge, yellow stem borer, and
leaffolder in sorjan with that in nearby sole cropped ricefields showed no consistent differences in
pest abundance (Ruhendi et al., 1976, Rochman et al., 1977).
4.3 PLANTING METHOD

Rice can be planted either by direct sowing into dry or moist soil with dry seeds, onto puddled soil
with pregerminated seeds, or by transplanting seedlings in fields with or without flood water. Direct
seeding onto dry or nonflooded soil is disadvantageous to aquatic pests. On the other hand,
transplanting into flooded fields prevents the establishment of dryland adapted pests. Planting rows
oriented north-south lessens intraplant shading and consequently is said to reduce brown planthop-
per incidence (Oka 1979).
Most of the effects of planting method on pest numbers are the result of differences in planting
time, plant density, or water level which occur as a result of changing planting methods.
Transplanted rice is often reported to have higher stem borer numbers than a direct seeded crop
(Jinnendradasa 1954, Koyama et al., 1968, Israel 1969) which can be attributed to later planting,
lower plant density, and a proportionally longer flooding period than a direct seeded crop.
Direct seeded rice often grows less vigorously without standing water during the vegetative
stage making it less attractive to yellow stem borers. Stems of direct seeded rice grow more hard and
narrow, and therefore cause higher mortality to first instar stem borers (Shiraki 1917). Transplanted
crops mature later in the season allowing stem borers to build up on earlier plantings.
A seedbed offers certain advantages for crop protection. The longer seedlings remain in the
seedbed the less time vegetative stage pests such as whorl maggot (IRRI 1980) and gall midge
(CRRI 1964) have to damage the transplanted crop. Because of their smaller area, seedbeds can be
more readily protected from insect vectors than a transplanted field so delayed transplanting is a
management tactic to control vector-transmitted diseases. This is particularly important because
delayed infection of a crop reduces disease severity (Kiritani 1983). Seedlings can be cultured on
plastic or concrete away from fields and thus isolated from insect vectors. In a cold climate,
seedlings can be grown in protected nursery boxes and planted earlier than a direct seeded crop.
Machine transplanted plants from seedbeds grown under cover had low incidence of dwarf virus
vectored by the green leafhopper in Japan (Kiritani 1983). Whorl maggot is not attracted to a
densely planted seedbed (Bandong and Litsinger 1986). Alternating flooding and draining of
seedbeds, which is practiced to enable seedlings to be easily pulled, is detrimental to both dryland
and aquatic adapted pests. Control of pests is less expensive in a seedbed than a field (Lippold
1972).
A transplanted crop has a shortened field maturation and with the exception of whorl maggot
is less damaged by vegetative stage pests. Bundled seedlings, prepared for transplanting, can be
held for two days to drive out stem borer larvae (Ramakrishna Ayyar and Anantanarayanan 1937)
or can be dipped in hot water to kill hispa (Bouriquet 1937).
4.4 AGE OF SEEDLINGS

Irrigated rice farmers have the choice to plant either young or old seedlings, but yields of modern
cultivars decline dramatically when older seedlings are transplanted. Rainfed farmers plant tradi-
tional cultivars that are little affected by seedling age. The rainfed farmer has selected such cultivars
because he must quickly transplant when rains arrive. Increasing seedling density per hill is
necessary to compensate for lack of tillering in older seedlings. The effects of seedling age on stem
borer abundance are however inconsistent (CRRI 1983).
4.5 PRUNING
Clipping the tops of bundled seedlings is primarily a practice to prevent lodging (of old seedlings)
and stimulate tillering of traditional cultivars but secondarily removes stem borer (Khan 1967) and
hispa eggs (Reddy 1968). The practice is not commonly done with modern semidwarf cultivars that
are 2-4 weeks old when transplanted and will actively tiller.
Removing the top third of standing crop foliage before flowering has been a practice with
traditional tall cultivars to obtain fodder for cattle during the wet season when fallow lands are few
(Otanes 1947). This practice removes stem borer egg masses (Catling and Alam 1977), hispa
(Agarwala 1955), thrips (Usman 1954), leaffolders (Otanes 1947) and caterpillars (Alam and Alam
1977). Pruning may stimulate higher yields even on a healthy crop (Tripathi et al., 1973) but should
not be carried out if the crop is under stress (Jepson 1954). This practice, on the other hand,
stimulates tillering making the plants more susceptible to gall midge (CRRI 1975). Cutting rice
foliage also makes wounds for entry of bacterial leaf blight (Chu 1971).
4.6 PLANT DENSITY

Dense planting increases populations of planthoppers (IRRI 1977), leafhoppers (Shukla and
Anjaneyulu 1981a), leaffolders (Kushwaha and Sharma 198l), gall midge (Pillai et al., 1979), black
bug (Dammerman 1929), and caseworm (Venugopal Rao et al., 1982). But whorl maggot (CRRI
1978), root aphids (Tanaka 1961), root weevils (Rolston and Rouse 1964), and leaf beetles
(Dhaliwal and Jaswant Singh 1979) are less abundant. Reports on stem borers are mixed; some say
stem borers are more abundant (Israel 1969), while others less (Kiritani and Iwao 1967). High
seeding rates generally guarantee high yield and offer greater tolerance from those pests that
remove seeds or kill seedlings (Pimentel and Goodman 1978). Effects of plant density on insect pest
abundance can however be more varied and complex. Dense plantings change crop growth,
development, and the microclimate which in turn affect pests and their natural enemies (Teetes
LITSINGER 559
1981). Sparse plantings encourage weeds, and indirectly affect insect abundance. Gall midge
numbers rise proportionally with the density of growing points (tillers), therefore greater incidence
of galls occurs in dense plantings (Vungsilabutr et al., 1972).
The complexity of the effects is further elaborated with stem borers. More moths hide during
the day in dense plantings, presumably because they prefer a high humidity (CRRI 1961). However,
at night they oviposit more in sparse plantings, but egg parasites will reach more egg masses in a less
dense crop. Both stem borers and parasites can easily penetrate the canopy in less dense plantings
(van der Goot 1925). Yellow stem borer moths are also more attracted to water which is more
detectable in sparse stands. Tillers elongate more and are softer in dense plantings — thus are more
susceptible to penetration by the first instar larvae ensuring higher survival and more deadhearts.
High tiller densities result in more infested tillers per area but a lower percentage (CRRI 1970).
Results, therefore, should be expressed as the number of infested tillers or deadhearts per area as
well as percentage. If there is competition among stem borer larvae for tillers, more tillers guarantee
more larvae will survive (Yamazaki and Hatai 1960). However if greater fitness occurs with wider
diameter tillers then more larvae will survive in sparse plantings.
Brown planthopper numbers dramatically increase in a dense planting (Hino et al., 1970). The
reason given by most researchers is that high relative humidity favors survival (Pillai et al., 1979).
However, phytotron studies show that brown planthopper mortality increases at humidities over
60% perhaps because of increased pathogen infection (IRRI 1976). Two other explanations are (1)
dense plantings provide more plant surface area and less competition for oviposition or feeding
sites (Kenmore et al., 1984) and (2) denser canopies impede egg parasitic wasps (Shepard and
Arida 1986).
Whorl maggots lay progressively fewer eggs per area as plant density increases (Bandong and
Litsinger 1986). Females may be first attracted to open water and secondarily to rice plants. If the
water surface in the field is covered with a dense crop, whorl maggots overfly the field.
4.7 CROP BACKGROUND

Low whorl maggot incidence when azolla water fern covers the water surface during the early
vegetative stage, supports the water attraction theory (Viajante and Heinrichs 1985). Reports that
azolla augments green leafhoppers and tungro virus are unfounded (IRRI 1983a). The orange
leaves of tungro-infested plants contrast more against a green azolla background than open water,
and therefore stand out more. Plastic mulch to conserve moisture in dryland rice in Japan resulted
in an outbreak of striped stem borer (Ichihara and Matsuo 1970). Perhaps soil dwelling stem borer
predators were impaired.
4.8 WATER MANAGEMENT

Aquatic pests — whorl maggot (Feron and Audemard 1956), root-feeding midges (Abul-Nasr et al.,
1970), water weevils (Isley and Schwardt 1934), yellow stem borer (Fang 1977), and caseworm
(Sison 1938) — are suppressed when fields are drained for 1 or 2 days. Caseworm larvae respire via
gills and midge larvae through their body wall. Larvae of water weevils respire through pointed
dorsal protuberances jabbed into the air spaces of roots. The air spaces collapse when the plant is
not in standing water (B.S. Vergara, IRRI, personal communication). Draining the field is a
common practice to control planthoppers (Das and Thomas 1977) and armyworms (Anan-
tanarayanan and Ramakrishna Ayyar 1937). Drainage as an insect control method was believed to
have been discovered because the water in an infested field smelled bad and farmers were primarily
concerned with eliminating the bad odor. Draining the field is reported to harden plant tissue
making the crop more resistant to insect feeding. Calcium is readily taken up when the field is
drained which may be an explanation for this observation (H.U. Neue, IRRI, personal communica-
tion).
Dryland adapted pests (which pass at least one stage in the soil) are highly vulnerable to
flooding. These pests include armyworms (Ingram 1927), grasshoppers (Katsumata and Nishikawa
1935), thrips (Commun 1937), ants (Dammerman 1929), white grubs (Berthet 1918), mole crickets
(Gonzales 1976), root aphids (Grist and Lever 1969), termites (Grist and Lever 1969), root weevils
(Tirumala Rao 1952), and seedling maggots (Dammerman 1929). Armyworms attacking a wetland
seedbed can be flooded with little effort if water is available. Flooding a dryland environment,
however, is normally not considered a rational alternative (Tjoa 1952).
Many wetland pests may be termed semiaquatic, and are controlled by alternate flooding and
draining. Semiaquatic insects include black bugs (Fernando 1961), planthoppers (Hinckley 1963),
gall midge (Yen et al., 1941), hispa (Bouriquet 1949), and most stem borers (Jepson 1954). Draining
minimizes these pests if carried out for 5-7 days but reflooding is necessary to control weeds and
prevent crop desiccation. Frequency of the action is important because alternative flooding and
draining causes high losses of nitrogen. The degree of control against these pests is enhanced by
other management factors acting at the same time.
Increasing the water level drives insects up plants to concentrate their numbers for more
effective chemical, biological or mechanical control (Ramakrishna Ayyar 1933). Sowing rice under-
water gives initial protection from hispa (Bouriquet 1949) but would inhibit tillering on modern
cultivars.
4.9 FERTILIZER
Stem borers (Subramanian et al., 1977), leaffolders (Jaswant Singh and Shahi 1984), planthoppers
(Uthamasamy et al., 1983), leafhoppers (Oya and Suzuki 1971), gall midge (Chelliah and Sub-
ramanian 1972), armyworm (Koyama 1966), root weevils (Bowling 1963)and leaf beetles (Prakasa
Rao et al., 1971) tend to do more damage and increase in number with fertilization, particularly
nitrogen. Nitrogen increases the nutrition of both plants and pests. Nitrogen augments plant growth
rates leading to softer tissues and quicker penetration by first instar stem borer larvae. This allows
more larvae to escape natural enemies (Nadarajan and Janandhanan Pillai 1985). Nitrogen
promotes tillering and the increased tiller density affects insects as described earlier. Fertilization
promotes even development of a crop in contrast to natural grasslands with varied tillering patterns
and consequently low insect incidence (van Emden 1966). Fertilization synchronizes pest develop-
ment and survival.
LITSINGER 561
The effects of well fertilized plants on insects are greater insect survival, increased tolerance to
stresses (pesticides, cold), large body size (greater fecundity), increased feeding rate (more
damage), and faster growth (more generations per crop) (van Emden 1966, Sogawa 1970, Tandon
1973, Pimentel and Goodman 1978). Some insect pests, such as thrips (Ghose et al., 1960), and
whorl maggot (IRRI 1980) become less abundant if nitrogen is applied.
Nitrogen accelerates plant physiological processes and the production of greater amounts of
volatile chemicals attracting more pests which locate their host by odor (Chandramohan and Jayaraj
1977). Nitrogen also reportedly causes, oryzanone, a stem borer attractant to exude from the leaves
(Seko and Kato 1950). Organic fertilizer also attracts seedling maggots (Dammerman 1929) and
white grubs.
The beneficial effect of nitrogen on plant yield outweighs the pest controlling effect of entirely
omitting its use (Oka 1983). Bigger plants from nitrogen application, allow greater tolerance of
insect damage (Otanes 1947). But splitting the application, first with a basal soil incorporation for
slow release, and using judicious amounts help meet the dual goals of high yields and low pest
incidence (Israel and Prakasa Rao 1968).
Phosphorous tends to increase abundance of stem borers but to a lesser degree than nitrogen
(Hirano and Ishii 1961, Ferreira 1983). Phosphorous is important in root development therefore
allowing the crop to tolerate root weevils (Tirumala Rao 1952).
Potassium however suppresses many pests (whorl maggot, green leafhopper, yellow stem borer,
brown planthopper, thrips, leaffolder) by lowering levels of plant sugars and amino acids as well as
leading to greater production of allelochemics, thicker cell walls, and greater silica uptake (Bas-
karan 1985).
Silica, provided by addition of slag, plowing under straw, or from river irrigation water, hardens
stems making the crop resistant to stem borer larval feeding (Sasamoto 1957). Zinc is reported to
minimize Elasmopalpus damage in dryland rice (Ferriera 1983) and bone meal is said to control gall
midge (Reddy 1967).
5. COMMUNITYWIDE CULTURAL METHODS
5.1 CROP ROTATION

Breaking the rice cycle with other crops as a method of insect control works best against species
with limited host ranges, limited dispersal powers, and long life cycles (Isely 1951). Crop rotation is
advocated to control gall midge (Krishnamurthy Rao and Krishnamurthy 1960), stem borers (Sauer
1939, Tian 1981), white grubs (Kalshoven 1981), termites (Ferreira 1983), grape colaspis (Rolston
and Rouse 1965), planthoppers (MacQuillan 1974), seed bugs (Uichanco 1921), and armyworm
(Am and Nurullah 1977). Crop rotation should be carried out in synchronized plantings over a
wide area (Khan 1967). The degree of control increases each year until pest numbers stabilize. The
most serious rice pests in Asia are monophagous. But the majority of pests are oligophagous
preferring graminaceous plants. Choosing crops outside the grass family therefore affects the
greatest number of pest species. Polyphagous armyworms which build up on rice, can attack a
following crop such as berseem fodder (Majid et al., 1978). Maize, wheat, and barley (common
crops in rotation with rice) encourage mole cricket (Novelli 1914), smaller brown planthopper
(Hokyo et al., 1976), and Diatraea, Chilo, and Sesamia stem borers (Ingram 1927, Koyama 1964,
Lippold 1972).
However two rice pests in the Americas – colaspis and Elasmopalpus – have legume hosts.
In North America colaspis builds up on lespedeza and soybean planted the previous year (Bowling
1967). Minimal irrigation of lespedeza is recommended to lower colaspis populations (Rolston and
Rouse 1965). They further state that planting species of lespedeza resistant to colaspis prevents
buildup on rice, and delaying planting of rice and plowing under resulting weeds reduce the impact
of colaspis.
Clover rotated with rice is claimed to lower stem borer numbers in untilled stubble (Khan
1967). Otherwise tillage during land preparation of any rotated crop is the significant mortality
factor affecting stem borer larvae, not the rotated crop species (Zhan 1977).
5.2 CROP AREA

Most insect pests undergo a period of aerial dispersal in their life cycle but have limited ability in
locating a rice crop. The larger the area grown to rice, the greater is the probability that dispersing
insects will descend on a rice crop (Loevinsohn 1984). Most pests cannot actively search out a crop
and die if they do not land on a suitable host. Increasing the crop area has been said to favor stem
borers (USA 1977) and brown planthopper (Kalode 1983). Continuously cropped rice gardens of
less than several ha escape insect pest buildup (Pantua and Litsinger 1981) but corporate ventures
of the scale of several hundred ha – C.P. Brewer, Solomon Islands (MacQuillan 1974, Ho and Taro
1985) and Philippine Packing, Mindanao – succumb to increasing pest pressure within 5-10 years.
But as the rice area decreased in Japan, green leafhopper and yellow dwarf increased as grassy
fallows favored their overwinter survival (Nakasuji 1974). Polyphagous mobile pests however are
least affected by crop area (Squire 1933).
5.3 NUMBER OF RICE CROPS PER YEAR

A dry season fallow typical of rice growing areas in the tropics lowers the carrying capacity of
rainfed environments (Loevinsohn 1984). Root weevils (Tirumala Rao 1952) and Scirpophaga
innotata (van der Goot 1948) are highly adapted tropical rainfed species surviving the dry season in
the soil or stubble but are eliminated during tillage of a dry season irrigated crop.
Dry season cropping of photoperiod-insensitive cultivars with irrigation extends the temporal
availability of rice. Continuous cropping throughout the year allows exponential insect population
growth (Loevinsohn 1984). The following pests are reported to be encouraged by sequential rice
cropping – whorl maggot (Kalshoven 1981), leaffolders (Tu et al., 1985), black bug (Krishna Ayyar
1929), leafhoppers (Chowdhury and Mukhopadhyay 1973), planthoppers (Oka 1979), thrips (Tu et
LITSINGER 563
al., 1985), and stem borers (Khan and Murthy 1955). Pest species that are particularly favored are
those that develop on all crop growth stages and are monophagous — Nilaparvata lugens (Kalode
1983), Nephotettix virescens (Anjaneyulu et al., 1982), Scirpophaga incertulas (Tian 1981), and
Maliarpha separatella (Appert 1967). Gall midge, now also believed to be monophagous (Gagne
1985), is increasing in abundance in dry season irrigated crops (Japan 1984).
On the other hand, continuous rice also sustains natural enemies and behaves as a perennial
crop (Balasubramanian and Ooi 1977). Increasing the number of rice crops per year is reported to
lower stem borer populations (Tsou 1947). Stem borers have longer generation times than most rice
pests and much of their life is spent as relatively immobile larvae. Many are removed and killed
during frequent harvests. One report from China claims lower pest incidence in triple than double
cropped rice due to the shorter field duration in triple crops (Wong and Kiang 1934). However
insect vectors and virus diseases normally build up to unmanageable levels if more than two rice
crops are grown.
5.4 PLANT MATURITY

Traditional, photoperiod-sensitive cultivars can only be grown during long day lengths of the wet
season. High yielding, photoperiod-insensitive cultivars shorten the maturation time of rices and
with irrigation allow yearround cultivation in the tropics (Balasubramanian and Ooi 1977). Early
maturing cultivars shorten Geld time, limiting the number of pest generations on each crop. Insect
numbers increase exponentially with each generation, therefore shortening crop maturation by only
two weeks can have a profound effect (Loevinsohn 1984).
Early maturing cultivars have a reduced vegetative stage. Therefore the negative effect of early
harvest has greatest impact on stem borers (Dinther 1971, Agyen-Sampong 1982), armyworms
(Alam and Nurullah 1977), gall midge (Patel et al., 1957), leaffolders (Lippold 1972), white grubs
(Chowdhuri 1966), leafhoppers (Misra 1920) and planthoppers (Kiritani 1979) which colonize the
vegetative stage and can develop over succeeding growth stages. Use of early maturing cultivars as
an escape mechanism is particularly effective if planted after a rice-free period induced by winter
or a dry season. The effect can be made even more pronounced by early planting and methods that
hasten maturity such as dense planting and reduced fertilizer (Isely 1951).
A shortened vegetative stage, however, limits the capability of the crop to compensate from
pest damage. Highest yield losses from insects have been recorded in the earliest maturing rice
cultivars (Litsinger et al., 1987). As a rule, early maturing cultivars are low yielding (Ghose et al.,
1960), but provide greater flexibility for crop managers to shift planting times to avoid seasonal pest
incidence. But in the tropics higher yields are attained by growing more crops per year, thus early
maturity is often desired. Delayed transplanting at the beginning of the monsoon season using early
maturing cultivars causes a suicidal flight of stem borers while still allowing for a timely harvest.
Having a contiguous area planted to a mosaic of different fields of early, medium, and long
maturing cultivars, breeds insect pest problems through staggered plantings (Tian 1981). In Taiwan,
mixing early (4 month) and late (7 month) maturing cultivars in the same field resulted in high
yellow stem borer numbers in the late crop (Shiraki 1917). If an area is dominated by a single-crop
planted to photoperiod-sensitive cultivars higher damage often occurs from stem borers than a
double crop using early maturing cultivars (Torii 1971).
5.5 PLANTING TIME

The effects of shifting the planting time of the rice crop are more highly site specific than perhaps
any other cultural control method. Generalizations are difficult as a shift of planting time of 1 week
is sufficient to significantly increase or decrease insect numbers (Jepson 1954). Both early or late
plantings may miss peak abundance of the damaging pest stage. Early plantings minimize gall midge
(Prakasa Rao 1975), thrips (China 1976), grasshoppers (Mochizuki 1957), leaf beetles (Breniere
1966), stem borers (Khan 1967), root weevils (Isely and Schwardt 1934), leafhoppers (Shukla and
Anjaneyulu 1981b), planthoppers (O’Connor 1952), seed bugs (Srivastava and Saxena 1967), whorl
maggot (Mochizuki 1957) and caterpillars (Okamoto and Abe 1960).
Late plantings likewise minimize many of the same pests-gall midge (Mallik and Behera
1965), thrips (Yau-i 1980), leaf beetles (Frappa 1938, Hirao 1978), stem borers (Tsutsui et al., 1957,
Sauer 1939), root weevils (da Silva Martins 1976), leafhoppers (Kiritani 1981), planthoppers
(Okamoto et al., 1967), seed bugs (Kiritani et al., 1961), leaf miner and black bug (Mochizuki 1957),
and caterpillars (Ishikura 1961).
Pests for which only early plantings resulted in low populations were midges (Grigarick 1984),
seedling maggots (van der Laan 1951), leaffolders (Okamoto and Abe 1960), whorl maggots
(Okamoto and Kashihara 1954) and armyworms (Hutson 1920). Pests for which only late plantings
were recorded as offering control were white grubs (Chowdhuri 1966), root aphids (Tanaka 1961),
leafminers (Ito et al., 1970), and grasshoppers (Mochizuki 1957).
The basis for planting time shifts is in the cyclical occurrence of insect stages, as not all stages
injure the rice plant. Shifting the planting date will disadvantage the pest when the prevalent insect
stage is vulnerable to air or water temperature extremes, heavy rainfall, a nonpreferred crop growth
stage, or the abundance of natural enemies (Teetes 1981). Although there are exceptions, often early
plantings of a first or single crop escape pest colonization (Pimentel and Goodman 1978) while late
plantings of a first crop are benefited by high levels of natural enemies and delayed plantings of a
second crop by suicidal flights. Planting time interactions are greatest if carried out over large areas
and against monophagous pests that attack one growth stage or are highly seasonal in appearance
(Isely 1951). Shifting planting time has little effect in an asynchronous area (Logothetis 1951). A
disadvantage is that shifting a planting date may reduce the number of crops that can be grown
annually (van der Goot 1948).
An effective use of planting time is to strategically time an areawide rice planting after a dry
season. The most classical example was practiced in Java in the 1920s and 1930s where planting after
the prolonged dry fallow period was delayed to control Scirpophaga innotata (van der Goot 1948).
S. innotata underwent a predictable aestivation period that was broken by the first monsoon rains.
Delayed planting until after the moths emerged and died — termed a suicidal stubble flight — sup-
pressed the stem borer for the entire crop season.
LITSINGER 565
On the other hand, timely planting with the onset of monsoon rains allows the crop to mature
past the early growth stage before pests can colonize the crop. Rain breaks the dry season dormancy
of the gall midge and planting early allows the crop to escape the vulnerable tillering period. Low
gall midge damage also occurs if the crop is planted later than normal because by that time its
parasites have become abundant. With photoperiod sensitive cultivars, late planting have a
reduced tillering period and less gall midge damage (Hidaka et al., 1974).
In temperate regions, S. incertulas emerge from field stubble sooner and more synchronously
than C. suppressalis which emerges from straw stored under various conditions and thus at various
temperatures. Two effects of shifting planting time were identified — (1) the timing of emergence of
overwintered moths in relation to the crop growth stage and (2) the stage of development of the
second generation larvae at harvest. Greatest reduction in stem borers occurred if both effects
acted negatively on the same crop. The shift in the 1950s to earlier planting in southern Japan to
avoid typhoons was studied intensively with regard to changes in pest abundance (Ishikura 1964).
The rice crop is particularly vulnerable to stem borers during tiller elongation and panicle exsertion
(van der Goot 1925, Tian 1981). Timing the crop so that eggs are laid either before or after these
stages substantially reduces damage (Suenaga 1966). Early planting minimized both stem borer
species that emerged to oviposit on a crop before or after tiller elongation. Heavy mortality also
occurred if harvest was timed when stem borer larvae were too small to enter diapause (Ono et al.,
1972).
Early planting in Japan increased green leafhopper and the rice dwarf disease they transmit.
They overwinter as mature nymphs in grassy fallow areas, and the crop is transplanted from covered
seedbeds before the nymphs molt. Colonizing adults then attack a favored rice crop instead of a
fallow.
The green leafhopper is highly seasonal in South Sulawesi, Indonesia, thus the tungro virus
disease it vectors is controlled in an irrigated double crop area carrying out strict planting schedules
(Manwan et al., 1985).
When only a portion of the farmers plant early and others delay, this creates staggered planting
and increased pest buildup (Oka 1979). Communitywide coordination therefore is necessary.
Shifting planting time also means growing the crop under different weather conditions and
often planting early in a rainfed area increases risk from drought or typhoons. An early planted
rainfed crop may take advantage of the flush of mineralized nitrogen which could either mean
greater tolerance to pest damage or make the crop more susceptible to stem borers. Although the
crop is more susceptible to stem borers few may find the crop.
5.6 SYNCHRONOUS PLANTING/FLOWERING
Staggered planting ensures greater survival of rice pests yearround by extending the temporal
availability of rice as a host in a farm community (Ramakrishna Ayyar 1933, Loevinsohn 1984).
Asynchrony of harvest in the community lessens the risk of crop failures due to typhoons or other
calamities. In Japan the introduction of early planting caused staggered planting and green leafhop-
pers increased five fold (Kiritani et al., 1970).
Synchronous planting allows a planned non-rice break to occur during the year to interrupt pest
cycles. Planting of contiguous crops should occur within a time span less than the generation time
of rice pests and over a spatial scale larger than the effective dispersal range, normally in the order
of 1 km. Therefore a minimal area of 1 km2 (314 ha) should be considered. Often synchronous
planting on an area-wide basis is enforced bylaw (Logothetis 1951, Khan and Murthy 1955, Manwan
et al., 1985). Synchronous planting is advocated for management of planthoppers (Oka and
Bahagiawati 1984), leafhoppers (Basu et al., 1976), stem borers (Chen et al., 1968), gall midge (Li
1982), leaf beetles (Iso 1954), caseworm and caterpillars (IRRI 1983) and mites (Ho and Lo 1979),
and root weevils (Tucker 1912). Monophagous pests are most affected by synchronous planting.
Also planting cultivars of differing maturities to ensure synchronous flowering reduces panicle
feeding pests (Uichanco 1921, Kiritani 1971).
On the other hand, asynchrony maximizes populations of natural enemies (Lim 1970b). How-
ever synchrony is a more reliable control measure than natural enemies especially in preventing
virus disease transmission by insects.
5.7 TRAP CROP

The trap crop technique relies on the attraction of insect pests to plantings other than the main rice
crop (Isely 1951). Timing is important in utilizing a trap crop (Jepson 1954) as the pests should not
be allowed to reproduce. A trap crop should not sacrifice field area.
The Chinese have used the trap crop technique to control stem borers by planting many small
fields comprising 0.7 to 5.0% of the rice area ahead of the main crop and increasing their attraction
by using susceptible cultivars, high amounts of nitrogen, and lights (Cheo et al., 1964, S-mei 1976,
Zhan 1977). Stem borers are controlled by manually removing egg masses from the trap crop plants
or with insecticide (van Dinther 1971). Trap crops were also used in Java to attract the highly
synchronized emergence of Scirpophaga innotata moths (Li 1961).
It is important to plant a trap crop over an area large enough to attract the resident pests. Trap
crops in India occupying too small an area (200 m2) did not show promise (Khan 1967).
Another successful method used in Japan was to delay spring land preparation of the weedy
fallow until after emerging green leafhopper and small brown planthopper adults laid eggs (Kono
et al., 1972). This method requires perfect timing. Leaving small patches of weeds to attract seed
bugs also has been used (Corbett 1930). The patches are frequently sprayed with insecticide while
fields are thoroughly weeded.
A final technique, only tested experimentally, is to plant 2-3 border rows of a field ahead of the
main crop to a highly susceptible cultivar to attract stem borers (Ghosh and Choudhury 1969),
brown planthoppers (Saxena and Justo 1984), or green leafhoppers (Saxena and Justo 1987). The
border areas are then sprayed. The problems with earlier plantings are that more extended water
LITSINGER 567
delivery is required, the pest-free fallow period is reduced, and synchronous planting is less
effective (Isely 1951).
5.8 FLOODING STUBBLE

In areas with sufficient water, harvested fields can be submerged to drown stem borers (Harukawa
1920), armyworms (Hutson 1920), and grasshoppers (Katsumata and Nishikawa 1935) that would
persist to attack a following crop. This method is particularly effective if carried out com-
munity wide.
Flooding the field after harvest has been used mainly for stem borer control. Yellow stem borer
is highly adapted to submerged conditions and is a serious pest of deepwater rice. The larva survives
only because of the triple membrane it builds to seal itself within the tiller (Shiraki 1917). It is
inadvisable to flood right after harvest unless the operation is combined with tillage to lacerate the
stubble allowing quicker entry of water (Tian 1981).
Hours and days rather than minutes are required to drown insects. Striped stem borer larvae
died within 1 hour in 45°C water but 28 hours were necessary at 30°C. The warmer the water, the
greater is the mortality (Harukawa 1920). However in many areas the stem borer larvae are in a state
of diapause and respire at very low levels. Reports from China state that flooding for at least
1 month is required to cause significant mortality of insects in dormancy (Li 1982). Very few
locations would have such quantities of water to manage pests. However if the flooding is done in
the spring after the larvae have terminated vernal diapause but before moth emergence, high levels
of mortality occur. Timing of the operation is obviously critical and the field should be underwater
for 1 week (van der Goot 1925).
Flooding the stubble before the monsoon rains is a way of initiating a suicidal Scripophaga
innotata flight. Again timing is critical because the larvae must pass 3.5 months in aestival diapause
before they become responsive. Flooding too early will have no effect. Flooding must be done only
several weeks before the main monsoon rains are expected and diapause is broken (Li 1972).
5.9 TILLAGE
Turning the soil over crushes insects, buries them, destroys their nests, and exposes them to
desiccation and predators. A moldboard plow gives best results particularly if combined with
harrowing to break up root masses (Khan 1967). Tillage can be performed most frequently in
dryland fields during land preparation or interrow weed cultivation of rice or crops preceding or
following rice. Soil insects prefer moderately moist soil. If the soil is wet, more insects will be within
the plow zone (Tjoa 1952). Dry season cultivation in the drylands is therefore of little benefit for
insect control as the insects have tunnelled deep into the soil. Soil-dwelling pests of dryland
rice-whitegrubs (Maxwell-Lefroy 1906), mole crickets (Novelli 1914), termites and ants (Dam-
merman 1929), root bugs (Headlee and McColloch 1913), and ants – become reduced in number
with each tillage operation. Dryland communities extensively cropped to annuals and where tillage
is practiced throughout the year have fewer soil insect pest problems than communities with large
areas of perennial grasses which remain a source of reinfestation.
Tillage of wetland fields is more limited and can be done only when the soil is moist, which
generally means after soaking during land preparation or after harvest (Chatterjee et al., 1979).
Tillage after rice harvest directly kills stem borers (Jepson 1954), armyworm pupae (Jhaveri 1920),
grasshopper eggs (Jones and Mackie 1913), black bugs (Shepard et al., 1988), and root weevils
(Kuwayama 1963). Tillage to destroy crop residues is of greater importance if more than one crop
is grown annually.
Tillage soon after harvest prevents unwanted crop regrowth (volunteer ratoon) which per-
petuates gall midge (Reddy 1967), leafhoppers (Hokyo 1975), and planthoppers (Sakanoshita and
Koide 1971), and virus diseases that may be transferred to neighboring seedbeds or younger fields
(Anjaneyulu 1975). Stubble plowdown kills the ratoon to stop pest cycles, particularly
monophagous pests, if carried out communitywide. Stubble plowdown should be combined with
synchronous planting for best results (Fernando 1967). Greatest benefit occurs in areas where more
than one rice crop per year is planted. In single cropping, high mortality normally occurs during the
long fallow with the exception of pests that undergo dormancy (e.g. stem borers and gall midge).
Turning over the stubble mat breaks the hibernacula of stem borer larvae allowing moisture and
pathogens to enter (Khan 1967). Undamaged stubble, even though plowed under, allows high stem
borer survival. The stubbles need to be broken open for greatest mortality (Khan 1967). Also, lower
stem borer survival is related to how long stubble is buried rather than depth of coverage (Shiraki
1917).
It is preferable agronomically to plow stubble under after harvest to increase soil organic matter
and recycle nutrients (Cleare 1927). But allowing the ratoon and weeds to grow is often done to
provide grazing for livestock during the off season. This practice allows insect pests to carryover
between crops. Tillage during land preparation can control dormant stem borer larvae before they
emerge as adults, but timing of these operations on a community wide basis is highly critical. If for
some reason tillage is delayed, pest problems worsen.
5.10 WEEDING
Most rice pests can develop on the common ricefield weeds, mostly grasses, that occur in fallow
fields, border areas, or in association with rice. Weed hosts act as a bridge between rice crops or
preferred stages of rice crops to sustain pest populations (Jepson 1954). Weedy rice fields often
have higher populations of some rice insects. Weeding is advocated for control of leaffolders
(Kalshoven 1981), leafhoppers (Abalos 1939), planthoppers (Oka 1979), seed bugs (Akbar 1958),
gall midge (Israel et al., 1970), caterpillars (Japan 1913) leaf beetles (Prakasa Rao et al., 1971), stem
borers (Catling and Alam 1977), black bug (Corbett and Yusope 1924), mealybug (Ramakrishna
Ayyar 1939), armyworms (Hutson 1920), caseworm (Shroff 1920), root aphids (Otanes 1947), root
bugs (Isely and Horsfall 1931), root weevils (Ishly 1975), colaspis (Rolston and Rouse 1965),
leafminer (Kuwayama 1950) and seedling maggot (Grist and Lever 1969). Dense vegetation is a
preferred habitat for leaffolder and stem borer moths. Weeds also interfere with insecticide
coverage. Hispa (Prakasa Rao and Israel 1970), seed bugs (Dammerman 1929) and water weevils
(Tucker 1912) build up on grassy weeds early in the season and later transfer to rice. Weeds
LITSINGER 569
therefore can act as a diversion but the effect is reversed once the weeds are controlled or senesce.
Armyworms prefer grassy weeds and transfer to rice after weeding operations (Grigarick 1984).
Early weeding therefore is important to kill the armyworms before they become large enough to
migrate. The gall midge does not occur in regions lacking wild rice alternate hosts.
Areawide weed control is a requirement for successful insect control as most pests are highly
mobile and can fly to distant fields. Good weed control also allows the rice crop to tolerate more
pest damage (Rolston and Rouse 1965).
5.11 HARVEST METHOD

The cutting height of the crop at harvest influences the control of stem borers. Panicle harvesting, a
traditional practice, allows 98% of the stem borer population to carryover to the next rice crop
(FAO 1979). Stem borer larvae descend to the base of the plants to pupate or to pass an unfavorable
season in larval diapause. How far they descend depends on the pest species and flood water depth.
Lowering the cutting height results in proportionally more larvae in the straw (Shiraki 1917). The
greatest degree of control occurs when the stubble is cut below 15 cm (Lippold 1972). The yellow
stem borer was claimed to be controlled in Taiwan when the stubble was cut at ground level (Jepson
1954). However, if the larvae were in diapause, ground level cutting only removed 14%(Islam 1976).
Harvesting before larvae descend below the cutting height is a practice to capture more larvae in
the straw (Li 1961).
5.12 STRAW AND STUBBLE DESTRUCTION

Management of postharvest crop residue is mainly directed at stem borers. Even though there is
normally a high natural mortality after harvest, small differences in numbers that carry-over from
one crop season to the next, can, through exponential increase, become significant. Burning straw
and stubble in the field is a common practice and is claimed to control stem borers (Commun 1934),
armyworms (Alam and Nurullah 1977), planthoppers (Oka 1979), and leaffolders (Korea 1973).
Piled straw burns thoroughly but standing stubble does not. Spreading the straw to allow the sun’s
heat to kill stem borers is just as effective as burning (Khan 1967) and allows natural enemies to seek
shelter to survive (Lim 1970) as well as conserve nutrients which would be lost in the smoke (Cleare
1927). Attempts to increase the burn of stubble by flame throwers (Israel and Prakasa Rao 1968) or
collecting more straw from neighboring fields (Khan 1967) are impractical. Weedy and direct
seeded fields produce better burns.
Often straw is stacked to be used as fodder, mulch, fuel, or in mushroom culture. Using straw
as fertilizer in greenhouses in Japan is claimed to help control striped stem borer (Kiritani 1979).
Stacked straw can be mixed with stems of maize or sorghum. Striped stem borer but not yellow borer
larvae will transfer to these plants that act as a trap crop (Shiraki 1917). Stacking straw in the open
overwinter confers greater mortality of stem borers particularly those exposed in the outer layers.
AS the straw is used, larvae in the newly exposed layers will be killed by the cold. In Japan, striped
stem borer moths emerge two weeks earlier in the spring from straw stacked in the open compared
to straw under a shelter (Ozaki 1940). Earlyemergence becomes a suicide flight as the seedbeds are
either covered or not yet sown. Another method is to cover the stacked straw with a cloth in the
spring to prevent moth emergence (Pang 1932).
Striped stem borers remain in the stubble at or below the ground level. A labor intensive
practice was to pull out the stubble by hand or by hand tool, roots and 211 (Shiraki 1917). A special
hand tool was developed in Japan to recut the stubble below ground level after harvest (Shiraki
1917). Stubble was also raked to kill stem borers (Khan 1967). Over 90% of striped stem borers were
controlled by these intensive methods. Stubble destruction before winter also reduced the number
of tillage operations needed in the spring.
5.13 RATOONING
A ratoon extends crop availability for additional pest generations. The higher the main crop is cut,
the sooner the ratoon appears, as the new tillers emerge from axillary buds. Reproductive stage
pests are favored in a ratoon. Stem borers survive in the stubble. Leaffolders thrive in a ratoon crop,
perhaps because of the large or physiologically older leaves (Chen and Wang 1978). Gall midge is
favored by tillering but normally parasite numbers are also high and temper the infestation (Rao
and Murthy 1961). Planthoppers and leafhoppers would be expected to multiply, but as their
predators and parasites are also spared, build up is normally curtailed. Seed bugs can increase as
they are long lived and can survive between the ripening stages of the main crop and ratoon.
The most serious pest problem is insect-vectored virus diseases-yellow dwarf and tungro-
which can be contracted late in the main crop without visible symptoms but later appear in a ratoon
(Prakasa Rao and John 1974). Vegetative pests — whorl maggot, caseworm, defoliators (Naranga,
Rivula) — are disadvantaged (dela Cruz and Litsinger 1988) because the vegetative stage is of
minimal duration in a ratoon.
5.14 GRAZING
Grazing livestock in a recently harvested rice field causes stem borer larvae lingering in the stubble
to be trampled. The beneficial effect is on the subsequent crop and is proportional to the stocking
rate and size of animal. Studies in the USA (Douglas and Ingram 1942) and Australia (Li 1972)
claim 57-75% control. But many cattle died from an unknown cause when grazed on rice during an
outbreak of fall armyworm (Brown and Mohamed 1972). Grazing also helps to remove weeds
(Hutson 1930).
6. SYNOPSIS
A wide range of mechanical, physical, and cultural methods are potential practices, and their use
will depend on existing alternatives and local conditions. A synopsis of the most practical methods
is presented.
Handpicking and netting are labor intensive but can be useful in certain situations. Because
they are food items, white grub adults, locusts, and mole crickets may be rewarding for the poorer
LITSINGER 571
farmers. It may be appropriate to net caterpillars, armyworms, and caseworms when abundant in
seedbeds or small dryland fields. Complete control is not necessary in these situations as the
seedlings can normally recover from moderate infestations. Stem borer egg masses pose little threat
to a seedbed but it may be attractive to remove them by hand from the field before stem elongation,
particularly eggs of those species that oviposit openly. Egg masses are most easily seen on young
plants. Egg parasites should be spared by allowing them to escape. Collecting egg masses just before
panicle exsertion is probably futile as they are difficult to see in fully grown plants.
Roguing is not practical as the damage is already done. Shaving rice bunds would not be done
solely to control insect pests but is a normal practice of most farmers for rodent and weed control.
It would be an exceptional case where constructing a moat or kerosene barrier to protect a seedbed
from invading armyworms would be practical. Nursery boxes are used in temperate regions primari-
ly to start an early crop, and insect control is secondary. Rollers and other heavy equipment are
generally unavailable and thus of improbable use of small scale farmers.
Kerosene or oil applied to flooded fields is expensive, phytotoxic, harmful to aquatic natural
enemies, and requires additional effort to dislodge pests from foliage. The treatment would have to
be repeated before any significant population reduction would occur. Light traps are expensive and
capture only a small proportion of a pest population.
Changing field geometry and intercropping are done for reasons other than controlling insects
and are usually not a viable pest control option. Changes from transplanting to direct seeding are
done because of labor costs, not to alleviate insect damage. But in a tungro infested area, farmers
may wish to establish dapog seedbeds away from infested fields.
In irrigated areas low yields are expected if transplanting of modern varieties is delayed beyond
one month. Irrigated farmers therefore have little incentive to transplant older seedlings. Rainfed
farmers, however, plant low tillering traditional cultivars that can be transplanted from 2 to 3
months old seedbeds without yield loss. Rainfed farmers transplant when the rains occur and
cannot risk delaying transplanting to achieve better insect control. Pruning seedlings is a practice
for traditional varieties but is usually done to prevent lodging and promote tillering. There is some
scope to support the practice of clipping foliage from rice plants attacked by thrips, hispa, or
leaffolders and to use the foliage as animal forage. Yield may be stimulated if pruning occurs before
tillering. Moderate plant densities appear to be a practical target for best agronomic yield and
lowest insect pest numbers in general.
Draining a field to control aquatic insect pests has merit if water is less expensive than other
control measures. Irrigation costs are often high making this option unattractive. Fields should be
reflooded within a few days, for weed control, if drained before canopy closure.
Excessive use of fertilizer invites insect and disease problems. Therefore optimal levels of
fertilizer with split applications are warranted.
It is impractical to consider regulating crop area. Ecologically, flooded rice is cultivated in
contiguous lowlying areas in which dryland crops would become waterlogged in the rainy season.
The effect of limiting the area planted on insect control is minimal except at low crop densities.
The most profound cultural control practice is to isolate the rice crop in time. This is ac-
complished by growing no more than two rice crops annually. Early (100-130 days) cultivars should
be grown. Cultivars less than 100 days seed-to-seed are too sensitive to stress. Crop maturity greater
than 140 days encourages stem borers and brown planthopper. If possible, the crops should be
planted at times of least pest abundance and the rice-free period should occur during the driest part
of the year so that stubble will not regrow, thereby killing insect pests and virus diseases. The stubble
should be plowed under during the rice-free period to control stem borers. The best effect is to plow
under the rice stubble before planting a non-rice third crop, and again before the wet season,
followed by prolonged flooding. Flooding should last one week during land soaking to ensure
greatest stem borer control at the start of the season. Each of the two rice crops should be planted
synchronously (within three weeks) over an area of more than 300 hectares. In temperate regions
small trap crop seedbeds may be planted at the beginning of the rainy season and then plowed
under.
Management of the cutting height and straw is directed towards stem borer control. If rice is
double cropped with early maturing cultivars and the stubble is plowed under between crops, then
cutting height and straw management are not important considerations. However if the rice crop is
long maturing then management of crop residues becomes important. The stubble should be cut
lower than 15 cm and the straw spread and dried in the sun. If the stem borer undergoes dormancy
in the off season then wet season plowing and flooding are advisable. Populations of insect pests are
not static and can adapt to stresses from pesticides (insecticide resistance), resistant cultivars
(biotypes), parasites (encapsulation), and predators (change behavior to avoid capture). Rice
insect pests have also adapted to changes in cultural practices.
Local biotypes of striped stem borer exist in various regions in Japan. These biotypes emerge
at different times in the spring in response to prevailing planting schedules rather than spring
temperatures per se. Moths began to appear earlier in areas that switched to early planting (Fukaya
1951). When Japanese farmers changed to early maturing cultivars, striped stem borers emerged
from hibernation earlier (Ishikura 1956). Longer maturing biotypes of striped stem borer developed
on early maturing late season rices in order to time their development so that the larvae were in a
pre-diapause stage right before harvest, thus ensuring the greatest off season survival. Ill-adapted
biotypes emerge near harvest to constitute a suicidal third generation that perishes overwinter. In
addition, when stubble heights were lowered, natural selection favored stem borer larvae that
descended below the prevailing cutting height (DuPort 1925).
However, cultural control practices that are preventative rather than corrective form a first line
of defense. Combining a rice-free period with crop rotation of botanically unrelated crops, for
example, poses a difficult obstacle for rice insect pests to overcome. However, mechanical, physical
and cultural methods will not by themselves prevent economic damaging levels from being reached.
These practices along with resistant cultivars and judicious use of chemicals to conserve natural
enemies form the foundation of insect pest management in rice.
LITSINGER 573
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7
Predators and Parasitoids of Rice Insect Pests
P. A. C. OOI,
CAB International Institute Of Biological Control
(CIBC) Malaysian Station, Batu Tiga
Selangor, Malaysia
B. M. SHEPARD
1. APPRAISAL AND DIRECTION

LONG HISTORIES OF RICE CULTIVATION in many parts of the world have allowed stable relationships
to evolve between rice insect pests and their natural enemies. Natural biological pest control
managed insect pests long before the advent of synthetic insecticides. That action of indigenous
predators, parasitoids and insect pathogens forms the cornerstone of modern Integrated Pest
Management (IPM) programs in rice.
When modern improved rice varieties were introduced in the mid-l960s, chemical insecticides
were embraced as part of the package of technologies of the green revolution. With increased
adoption of new high yielding varieties, use of insecticides also increased. Planting resistant
varieties and using chemical insecticides were the primary insect control tactics.
The destruction of predators and parasitoids that followed insecticide misuse resulted in
resurgence of several rice pests including the brown planthopper (BPH), Nilaparvata lugens (Stal)
(Homoptera: Delphacidae) (Heinrichs and Mochida 1984, Kenmore, et al., 1984), green leafhopper
(GLH), Nephotettix spp. (Kobayashi l96l), and rice stem borers (Lim et al., 1980). Chronic
outbreaks of the brown planthopper in Indonesia in the mid 1980s were attributed to excesive use
of insecticides in rice fields. The result was the Presidential decree in 1986 banning use in the rice
586 PREDATORS AND PARASITOIDS OF RICE INSECT PESTS
environment of more than 57 insecticides (England 1987). The story of mismanagement of natural
enemies, particularly predators and parasitoids, has repeated itself many times before in other
crops, for example cotton in Peru (Boza Barducci 1972), cocoa in Sabah (Conway 1971), and oil
palm in Peninsular Malaysia (Wood 1964). Pest outbreaks were reduced following a conscious
effort to conserve the natural enemies in all these cases.
Classical and inundative biological control approaches tried so far have met with dismal
failures, and there is little potential for this approach in rice. Here we emphasize maximizing the
impact of indigenous natural enemies as an essential part of IPM programs. We believe biological
control of rice insect pests in tropical south and southeast Asia lies in the identification and
conservation of their natural enemies.
Because much of the scattered literature related to the biological control of rice insect pests is
anecdotal, with few quantitative or definitive studies, we make no attempt to include an exhaustive
literature search. Our objective is to focus on biological control issues and concepts drawn from
selected literature and our own experiences. This chapter covers arthropod predators and
parasitoids in rice in tropical south and southeast Asia and is oriented toward plant protection
specialists, particularly those working in rice. The important insect pathogens are covered in
chapter 8 and the taxonomy of predators and parasitoids is included in chapter 2.
2. CASE STUDIES
In the past, biological control of rice insect pests emphasized the importation of parasitoids and/or
mass rearing of egg parasitoids for release at regular intervals. Little attention was paid to the use
of predators. Two groups of insect pests were involved, stem borers and defoliators. The examples
of attempts at biological control of rice insect pests listed in Table 1 were selected to show the
limited successes of introduction, augmentation, and inundation of natural enemies in controlling
pests.
2.1 EXOTIC PARASITOIDS FOR EXOTIC PESTS

In two cases, introduction of exotic parasitoids resulted in successful biological control (Table 1).
The first was the biological control of Chilo suppressalis (Walker) (Lepidoptera: Pyralidae)
(referred to as C. simplex) in Hawaii. C. suppressalis had been accidentally introduced into Hawaii,
probably in rice straw used for packing. Three parasitoids were introduced in 1929, an egg
parasitoid, Trichogramma japonicum Ashmead (Hymenoptera: Trichogrammatidae), from Japan
and two larval parasitoids, Eriborus sinicus (Holmgren) (Hymenoptera: Eulophidae) and Bracon
chinensis Szepligeti (Hymenoptera: Braconidae), from China. All three parasitoids helped reduce
the infestation of this stem borer (Sweezey 1931).
In the second case, reported by Hinckley (1963), the leaffolder, Marasmia (Susumia) exigua
(Butler) (Lepidoptera: Pyralidae), had arrived in Fiji sometime before 1944. A larval parasitoid,
Trathala flavoorbitalis (Cameron) (Hymenoptera: Ichneumonidae) was introduced from Hawaii.
Apparently, the introduction reduced leaffolder population density.
OOI AND SHEPARD 587
TABLE 1
Some examples of past attempts at classical, augmentative and inundative biological control of rice insects pests
Pest group/species Parasitoid species Origin Introduced into Year Stage Status*
attacked
Stem borers
Chilo suppressalis T. japonicum Japan Hawaii 1929 egg S
E. chinensis China Hawaii 1929 larva S
E. sinicus China Hawaii 1929 larva S
Chio polychrysus P. claripalpis Trinidad Malaysia 1951 larva F
Scirpophaga T. japonicum Japan Philippines 1934 egg E
incertulas S. inferens India Philippines 1966 larva E
Chilo suppressalis Trichogramma sp. Philippines Japan 1929 egg F
Chilo polychrysus T. japonicum indigenous Malaysia 1931 egg F
Defoliator
Susumia exigua T. flavooribitalis Hawaii Fiji 1928 larva S
* Status of biological control:
S - successful biological control
F - failed to provide control
E - established but no control exerted
However, not all attempts at classical biological control of rice pests have sucessful conclusions.
The sucesses mentioned here were dependent on the types of pests involved. Both were exotic
pests, suitable candidates for classical biological control.
2.2 EXOTIC PARASITOIDS FOR INDIGENOUS PESTS
Several attempts have been made to introduce exotic parasitoids to control indigenous rice insect
pests in south and southeast Asia (Table 1). One case involved the introduction of Paratheresia
claripalpis (ven der Wulp) (Diptera: Tachinidae) to control Chilo polychrysus (Meyrick) (Lepidop-
tera: Pyralidae) in Malaysia. P. claripalpis is a parasitoid of sugarcane stem borers in the new world.
The rice environment and the rice stem borer it was meant to control were very different in
Malaysia. The tachinid apparently failed to adapt and, after the release of 8,187 adults, none were
recovered.
A more recent introduction of parasitoids from a different host to control a rice insect pest also
failed. B. M. Shepard introduced two egg parasitoids to control the Malayan black bug,
Scotinophara coarctata (F.) (Hemiptera: Pentatomidae) on the island of Palawan in the Philippines.
The parasitoids, Psix lacunatus Johnson and Masner (Hymenoptera: Scelionidae) and Telenomus
cyrus (Nixon) (Hymenoptera: Scelionidae), were collected from the eggs of an economically
unimportant pentatomid, Scotinophara latiuscula Breddin (Hemiptera: Pentatomidae) on the is-
land of Luzon. A total of 12,840 P. lacunatus and 19,330 of T. cyrus were released in 1985-1987; to
date, none have been recovered.
While a few introductions of parasitoids from a different species of pest related to the target
insect have been successful, examples are not common and none are known in rice. The best known
case is the biological control of the coconut moth in Fiji (Tothill et al., 1930). In this case a larval
tachinid parasitoid, Bessa remota Aldrich (Diptera: Tachinidae) which parasitised Zeuxippa
catoxantha Hampson (Lepidoptera: Zygaenidae) in Malaysia was introduced to control Levuana
iridescens Bethune-Baker (Lepidoptera: Zygaenidae) in Fiji. Both L. iridescens and 2. catoxantha
caused similar damage to coconuts and it is likely that the parasitoid had little difficulty adapting to
a similar environment.
Kamran and Raros (1971) reported the introduction of Sturmiopsis inferens Townsend (Dip-
tera: Tachinidae) from India in an attempt to control rice stem borers in the Philippines. Although
the parasitoid was established, it is not an important mortality factor. This is another example of
the failures experienced with the classical biological control approach of indigenous rice insect
pests in tropical south and southeast Asia.
In 1929, Shibuya and Yamashita (1936) imported an egg parasitoid from the Philippines into
Japan and experimented on its mass rearing and release for the control of C. suppressalis (known as
C. simplex). This introduction was carried out despite an existing indigenous egg parasitoid,
Trichogramma japonicum Ashmead (Hymenoptera: Trichogrammatidae). The introduced
parasitoid could be differentiated from the black native one by its yellowish body color. The
researchers found that the imported egg parasitoid did not perform as well as the indigenous one,
and concluded that there was no possibility of success in using the imported species. This example
suggests that the introduction of additional species when there are already effective indigenous
parasitoids is inappropriate.
The above cases suggest that there is no justification for importations of parasitoids. Most rice
insect pests in tropical south and southeast Asia already are under biological control by indigenous
predators and parasitoids. A more viable approach may be to determine the most important
natural enemies and investigate their possible use against rice pests in other continents.
2.3 INUNDATIVE RELEASES

The case study for this approach is taken from an attempt in the early 1930s to rear T. japonicum
for the control of C. polych rysus in Malaysia. Having recognized that egg parasitoids are important
mortality factors, Miller and Pagden (1930) recommended that large numbers of T. japonicum
(referred to as T. nanum) be reared and released. Up to 1,300,000 parasitoids were released per
acre. However, Corbett and Pagden (1933) concluded that T. japonicum was not effective, even
when released in large numbers. Pagden (1932) suggested that two other egg parasitoids, Tetras-
tichus schoenobii Ferriere (Hymenoptera: Eulophidae) and Telenomus rowani (Gahan)
(Hymenoptera: Scelionidae) were more effective.
The failure of inundative releases and subsequent realization of the importance of existing
biological control of the rice stem borer suggests that this approach is not appropriate for in-
digenous rice insect pests in the humid tropics. This conclusion concurs with that of Nickel (1964).
OOI AND SHEPARD 589
2.4 CASUAL INTRODUCTIONS

An examination of introductions of natural enemies into the Philippines indicates that many of them
were introduced without adequate evaluation. Baltazar (1964) reported on the introduction of T.
japonicum from Japan in 1934. Nickel (1964) pointed out that the parasitoid was already an
indigenous species in the Philippines. This species is also indigenous in Thailand (Yasumatsu et al.,
1982) and Malaysia (van Vreden and Abdul Latif 1986).
Casual introductions result in wasted effort and may create problems which seriously retard
advancement in the field of biological control. For example, the casual introduction of house crows
to control insect pests on coffee in Malaysia (Carey 1903) led to the establishment of a nuisance.
Several other examples of the adverse consequences of this careless, non-scientific approach can be
found. These incidents have haunted attempts at biological control of insect pests in Malaysia, and
probably other developing countries.
While studies of the ecology of both pests and natural enemies are recommended, unnecessary
prolongation of such studies may seriously hamper IPM efforts. The procedure recommended by
Nickel (1964) is a useful guide. It involves an extensive survey of natural enemies of the pests
followed by detailed ecological studies. Studies of rice insect pests and their natural enemies should
be a continuous part of a research program, both to identify important natural enemies and to
promote practices that enhance their conservation.
From the evidence so far, biological control in the classical sense is suitable only for exotic rice
pests. Attempts to enrich parasitoid species and inundative releases mostly have met with failures.
Casual introductions of beneficials in the past, probably stimulated by sucessful classical biological
control in the United States, have resulted in more failures, and may have discouraged plant
protection scientists from pursuing other approaches to biological control, such as conservation.
Most rice insect pests in tropical south and southeast Asia are indigenous and have their own
indigenous natural enemies.
3. ROLE OF PREDATORS AND PARASITOIDS

Although species diversity and total numbers of predators and parasitoids in tropical rice are
impressive, generalizations about the precise role and relative importance of individual species are
difficult. Clearly, abundance shifts seasonally and geographically, but in general only relatively few
species have been shown to impact heavily on target insect pests. This is primarily due to the fact
that little definitive work has been done and techniques that allow researchers to dissect these
communities of parasitoids and predators and determine their individual roles are not available.
Their collective role can be clearly demonstrated by observing outbreaks of insect pests after a
broad spectrum insecticide has removed the natural enemy community. A comprehensive review
of techniques for evaluating arthropod natural enemies has been published by Luck et al., (1988),
although relatively few of the techniques have been used in rice. Examples of specific techniques
for evaluating predators and parasitoids are discussed by Shepard and Ooi (in Prep.).
We have selected three groups of pests to illustrate the role of predators and parasitoids; stem
borers, leaffolders, and leaf- and planthoppers. They were selected because their natural enemies
have been more extensively studied than those of other rice insect pests.
3.1 RICE STEM BORERS
3.1.1 Stem Borer Parasitoids
Before chemical insecticides were introduced, natural enemies played an important role in main-
taining stem borer populations at low levels (Yasumatsu 1967a). The major focus of early work on
natural enemies was on parasitoids. Nickel (1964) reported over 100 species of stem borer
parasitoids. Of the 99 species of natural enemies reported by Yasumatsu (1967b) most were
parasitoids. Other authors have identified many stem borer parasitoid species (Kamran and Raros
1969, Rothschild 1970, Yasumatsu and Torii 1968). In general, egg parasitoids are considered the
most important; only three groups, namely, Telenomus spp., Tetrastichus spp. and Trichogramma
spp. dominate the complex (Catling et al., 1983, Delfinado 1959, Nickel 1964, Rothschild 1970,
Shepard and Arida 1986).
In a study in the Philippines in both direct seeded and transplanted rice, parasitism of yellow
stem borer, Sciropophaga incertulas (Walker) (Lepidoptera: Pyralidae), eggs was found to reach
more than 60% then declined with crop age (Shepard and Arida 1986). In Bangladesh, Catling
(1979) found an average of 75.5% of egg masses and 49.7% of eggs were parasitized. T. schoenobii
was the most important parasitoid. Catling et al., (1983) reported a similar situation for deepwater
rice in Bangladesh, where 61-89% of yellow stem borer egg masses were parasitized. The two most
important parasitoids were T. rowani, which was able to parasitize 64% of the eggs in parasitized
egg masses, and T. schoenobii, which destroyed 98% of the eggs. Subba Rao et al., (1983) reported
77% egg parasitism by Tetrastichus spp., Telenomus spp. and Trichogramma spp. in India. On the
IRRI farm in the Philippines, egg parasitism by T. schoenobii, T. rowani, and T. japonicum averaged
84, 42 and 24 per cent, respectively (Kim et al., 1986).
Several reports have suggested that T. schoenobii is the most important yellow stem borer egg
parasitoid (Rothschild 1971, Ooi 1974, Than Htun 1976, Catling et al., 1983, Shepard and Arida
1986) because it attacks several eggs during its development, a behavior not unlike that of a
predator.
Although the complex of larval and pupal parasitoids is large, they are not as abundant as egg
parasitoids (Table 2). Subba Rao et al., (1983) reported that the action of larval/pupal parasitoids
and microbial action caused as high as 58% mortality in the Warangal region of India. In our
experience, larval and/or pupal parasitism has not approached this level. More detailed records
can be found in Nickel (1964), Rothschild (1971), Yasumatsu (1967b), and Yasumatsu and Torii
(1968).
OOI AND SHEPARD 591
TABLE 2
Some common parasitoids of stem borers
Borer species Stage attacked

Egg Larva Pupa
S. incertulas Tetrastichus schoenobii Temelucha philippinensis Tetrastichus ayyari
Ferriere*** (Ashmead)* Rohwer*
Telenomus rowani Stenobracon nicevillei
(Gahan)** (Bingham)*
Trichogramma japonicum Bracon chinensis
Ashmead. * Szepligeti*
C. polychrysus T. japonicum * * Cotesia flavipes
Cameron* *
Tropobracon schoenobii
(Viereck).
C. suppressalis Telenomus dignus C. flavipes * Xanthopimpla stemmator
(Gahan)* (Thunberg)**
B. chinensis*
* common; ** important; ***very important
3.1.2 Stem Borer Predators

The importance of predators in natural control of stem borers has received little attention. Pantua
and Litsinger (1984) reported up to 65% of the eggs of yellow stem borer were consumed by
Conocephalus longipennis (de Haan) (Orthoptera: Tettigoniidae). Predation on eggs of the yellow
stem borer averaged about 30% during an entire season at the IRRI farm (Shepard and Arida
1986). The single most important predator was the grasshopper, C. longipennis. Although C.
longipennis has been reported to damage rice panicles (Rothschild 1971, Barrion and Litsinger
1987, van Vreden and Abdul Latif 1986), its predatory role is more significant. Rubia et al., (1990)
showed that C. longipennis could consume over eight yellow stem borer egg masses in three days.
The grasshopper population increased late in the season, when yellow stem borer is usually a more
serious problem.
Other predatory orthopterans such as Metioche vittaticollis (Stal) (Orthoptera: Gryllidae) that
feed readily on eggs of the striped stem borer, C. suppressalis, would not feed on yellow stem borer
egg masses (Rubia and Shepard 1987). Manti (unpublished information) found that Cyrtorhinus
lividipennis Reuter (Hemiptera: Miridae) had a similar habit, feeding on egg masses of striped stem
borer but not that of yellow stem borer. Than Htun (1976) reported that the wolf spider, Lycosa
pseudoannulata (Boesenberg and Strand) (Araneae: Lycosidae), confined in a small arena in the
laboratory killed up to 90% of 130 borer larvae in one day. This wolf spider is also capable of
attacking stem borer moths (Figure 1).
An anthocorid, Euspudaeus sp. was the only known important predator of the striped stem
borer in Japan (Oho 1954-55). This predator alledgedly attacked 90% of C. suppressalis adults in
rice straw.
Fig. 1. Lycosa sp. feeding on the moth of a rice stem borer.
A sizeable but seldom recognized predator community attacks first instar stem borers before
they enter the stem. Pena (1987) found very little correlation between egg mass density and
resulting damage by stem borer that could not be explained entirely by the action of egg parasitoids
or predators. Small predators such as Micraspis spp. and other lady beetles, carabids such as
Ophionea, ants, etc., attack and consume first stage stem borer larvae before these larvae enter the
stems. In laboratory trials, we have observed that the water bugs, Microvelia douglasi atrolineata
Bergroth (Hemiptera: Veliidae), Mesovelia vittigera (Horvath) (Hemiptera: Veliidae) and Lim-
nogonus fossarum (F.) (Hemiptera; Gerridae) feed voraciously on early stage stem borer larvae
(unpublished information). Larter (1955) observed Paederus fuscipes Curtis (Coleoptera:
Staphylinidae) attacking newly hatched stem borer larvae in the field.
3.13 Stem Borer Management Strategy

There is ample evidence that if stem borer attack occurs during the vegetative stage the rice plant
is able to compensate for damage by producing new tillers (IRRI 1987). More than 40% of the
tillers can be removed during the early vegetative stage with no significant effect on yield. This
OOI AND SHEPARD 593
factor is important in developing management strategy, because a buildup of stem borer predators
and parasitoids can be allowed early in the season without fear of yield loss. Insecticide use at this
time is usually not considered as a viable tactic. Later in the season, when the crop cannot
compensate as easily for damage, dense communities of arthropod natural enemies will be present.
In our judgement, losses by stem borers have been grossly exaggerated. Catling et al., (1978)
reported that crop losses due to stem borers in Bangladesh were lower than most estimates for rice
in India. There is evidence to show that the high yielding modern varieties are not as severely
damaged by stem borers as older traditional varieties (Balasubramanian and Ooi 1977). This is
likely due to the action of natural enemies and the re-tillering capacity of modern varieties which
have thinner stems that are less suitable for stem borer development. Khoo and Lee (1978)
suggested that the shorter crop development time of modern varieties may also reduce the in-
cidence of stem borers.
3.2 LEAFFOLDERS
3.2.1 Parasitoids of Leaffolders
The major parasitoids of leaffolder eggs in the Philippines are Copidosomopsis nacoleiae (Eady)
(actually an egg-larval parasitoid)(Hymenoptera: Encyrtidae) and Trichogramma spp. The larval
and pupal parasitoid complex is richer. Rao et al., (1969) listed 32 species of larval and eight species
of pupal parasitoids. Parasitism averaged about 40% on the IRRI farm and in nearby farmer’s fields
in Laguna, Philippines (Arida and Shepard 1990). From a total of 4,904 leaffolder larvae collected,
15 species of parasitoids belonging to nine families were found. The seasonal incidence of leaf-
folder larval parasitism is shown in Figure 2. Some of the most common parasitoid species
encountered in the Philippines are listed in Table 3.
Fig. 2. Seasonal incidence of leaffolder larval parasitism on the IRRI farm and
in nearby farmer’s fields. 1985.
TABLE3
Some common parasitoids of leaffolders (Cnaphalocrocis medinalis and Marasmia patnalis)
Egg parasitoids Larval parasitoids Larval parasitoids

Copidosomopsis nacoleiae Goniozus nr. triangulifer Xanthopimpla flavolineata
(Eady) ** Kieffer* Cameron*
Trichogramma japonicum Cotesia angustibasis Tetralichus ayyari
Ashmead** (Gahan)* Rohwer*
Cardiochiles philippinensis
Ashmead*
Macrocentrus philippinensis
Ashmead*
Trichomma cnaphalocrosis
Uchida*
Temelucha philippinensis
(Ashmead)*
*common; **important
32.2 Predators of Leaffolders

Kamal (l981) reported 70% leaffolder mortality by predators. The predator complex that feeds on
eggs and larvae of leaffolders is nearly the same as for stem borers. A notable exception is the
cricket, M. vittaticollis, which feeds voraciously on leaffolder eggs but will not attack eggs of yellow
stem borer (Rubia and Shepard 1990). Leaffolder egg predation averaged about 50% in both direct
seeded and transplanted rice (Arida and Shepard 1990). These levels were detected by placing
plants with leaffolder eggs in the field and recording the missing eggs two days later.
C. lividipennis also attacks leaffolder eggs. Because the egg chorion is left intact, predation by
this mirid is easily overlooked and underestimated. Although both lady beetles feed on eggs of
Fig. 3. Laboratory study to evaluate consumption of leaffolder (LF) eggs by two rice field
coccinellids, Synharmonia octomaculata (F.) and Micraspis crocea Mulsant.
OOI AND SHEPARD 595
leaffolders, in laboratory experiments Micraspis crocea Mulsant (Coleoptera: Coccinellidae) con-

sumed more eggs than another common rice field coccinellid, Synharmonia octomaculata (F.)
(Coleoptera: Coccinellidae) (Figure 3). Predation on newly hatched larvae is more difficult to
assess, but mortality caused by the same complex of predators that attack young stem borer larvae
must be high. An Ophionea sp. is shown eating a leaffolder larva in Figure 4. After a leaffolder rolls
the leaf, the pest is partially protected from predators. Parasitoids then play a more significant role
in suppressing populations.
Fig. 4. Ophionea sp. feeding on a leaffolder larva.
3.3 LEAF- AND PLANTHOPPERS

Greathead (1983) reviewed natural enemies of leaf- and planthoppers, and Subba Rao et al., (1983)
catalogued the important parasitoid species. Table 4 presents a list of the major predator and
parasitoid species. A substantial amount of attention has been given to parasitoids of leaf- and
planthoppers (Otake, 1977; Chandra 1980a, b; Bentur et al., 1981; Chang 1980; Subbao Rao et al.,
1983; Chua and Dyck 1982; Pena and Shepard 1986). However, our experiences and those of
Kenmore et al., (1984) have led us to believe that, in general, predators impact hopper populations
more strongly than do parasitoids. In at least one instance, failure to assess predation led to
conclusions that a decline in hopper population could not be due to natural enemies (Otake 1978).
TABLE 4
Some common predators and parasitoids of brown planthopper and green leafhopper
Hopper species Predator/Parasitoid species Stage of host attacked

Predators +
N. lugens Cyrtorhinus lividipennis Egg/Nymph
Reuter***
Lycosa pseudoannulata Nymph/Adult
(Boesenberg and Strand)***
Microvelia douglasi atrolineata Nymph/Adult
Bergroth**
Synharmonia octomaculata Nymph/Adult
(F.)**
Paederus fuscipes Nymph/Adult
Curtis* *
Nephotettix spp. L. pseudoannulata*** Nymph/Adult
C. lividipennis' * Egg/Nymph
Tetragnatha spp.* Adult
Oxyopes spp.* Nymph/Adult
Parasitoids +
N. lugens Oligositayasumatsui Egg
Viggiani et Subba Rao*
Anagrus spp.* Egg
Pseudogonatopus spp.* Nymph/Adult
Nephotettix spp. Gonatocerus spp.* Egg
O. yasumatsui* Egg
Haplogonatopus apicalis Nymph/Adult
Perkins*
Tomosvaryella spp.** Nymph/Adult
Pipunculus spp.** Nymph/Adult
* common; * * important; * * * very important
3.3.1 Parasitoids
Seasonal abundance and incidence of parasitism in nymphs and adults of brown planthopper, N.
lugens, whitebacked planthopper, Sogatella furcifera (Horvath) (Homoptera: Delphacidae) and
green leafhopper, Nephorettix spp. were monitored in Laguna Province, Philippines (Pena and
Shepard 1986). The major parasitoids of N. lugens were dryinids (Pseudogonatopus spp.,
Haplogonatopus spp. and Echthorodelphax fairchildi Perkins) (Hymenoptera: Dryinidae) and strep-
sipterans (Elenchus yasumatsui Kefune and Hirashima) (Hymenoptera: Strepsiptera). At 12 loca-
tions parasitism averaged below 10%. Host densities were normally low throughout the study.
OOI AND SHEPARD 597
Chandra (1980b) reported that parasitism rates by dryinids reached 40% when host densities were
high during the wet season (July-October) in the Philippines. Normally, parasitism levels were
similar to those reported by Pena and Shepard (1986).
Similarly rates of parasitism of the S. furcifera by dryinids and strepsipterans were low (Pena
and Shepard 1986; Chandra, 1980a, b). Parasitism of S. furcifera by strepsipterans reached some-
what higher levels (26%), but this group did not impact strongly on N. lugens or Nephotettix spp.
lncidence of parasitism by pipunculids on Nephotettix virescens (Distant) (Homoptera: Cicadel-
lidae) and Nephotettix nigropictus (Stal) (Homoptera: Cicadellidae) was higher than for any of the
parasitoid species (Pena and Shepard 1986) and pipunculids may be an important source of
irreplaceable mortality in green leafhoppers. During the 1984 dry season (January-April),
parasitism reached more than 50% on N. nigropictus.
33.2 Predators
The most abundant predator of hoppers is the mirid, C. lividipennis. When offered eggs of brown
planthopper, whitebacked planthopper and green leafhopper, C. lividipennis exhibited a preference
for brown planthopper eggs (Figure 5). We have found that increasing the ratio of brown planthop-
per to C. lividipennis in cage experiments resulted in a population increase of the hopper (Manti
and Shepard, unpubl.) (Figure 6). Based on the results of laboratory studies and field observations
of a strong numerical response of the mirid to brown planthopper populations, we disagree with
Kenmore (1980) that C. lividipennis is an opportunistic predator that exploits outbreaks of hoppers
only after other control factors have failed. Clearly, C. lividipennis must rate as one of the most
important predators of hoppers in the rice ecosystem. Observations by Stapley (1975, 1976) in the
Solomon Islands support this conclusion Although primarily an egg predator, C. lividipennis also
feeds on young hopper nymphs.
Fig. 5. Feeding preference by Cyrtorhinus lividipennis Reuter on eggs of green leafhopper (GLH).
whitebacked planthopper (WBPH) and brown planthopper (BPH).
Fig. 6. Results of an experiment to determine the effect of different ratios of the brown planthopper
to Cyrtorhinus lividipennis Reuter. The resulting populations at the end of 60 days are shown for each
treatment. (A) 2 pair BPH O pair C. lividipennis; (B) 1 pair BPH: 1 pair C. lividipennis; (C) 2 pair
BPH 1 pair C. lividipennis; (D) 3 pairs BPH: 1 pair C. lividipennis.
The wolf spider, L. pseudoannulata is also recognized as a major regulator of brown planthop-
per populations, and probably is its most important predator. Kenmore et al., (1984) reported a
strong correlation between L. pseudoannulata density and peak populations of brown planthopper.
Studies using arenas in the laboratory reveal that a single wolf spider can easily consume between 7
to 45 hoppers per day (IRRI 1975,1985).
In addition, Nakasuji and Dyck (1984) suggested that M. d. atrolineata is an important brown
planthopper mortality factor. In one of their field studies, 67% of the Yrown planthopper adults fell
Fig. 7. Micraspis crocea Mulsant feeding on a brown planthopper.

OOI AND SHEPARD 599
from the rice plant. If this phenomenon is common, the hoppers would be subjected to high levels
of predation by water bugs. Other surface dwelling bugs such as Mesovelia sp. and L. fossarum feed
on both whitebacked planthoppers and brown planthoppers or almost any small, soft-bodied insect
that falls into the paddy water. The complex of predators that attack the whitebacked planthopper
and green leafhopper, Nephotettix spp., is similar to that of brown planthopper. The most important
are various species of spiders which occupy the different strata of the rice canopy. Tetragnathids
and oxyopids, which inhabit mostly the upper canopy, attack Nephotettix spp. Beetles, such as
Micraspis sp. (Figure 7), Ophionea nigrofasciata (Schmidt-Goebel) (Coleoptera: Carabidae) and P.
fuscipes also feed on green leafhoppers, but probably do not cause major mortality.
4. MAXIMIZING THE IMPACT OF PREDATORS AND PARASITOIDS

Natural enemies are always present in the rice field, working to keep pests in check. However, little
effort has been spent to maximize their impact. We recommend the following procedures to realize
the full benefit of the action of predators and parasitoids.
4.1 SURVEYS
Maintaining an inventory of natural enemies is the first step towards recognizing their existence.
Many countries already possess some information about indigenous predators and parasitoids. An
example on parasitoids of rice pests was reported by Pagden (1934). This subsequently developed
into a larger volume compiled by Ahmad and Ho (1980).
Such an inventory must be constantly revised and updated in order to correct incomplete or
sometimes inaccurate records. Often, predatory insects are mistaken for pests. For example, the
cricket Anaxipha longipennis (Serville) (Orthoptera: Gryllidae), is predatory but many members of
the family are plant feeders (Canapi et al., 1988). Few rice extension workers would recognize it as
a beneficial species.
When an inventory of beneficial species is not available, a system for collecting this information
should be initiated immediately. Reference to entomological literature would be the first step to
determine available information from within a country, region or climatically similar areas, This
should be complemented with extensive field collections. Rice field arthropods can be collected
using a variety of techniques, including sweep nets, suction machines or collecting live arthropods
individually with vials.
Individual collection is important for identifying suspected predators or rearing parasitoids to
the adult stage. Suspected predators should be tested in laboratory cages or arenas by offering them
different hosts. Samples of live insect pests can be reared in individual containers to provide
accurate records of parasitism. When insects are reared together, records may be inaccurate.
Arthropods should be catalogued and the asistance of well known insect identification services
consulted to identify them. Many of the rice insects can be identified by the Entomology Depart-
ment of the International Rice Research Institute. However, for many predators and parasitoids,
the assistance of established taxonomic services would be most helpful (e. g. the CAB International
Institute of Entomology).
When possible, outbreaks should be investigated to determine their causes. Often, outbreaks
of rice insect pests in tropical Asia result from adverse farm practices, such as unnecessary and
intensive insecticide use. These records are important in providing directions for research into IPM
implementation.
4.2 SAMPLING AND SURVEILLANCE

After gathering information on the composition of beneficial species, the seasonal and relative
abundance of rice pests and their natural enemies should be determined. Sampling for the
abundance of rice pests alone can result in a change in agricultural practices that adversely affect
natural biological control.
For example, rice stem borers were considered the most important pests of rice in Malaysia
since 1887. When two attempts at biological control failed, emphasis was placed on using insec-
ticides for control (Wyatt 1956, 1957; Koyama 1964; Kimura 1965). Those studies resulted in the
recommendation by the Department of Agriculture to treat rice plants prophylactically for stem
borer control. Lim (1972) recognized the integration of chemical and biological control and
recommended the use of granular insecticides instead of liquid formulations. However, surveys by
Ooi (1974,1976) showed low incidence of rice stem borers in the Muda Scheme and the Tanjung
Karang Irrigation Scheme. This low incidence was subsequently confirmed by Khoo and Lee (1978)
and Zam and Ooi (1978). With percent bored tillers being generally less than 5% nationwide, there
was no justification for preventive applications of insecticides for rice stem borer control.
Similar trends were observed in Bangladesh, where a crop loss assessment study by Catling et
al. (1978) confirmed that the stem borer is under natural control. Evidence that egg parasitism and
predation are important mortality factors is given in Catling (1979), Catling et al., (1983), Hikim
(1988), Khoo and Lee (1978), and Shepard and Arida (1986).
These examples show the importance of field surveys which, when properly conducted, can
provide strong circumstantial evidence of the impact of natural enemies. At the same time, the
overall importance of the target pest can be assessed.
Surveys of predator populations are useful in establishing their relative importance. A study of
the abundance of the brown planthopper and its predator, C. lividipennis shows a strong correlation
between the populations of pest and predator in the rice field (Figure 8). While the figure does not
show direct causal effect of the action of the predator on the pest population, it indicates a strong
numerical response between predator and prey numbers in the field even at low pest densities.
Not all predators in the rice field respond to prey populations like C. lividipennis. Indeed,
Figure 9 shows that, there is little relationship between the spider predator (Lycosa sp.) and prey
numbers. However, this important predator of brown planthopper removes large numbers of this
pest in the field and serves as the first line of defence against a pest buildup.
OOI AND SHEPARD 601
Fig. 8. Population trends of the brown planthopper (BPH) and Cyrtorhinus lividipennis in an
insecticide free field in Sungai Burong, Malaysia (Season 2,1983). Data were collected using sticky
boards. There is a significant correlation between Cyrtorhinus and BPH populations.
Fig. 9. Population trends of the brown planthopper (BPH) and Lycosa spp. in an insecticide free field
in Sungai Burong, Malaysia (Season 2, 1983). Data were collected using a sticky board. While there
is no correlation between populations of the spider and BPH, the spider's presence in the field
throughout the season will limit the populations of the hoppers and other insect pests.
Field studies of both pests and predators may lead to the development of practical pest
surveillance and forecasting techniques. A system developed in Malaysia has been described by
Ooi (1982a). This surveillance system requires that the field scout counts and records planthop-
pers, number of damaged tillers and leaves and predators. Although many predators of the brown
planthopper have been recorded (Otake 1977, Chiu 1979, Kenmore et al., 1984), only those
considered to be important in the regulation of the brown planthopper in Malaysia (Lim et al., 1978)
are counted. These predators are C. lividipennis, Lycosa spp. and other spiders, P. fuscipes,
Cusnoides interstitialis (Schmidt-Goebel) (Coleoptera: Carabidae) and S. octomaculata. After an
examination of the data collected by the surveillance system, Ooi (1982b) concluded that the brown
planthopper population is checked by predators as long as the ratio of brown planthopper to the
combined predators does not exceed 50 : 1. Even if the population of the brown planthopper passes
the economic threshold level, the recommendation for chemical intervention takes into considera-
tion the prey:predator ratio.
Incorporting predators into decision making has helped maximize the impact of predators in the
rice field. Farmers are encouraged to use insecticides only when the ratio is in favor of the pest. Such
an approach has been adopted by some rice farmers in Malaysia. Absolute counts are not necessary.
Relative numbers of predators to pest numbers indicate quickly whether natural control is working.
Another way to maximize the action of predators is provided by Shepard et al., (1986) who
recommended the use of a sequential sampling program for hopper pests. This program is
particularly useful in saving the time needed to reach a decision about whether or not to make an
insecticide application. Further refinement of the model was made by incorporating predators
(Shepard et al., 1988). In the model, numbers of hoppers are adjusted according to numbers of
predators. For each predator counted, five hoppers are subtracted from the total number of large
nymphs and adult hoppers. The adjusted number of hoppers is then used in deciding whether or
not to treat a field. This improved sequential program is useful in preventing unnecessary insec-
ticide treatment.
4.3 RESEARCH
Successful IPM programs in the tropics have developed from observations that natural enemies play
an important role and their conservation through reduced insecticide usage results in cost effective
pest management (Conway 1971, Wood 1964). Way (1973) described this empirical approach in
recognizing the importance of existing natural enemies as an important prerequisite to successful
pest management. However, qualitative observations must lead to quantification of the action of
natural enemies to ensure that it continues to be an important paradigm in tropical rice IPM
programs. Some research carried out to quantify the impact of predators. and parasitoids has been
reviewed here to provide some directions on how such studies can benefit IPM programs.
In the past, the study of parasitoids of rice pests was emphasized more than that of general
predators. In the case of rice stem borers, such emphasis was justifiable, because egg parasitoids
are very important in checking the borer population. However, general predators are also impor-
tant. For example, Pantua and Litsinger (1984) reported that in a study to determine levels of
parasitism of the yellow stem borer, the experiment was disrupted by extensive egg predation
attributed to C. longipennis. Rothschild (1970), Sands (1977) and Manley (1985) reported that C.
longipennis is an important predator of rice insect pests. However, Grist and Lever (1969) and van
Vreden and Abdul Latif (1986) reported that C. longipennis also damages rice plants. Research is
needed to determine the role of this longhorned grasshopper as a predator and as a pest.
OOI AND SHEPARD 603
Besides the study of the biology of predators in the laboratory and the types of prey they feed
on, research into prey preference and consumption rates are important to indicate their relative
importance. General predators have been found to exhibit preferential feeding. This has been
shown with M. virraticollis (Sec 3.1.2), rice field lady beetles (Figure 3) and C. lividipennis (Figure
5). Rate of feeding is another important aspect of predation that can be evaluated in the laboratory.
Lim (1970) pointed out the possible importance of weeds along field levees that serve as alternate
hosts for stem borers. These weeds can sustain a reservoir of natural enemies during the crop and from
one crop to the next. Thus, manipulation of weed communities may offer possibilities for increasing
populations of stem borer parasitoids and predators. Studies by Rubia and Shepard (1987) showed that
M. vittaticollis oviposit in four species of weed in addition to rice plants. In a free choice experiment, it
was found that the different weeds were as attractive as rice plants for oviposition.
One method for laboratory evaluation of predators and parasitoids is to study the functional
response of the natural enemies to different population densities of prey. Nakasuji and Dyck (1984)
confirmed that M. d. atrolineata is one of the more important natural enemies of brown planthopper
on the basis of a study of its functional response and the high populations of the veliid in rice fields.
Field experiments are a necessary extension of the laboratory studies. A comparison of sprayed
and unsprayed fields indicates the population trends of both pests and natural enemes. Ooi (1986)
showed that mirid predator and spider populations are adversely affected, resulting in an increase
in the brown planthopper population. In an adjacent field, where only one early spray was applied,
the brown planthopper was kept in check by the same predators (Figure 10). Kenmore et al. (1984)
demonstrated that pest outbreaks can be cawed by insecticide sprays that kill predators.
Fig. 10. Comparison of two adjacent fields in Sekinchan, Malaysia to show the difference of a field
that received a single spray early in the season (left), with a field that received at least three sprays
(right). Insecticide sprays killed predators and allowed the brown planthopper (BPH) to develop a
high population.
604 PREDATORS A N D PARASITOIDS OF RICE INSECT PESTS
Using cages to exclude predators is another effective method to demonstrate the importance
of predators in regulating pest populations (Kenmore et al., 1984). The results of an exclusion cage
study in Tanjung karang, Malaysia is shown in (Figure 11). One gravid female per hill was
introduced into a field cage enclosing 9 hills. In one cage, predators were excluded while in the
second, the sides of the cage were opened 24 hours after introduction. The resulting large
population of brown planthopper in the absence of predators mimics an outbreak situation. In
contrast the population in the cage with the predator shows the importance of predators in keeping
brown planthopper populations down.
Fig. 11. Exclusion cage study conducted at Tanjung Karang, Malaysia. (A) One gravid female brown
planthopper (BPH)/hill was allowed to develop within a cage free of predators. After 45 days, the
population increased to513 BPH/hill. (B) A similar experiment but the sides of the cage were opened
24 hours after introduction of the gravid females. After 45 days, the population was 10 BPH/hill.
BPH= adults + numphs; Predators= C. lividipennis + spider.
To demonstrate the action of predators and parasitoids on the eggs of rice pests, van den Berg
et al., (1988) exposed eggs of Rivula atimeta (Swinhoe) (Lepidoptera: Noctuidae), Naranga aenes-
cens Moore (Lepidoptera: Noctuidae) and Hydrellia philippina (Ferino) (Diptera: Ephydridae) on
rice seedlings in the field. Egg predation in all three insect pest species increased as the crop
matured (Figure 12). Their study shows that predators should be conserved because of their
importance in controlling these early season pests.
We have discussed ways that entomological research in rice can contribute to maximizing the
impact of predators and parasitoids. The examples were selected to illustrate the possible outcome of
such studies. Transferring this knowledge from research to the farmers through extension programs is the
next important step. It is our firm belief that, as we unravel the complexities of natural biological control in
rice fields, the beneficiary will be the rice farmers. These farmers have lived with the predators and
parasitoids for many years and it is appropriate that they learn about them and work to conserve them.
OOI AND SHEPARD 605
Fig. 12. Per cent predation on Hydrellia philippina, Naranga aenescens and Rivula atimeta in a
farmer’s field, Victoria, Laguna, Philippines, 1987. (after van den Berg et al, 1988)
4.4 DEMONSTRATION
After going through the procedure of surveying for natural enemies, estimating their impact
through sampling and surveillance and confirming this through research, the next step is to
demonstrate the importance of natural enemies to the rice farmers. While we recognize that
farmers are the eventual recipient of the information about natural enemies, we also appreciate that
support from policy makers, agricultural administrators and technicians is essential.
A first step toward explaining the possibilities of natural control is the publication of ap-
propriate materials. A field guide developed by Shepard et al. (1987) illustrates that the natural
enemies of rice pests are actually friends of the rice farmers. Another useful publication is the book
by van Vreden and Abdul Latif (1986) about rice pests and their natural enemies. The next step is
to train the agricultural technicians how to demonstrate the action of predators and parasitoids to
farmers. A manual to do this has been prepared by Ooi, Shepard and Kenmore (in preparation).
This manual suggests that the extension worker and farmers work together in the rice fields.
The first step is to visit rice fields with farmers to show them the important “friendly insects”,
particularly predators. As parasitoids are generally small, the manual suggests using predators to
bring home the message. Predators, such as L. pseudoannulata are collected, starved for a day or
so and fed pests in the presence of rice farmers. This step is supplemented by field cage experiments
where predators are excluded and farmers are shown what would happen in the absence of predators.
More sophisticated procedures, such as insecticide disruption of natural biological control are
described to demonstrate that insecticides can be a disadvantage if used indiscriminately. Placing
known numbers of prey insects in the field and returning later to determine predation and
parasitism is also a useful techinque. If agricultural technicians foIlow the procedures outlined,
both the technicians and farmers should feel as secure about natural biological control as they now
do about using chemical insecticides.
5. SUSTAINING THE NATURAL ENEMY CORE OF IPM

Integrated Pest Management (IPM) has its origin in the integration of biological and chemical
conrol (Stern et al., 1959). Central to this is the concept that natural biological control is the core
of IPM. Hence, good pest management should start with a knowledge about the natural enemies of
the pest and their role in controlling pest numbers, and how other control measures can be
integrated with this core.
For rice, it is impossible to develop an IPM program without understanding the role of
predators and parasitoids. These "friends of the rice farmer" (Shepard et al., 1987) were associated
with rice pests long before rice was cultivated in its present form. From our review of these natural
enemies, we conclude that predators and parasitoids in the rice fields of tropical south and
southeast Asia are essential for maintaining insect pest populations at low levels.
However, methods of rice cultivation are undergoing great biotic and abiotic changes. Some of
these changes may adversely affect the indigenous biological control communities. IPM prac-
titioners should recognize that changes in rice cultivation are inevitable. They should involve
themselves with these changes to ensure that, although other crop production practices may change,
predators and parasitoids are not adversely affected. We suggest some steps to sustain the action
of natural enemies.
Ongoing research activities enable integration of natural enemy research into other areas of
activities such as breeding for pest resistance. Studies by Kartohardjono and Heinrichs (1984) and
Joshi (1988) have indicated that action of natural enemies combined with resistant rice varieties
resulted in lower pest populations, thereby slowing down the development of adaptations to these
varieties. Joshi (1988) showed that when insecticides were sprayed onto fields of resistant and
susceptible rice varieties, hopper population resurgence was less in the resistant than in the
susceptible variety (Figure 13). Clearly, resistance in the rice variety was masking the impact of
Fig. 13 Mean numb of macropterous brown planthopper (BPH) adults sampled by D-Vac in the
wet season, 1986, Koronadal, Mindanao, Philippines to demonstrate the effects of insecticide spray
on the BPH population in resistant (IR62) and susceptible (IR 1917) rice varieties (Joshi 1988).
OOI AND SHEPARD 607
natural enemies. The importance of this interaction between natural enemies and resistant varieties
is slowly being recognized.
Another related aspect is the potential for developing new plant types which possess mor-
phological characters that affect the efficacy of some natural enemies. Changes in plant morphol-
ogy (e.g. leaf pubescence) have been reported to have negative effects on some parasitoids in
tobacco (Rabb and Bradley 1968, Elsey and Chaplin 1978) and in soybean (Turner 1983). Similarly,
there could be changes in plant chemistry that could affect natural enemies.
Mechanical drilling and broadcasting of seed and mechanical transplanting are replacing
traditional transplanting in some areas. The implications of these changes on the efficacy of
predators and parasitoids should be determined. In the study of parasitism and predation of stem
borer eggs, Shepard and Arida (1986) suggested that the higher incidence of egg parasitism in
transplanted rice was due to a more open canopy.
A cultural practice that may affect the efficacy of predators and parasitoids involves planting
synchrony. The decline in incidence of rice stem borer in Malaysia following double cropping was
attributed to the survival of natural enemies provided for by the asynchrony that developed
(Balasubramanian and Ooi 1977). However, Loevinsohn (1984) suggested that asynchrony is
responsible for increased incidence of rice pests. Oka (1979) reported synchronous planting was
critical to the management of the brown planthopper. We believe that there is insufficient evidence
to prove that either synchrony or asynchrony are important factors affecting the efficacy or natural
enemies. More studies are required in this field.
The importance of weeds on rice field levees as reservoirs of natural enemies should be further
investigated (Sec 4.3). The possibility of using or manipulating alternate plant hosts or even
alternate arthropod hosts of important natural enemies to provide effective biological control early
in the season is another area for future research. Such an approach may overcome the problem of
late stem borer egg parasitism reported by Catling (1979) and Catling et al., (1983). Surely, we are
only on the threshold of understanding the impact of natural enemies in a dynamic rice ecosystem.
Further understanding of natural enemies may occur through modelling. This approach should
include a description of the ‘problematique’ in terms of its biological, technical, social and economic
dimensions (Norton and Heong 1988). Such a holistic approach to problem solving will encourage
development of a pest control concept such as ‘inaction’ levels (Sterling 1984) which will feature
strongly in rice integrated pest management with a core founded on natural biological control.
6. SUMMARY
A review of past attempts at biological control of rice insect pests provides evidence that importa-
tion of exotic parasitoids is effective when the pests we also exotic. Failures were recorded with
inundative releases and importation of exotic parasitoids against indigenous insect pests. casual
introduction of natural enemies is an antithesis to good biological control. Rice insect pests in the
tropical south and Southeast Asia have a long and close association with their natural enemies.
Predators and parasitoids of these pests are most important in keeping them under biological
control.
Conserving the natural enemies is the strategy recommended. In the past, emphasis has been
on the qualitative studies of parasitoids of rice pests. Recent quantitative studies have confirmed
the importance of predators and parasitoids of rice stem borers, leaf- and planthoppers and
leaffolders. In recognition of their importance, a procedure is recommended to maximize the
impact of these beneficial arthropods. The steps are (1) surveys for identification, (2) sampling and
surveillance, (3) research to assess predator and parasitoid impact, and (4) demonstration.
Natural biological control as the center of integrated pest management is the paradigm
maintained in this review. Hence studies of the action of predators and parasitoids should be part
of an ongoing research program. To ensure sustained efforts in maintaining this core of IPM, it is
advised that research into natural enemies should take cognizance of changing plant types and a
dynamic rice ecosystem.
ACKNOWLEDGEMENTS
This chapter was completed when the first author was a Visiting Scientist at IRRI. We are
grateful to our respective institutions for their support in this endeavor. Our thanks to friends and
colleagues who have willingly shared their views with us. We thank H. D. Catling and L. Pollard for
their technical and editorial comments, respectively.
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8
Pathogens of Rice Insects 1
MICHIEL C. ROMBACH 2 AND D. W. ROBERTS

Boyce Thompson Institute for Plant Research at Cornell University
Ithaca, NY
REMEDIOS M. AGUDA
1. INTRODUCTION
SEVERAL INSECT GROUPS OCCASIONALLY cause damage to rice in Asia. These include the stem
borers, leaffolders, stink bugs, and a few plant- and leafhopper species. However, in undisturbed
rice fields most pests are effectively suppressed by indigenous populations of natural enemies such
as predators, parasites and entomopathogens. Conservation of these natural enemies is the key to
stable rice pest control in southeast Asia. Most pest problems occur when agricultural practices
such as excessive pesticide use destroy these populations of natural enemies. The disruptive
influence of broad spectrum chemical insecticides on natural enemies is well documented (Ken-
more et al., 1984). Several selective chemical insecticides are available, but these chemicals are often
1. Disclaimer notice: In this article only research is reviewed. Mention of any proprietary product does not constitute an
endorsement or a recommendation for its use by the Boyce Thompson Institute for Plant Research at Cornell or by the
International Rice Research Institute.
2. Insect Pathology Resource Center, Boyce Thompson Institute for Plant Research at Cornell, Tower Road, Ithaca, New
York 14853, U.S.A. Seconded to Department of Entomology, International Rice Research Institute, Los Baños,
Laguna, Philippines. Postal address: IRRI, P.O. Box 933, Manila, Philippines.
614 PATHOGENS OF RICE INSECTS
too expensive for use by the small scale rice farmers in the developing Asian countries where rice is
mostly a subsistence crop and profit margins are small.
Insect pathogens may offer an alternative to selective chemical insecticides. Microbial insec-
ticides are already widely used in agricultural systems (Burges 1981, Fuxa and Tanada 1987), and
knowledge of the biology and ecology of the diseases has greatly increased in recent years (Samson
et al., 1986). Microbial control agents are more easily manipulated for field use than are predators
and parasites and their mass production is comparatively easy. Many entomopathogens, notably the
insect bacteria and fungi, can be produced in industrial type fermentors in inexpensive liquid media.
The products are formulated, packaged and can be stored for several months. These features make
production of some of the insect microbials at the community level a viable approach. Insect
pathogens are safe to humans and other non-target species so they can be used harmoniously with
other control tactics. Application of these materials can usually be made with conventional spraying
equipment.
New biotechnology techniques can be used to induce changes in insect microbes that increase
their efficacy (Kirschbaum 1985). This biotechnological approach can be carried out on a wide
range of insect pathogens, with bacterial and viral pathogens being prime candidates for genetic
manipulation.
The incorporation of the genetic material from the insect pathogens into the plant may also
offer promise (Meeusen and Warren 1989, Vaeck et al., 1986). For example, incorporation of
Bacillus thuringiensis Berliner genes into plants induces production of the Bt endotoxins by the plant
cells. The toxins affect foliage feeding caterpillars as well as stem borer larvae. This may become an
effective and economical way to reduce the necessity for chemical insecticides.
The literature is replete with anecdotal accounts of infections of rice insects by en-
tomopathogens and, more rarely, reports on epizootics of viruses and fungi. However, few studies
report on the manipulation of insect pathogens for rice pest control.
There is no doubt that naturally occurring microbial agents are important in the regulation of
rice insect populations. However, the environmental conditions triggering natural epizootics are ill
defined. In rice the natural outbreaks of entomopathogens are positively correlated to pest density.
Usually this density is above the damage threshold and naturally occurring pathogens do not
prevent crop damage. Manipulation (i.e. mass production and application) of some of the
microbials should be attempted to suppress pest populations below threshold levels.
This chapter reviews some of the most pertinent literature on pathogens of rice insects. More
references are provided in Rombach et al., (1987c). Identification and practical application of
several groups of rice insect pathogens are discussed. Mycological terminology used in this chapter
is defined in Table 1.
ROMBACH ET AL. 615
TABLE 1
Explanation of mycological terms used
Basipetal chain, chain of conidia with the youngest conidium at its base (e.g., as in Metarhizium, Paecilomyces, and
Nomuraea).
Blatospore, a spore (conidium) which is differentiated from a hypha by budding.
Byssoid, cotton-like, made up of delicate threads.
Cerebriform, brain-like or convoluted.
Conidium, asexsual and nonmotile spore of a deuteromycetous fungus.
Conidiogenesis, process of conidium formation.
Conidiogenous cell, fertile cell from which or inside which the conidia are produced.
Conidiophora, simple or branched hypha with conidiogenous cells which produce the conidia.
Epizootic, epidemic of insect disease.
Fungi Imperfecti, anamorphs of higher (sexually reproducing) fungi in the classes Blastomycetes (yeasts), Hyphomycetes
(molds), and Coelomycetes (pycnidial fungi).
Hypha fungus filament.
Hyphal body, short hyphal fragment.
Hyphomycetes, class of Fungi Imperfecti, molds. Mycelia can be sterile or bear conidia on separate hyphae which can be
agregated (e.g. in a synnema), but not in distinct conidiomata.
Marcescens process, production process for filamentous fungi in which mycelium mass is produced by submerged fermen-
tation, filtrated, dried, milled, and formulated. The mycelium particles are applied in the field and sporulate on the
plants.
Mycelium, mass of hyphae.
Mycoparasite, fungus parasitic on another fungus (as in Calcarisporium ovalisporum).
Phialide, a conidiogenous cell which produces basipetal chains of conidia from one or more open ends.
Polyphialide, phialide with more than 1 neck.
Rachis, the zig-zag extension on the conidiogenous cell (as in Beauveria bassiana) resulting from sympodial conidiogenous
cell development.
Ramificated, branched.
Resting spore, a spore which germinates after a resting period, an “overwintering” spore (e.g. large thick walled spores in the
Entomophthorales).
Stroma, compact mass of hyphae.
Sympodial proliferation, a mechanism of conidium production in which each new conidium is formed just behind the
previous conidium, resulting in elongation of the cell (e.g., as in Beauveria bassiana).
Synnema, aggregation of erect hyphae, conidiophores, and conidiogenous cells. This structure can be sturdy (e.g., as in
Hirsutella citriformis) or loose (e.g., as in H. strigosa).
Verticillate, having parts in rings (verticils), whorled.
Zygospore, large, thick walled spore produced by conjugation of hyphae (as in the body cavity of insects infected by
entomophthoralean fungi).
2. ENTOMOGENOUS FUNGI
Various entomogenous fungi have been collected from rice arthropods including planthoppers and
leafhoppers (Homoptera), caterpillars (Lepidoptera), stink bugs (Hemiptera), and spiders (Arach-
nida). Some of these fungi can cause devastating epizootics, but other fungi (ie. most spider
Fig. 1. Metarhizium: A. M. anisopliae var. majus Johnston, from coconut beetles; B. M. anisopliae (Metsch.)
Sorokin var. anisopliae, from the brown planthopper Nilaparvata lugens (Stal); C. M. album Petch from the
rice white leafhopper, Cofana spectra (Distant); D. M. flavoviride var. flavoviride Gams & Roszypal, from
soil; E. M. flavoviride var. minus Rombach, Humber & Roberts, from the brown planthopper; and F in
culture.
ROMBACH ET AL.. 617
pathogens) are mostly collected on individual spiders, and epizootics have not been observed. The
reason for the different epizootic potential of the fungi is not known, and offers an interesting area
for ecological research.
Virtually all fungi of agricultural insect pests are Hyphomycetes or Entomophthorales (Roberts
and Humber 1981), although entomogenous fungi occur in nearly all fungal orders. Collections of
rice insects infected with the fungi and the isolation, mass production, and application of the fungi
Fig. 2. A. Metarhizium anisopliae (Metsch.) Sorokin, palisade layer of phialides and conidia; B. Nomuraea releyi
(Farlow) Samson infecting a rice looper; C. Erynia delphacis (Hori) Humber on the green leafhopper
Nephotettix virescens Distant; D. Conidiobolus coronatus (Cost.) Batko on the green leafhopper, Nephotettix
virescens (Distant). Note the halo of conidia on the lower part of the rice leaf.
will be discussed. A practical guide on handling fungi is given by Aguda and Rombach (1986) and
a review on application and mass production by Hall and Papierok (1982). Fungal pathogens will be
discussed by host group.
2.1 ENTOMOGENOUS HYPHOMYCETES

2.1.1 Hyphomycetes on Planthoppers and Leafhoppers
Few species of entomogenous Hyphomycetes are found in agricultural crops as compared to
natural tropical habitats (Evans 1974, 1982). Fungi most commonly collected in agricultural crops
include species of Metarhizium, Beauveria, Hirsutella, Nomuraea, and Paecilomyces (Samson 1981).
Most of the pathogens infecting planthoppers and leafhoppers belong to these fungus genera
(Soper 1985).
The planthoppers Nilaparvata lugens (Stal), leafhoppers Nephotettix virescens (Distant), and
Recilia dorsalis (Motschulsky), rice stink bug, Leptocorisa oratorius (F.), and black bugs
(Scotinophara spp.) feed by inserting their mouthparts into the plant. These insects are free of
pathogens which require oral uptake, such as insect protozoans, viruses, and bacteria.
Common pathogens of rice planthoppers and leafhoppers are Metarhizium anisopliae
(Metsch.) Sorok., M. flavoviride Gams & Rozsypal var. minus Rombach, Humber & Roberts,
Metarhizium album Petch, Beauveria bassiana (Bals.) Vuill., and Hirsutella citrifomis Speare.
Saprophytic species of Penicillium,Aspergillus and Fusarium were also reported from these insects.
However, these fungi mostly invade dying or dead insects, and are of no significance in biological
control.
Metarhizium Species.- M. anisopliae, the “green muscardine fungus”, is a common insect
pathogen. Veen (1968) compiled a list of more than 200 insects infected with M. anisopliae. Ferron
(1978, 1981) discussed the ecology and practical use of M. anisopliae. The fungus is used in Brazil
for control of cercopid spittlebugs such as Mahanarva postica (Wlk.) on a large scale.
Tulloch (1976) reports on M. anisopliae var. anisopliae Tulloch (Figs. 1B, 2A, 3A, D) and M.
anisopliae var. majus Johnston (Fig. 1A). These varieties differ in morphology and host range.
Brady (1979d) provides detailed descriptions of these varieties.
M. anisopliae var. anisopliae commonly infects planthoppers and leafhoppers, rice black bugs
(Scotinophara sp.) (Fig. 3D), and various other rice insects. It can decimate cultures of rice plant-
and leaf-hoppers (Aguda et al., 1984a, Medrano et al., 1984a). It has powdery masses of dark green
to yellow-green columns of conidia which arise from white mycelium. Cylindrical phialides, 6-13 ×
2-4 µm are located on stout conidiophores. The conidia are cylindrical to broadly ellipsoid with
rounded ends, 4.5-8.5 × 2.5-4,um, and are produced in long dry parallel chains. The fungus looks
the same in pure culture as on the insect.
M. flavoviride was collected from curculionid larvae and was isolated from soil (Gams and
Rozsypal 1973) (Fig. 1D). Recently M.flavoviride var. minus was described from planthoppers and
leafhoppers on rice in the Philippines and Solomon Islands (Rombach et al., 1987b) (Figs. 1E, 4A).
It produces greyish green conidial masses. Phialides, 9-15 × 3-4 µm, are club shaped and placed on
ROMBACH ET AL. 619
Fig 3. A. Metarhizium anisopliae (Metsch) Sorokin in culture. Note difference between the mycelium growth
(white) and sporulation (dark green); B. Paecilomyces farinosus (Holm ex S. F. Gray) Brown and Smith,
synnemata formation in culture; C. Hirsutella cidiformis Speare, synnemata formation in culture; D.
Metarhizium anisopliae (Metsch.) Sorokin infecting the Malayan black bug, Scotinophara coactata (F.) E.
Beauveria bassiana (Bals.) Vuill. on the rice bug, Leptocorisa oratorius (E); F. Paecilomyces tenuipes (Peck)
on a lepidopteran pupa. Note the synnemata lifting fertile area above the soil surface.
conidiophores on a white mycelium mat. Conidia, 4.5-7.5 × 2-3µm, are ellipsoid. Synnemata are
produced in culture.
Fig. 4. A. Metarhizium flavoviride var. minus Rombach, Humber & Roberts infecting the brown planthopper,
Nilaparvata lugens (Stal); B. M. flavoviride var. minus in culture; C. Gibellula leiopus (Vuill.) Mains on a
spider; D. Mythimna separata (Walker) infected by a nuclear polyhedrosis virus (NPV); E. Gibellula pulchra
(Sacc.) Cavara on a spider; F. Metarhizium album Petch on Cofana spectra (Distant).
M. album infects the cicadellid Cofana spectra (Distant) (Petch 1931) (Figs. 1C, 4F) and the
green leafhopper in the Philippines and in Indonesia. It differs from M. anisopliae in its white color,
ROMBACH ET AL. 621
cerebriform structure of the stroma, and in size of the conidia. M. brunneum (Petch 1935) was found
on a cicadellid insect in the Philippines. However, recent collections on the green leafhopper and
C. spectra show that M. album and M. brunneum are conspecific (Rombach et al., 1987a) and that
M. album is the correct name for this species. It produces bulging masses of hyphal bodies covered
by a layer of white mycelium. Up to 80 µm long conidiophores have verticillate ramifications near
the top (Fig. 1). Each branch bears 2-5 cylindrical to club shaped phialides, 10-12.5 × 2-3.5 µm.
Conidia, 4-6 × 1.5-2.5 µm, are broadly to narrow ellipsoidal and whitish brown. Cylindrical conidia
of 8-11 µm length are found in culture.
The Russian scientist Eli Metschnikoff (1845-1916) pioneered mass production and field
application of M. anisopliae. Metarhizium species can be easily isolated and grown in culture (Fig.
3A). On Sabouraud dextrose agar (SDA) mycelium grows abundantly but conidiation is sparse. On
less nutritive media such as Emerson's YpSs (YpSs), oatmeal agar (OA), and malt extract agar
(MEA) mycelium growth is flat and conidiation abundant. YpSs and MEA are the standard media
for Metarhizium cultivation.
Solid media are commonly used for Metarhizium mass production (Soper and Ward 1981).
Plastic bags or flasks with rice or bran are inoculated, and the conidia harvested after a few weeks.
This is the common mass production procedure in Brazil and China (Ferron 1981) where the fungus
is used to control spittlebugs in pasture land and sugarcane leafhoppers. Marques et al., (1981)
provides detailed information on small production plants and production procedures.
M. anisopliae also grows in liquid culture where mycelium pellets as well as hyphal bodies
(blastospores) are produced. This material can be processed with the marcescent process. This
method is develped for production of Entomophthora-like fungi (McCabe and Soper 1985), but it
can also be used for Hyphomycetes (Rombach et al., 1986c) (Fig. 5). The fermentor slurry is
filtrated and the mycelium dried and milled. The particles can be stored for several months at low
temperatures. When applied in the field the particles adhere to the plant where they produce
conidia after a few days. These conidia infect the insects, Rombach et al., (1986a) demonstrated the
effect of M. anisopliae marcescent particles in brown planthopper control. The method has not yet
been tested on a large scale.
The ecology of Metarhizium on rice insects is not well known. M. flavoviride var. minus is
restricted to planthoppers and leafhoppers, while M. anisopliae var. anisopliae has a wide host
range. Ferron et al., (1972) show that different M. anisopliae strains infect different hosts. There-
fore, infectivity tests with different strains have to be carried out before large scale field experiments
are started.
Metarhizium spp. produce dry conidia which are easily dislodged by air movements. Wind is
therefore important for dispersion. M. flavoviride and M. anisopliae are commonly isolated from soil
(Domsch and Gams 1970, Domsch et al., 1980) where conidia can survive. Soil is probably the
source of inoculate for natural epizootics.
Few experiments have been conducted using M. anisopliae for control of rice insects. However,
it has been shown that epizootics of M. anisopliae and M. flavoviride can be initiated in brown
Fig. 6. A. Schematic representation of the production and application of dry mycelium of entomogenous Hyphomycetes; B. Infection of the brown
planthopper in the field by different entomogenous fungi at 7, 14, and 21 days after application, Mf = Metarhizium flavoviride var. minus Rombach,
Humber & Roberts, conidia; Ma = M. anisopliae (Metsch.) Sorokin, conidia; Hc = Hirsutella citrifomis Speare, conidia; Bb and Bb 252 =
Beauveria bassiana (Bals.) Vuill., conidia; Plm = Paecilomyces lilacinus (Thom) Samson, mycelium; MaM = M. anisopliae, mycelium; Control -
untreated; C.B. bassiana mycelium production in different liquid media (CS = saccharose-yeast extract medium at different concentrations, CM =
maltose-yeast extract medium at different concentrations, YS = yeast extract medium (1%) with increasing concentrations of saccharose, and
YM = yeast extract medium (1%) with increasing concentrations of maltose); D. Suppression of the Malayan black bug in the field by different
entomogenus fungi, — M. anisopliae (Metsch.) Sorokin, conidia; .... = Beauveria bassiana (Bals.) Vuill., conidia; -..- = Paecilomyces lilacinus
(Thom) Samson, mycelium; -.- = P. lilacinus, conidia; — = M. anisopliae, mycelium; ®® = untreated control.
Fig. 7. A. Beauveria bassiana (Bals.) Vuill. on the brown planthopper; B-C. Beauveria bassiana, sympodial con-
idiogenous cell and cluster of conidiogenous cells; D. Nomuraea rileyi (Farlow) Samson infecting the green
semilooper, Naranga aenescens Moore; E. Paecilomyces farinosus (Holm ex S.F. Gray) Brown & Smith,
whorl of phialides and conidia.
planthopper populations by a single conidia application (Rombach et al., 1986b) and by applications
with dry mycelium (Aguda et al., 1987) (Fig. 6B). Mortality levels exceeded 80% at 3 weeks after
ROMBACH ET AL. 625
spraying. Albornoz and Parada (1984) tested M. anisopliae against Sogatodes oryzicola (Muir). One
application of 109 conidia/ml caused 100% morality after 12 days. A lower dosage of 107 conidia/ml
resulted in a 50% mortality.
Beauveria Species. - Beauveria bassiana, or ‘white muscardine fungus’, is being used for agricul-
tural pest control worldwide (Ferron 1978,1981). Brady (1979c) provides a description, illustration,
and discussion on B. bassiana. In the USSR the fungus is applied against the Colorado potato
beetle, Leptinotarsa decemlineata (Say), the potato ladybug, Epilachna sp., and several forest pests
(Yevlakhova 1979). In China, B. bassiana is used against the European corn borer, Ostrinia nubilalis
(Huebner), and forest pests (Franz and Krieg 1980). B. bassiana commonly infects rice planthop-
pers and leafhoppers. It is found on the leafhoppers Nephotettix nigropictus (Stal) and Nephotettix
virescens (Distant) (Nayak and Srivastava 1979b), and on the brown planthopper (Rao 1975,
Srivastava and Nayak 1978c, Rabindra 1973) in India. Chu and Hirashima (1981) report the fungus
from rice leafhoppers in Taiwan. We collected and isolated B. bassiana from all species of rice
leafhoppers and planthoppers in the Philippines. B. bassiana collections on rice insects are also
reported as ‘white muscardine’ fungus, Spicaria densa (Link ex Pers.) Vuill., and B. densa (Link per
Pers.) Picard.
Beauveria brongniartii (Sacc.) Petch is occasionally reported from rice insects as B. tenella
(Sacc.) McLeod. It differs from B. bassiana in its ellipsoid conidia and absence of dense clusters of
conidiogenous cells. B. brongniartii is, in general, restricted to melolonthid beetle larvae and adults.
We collected the species on white grubs (Holotrichia sp.), serious soil pests in tropical upland rice.
Collections of B. brongniartii from other rice pests are probably misidentifications of B. bassiana.
Hoog (1972) provides detailed descriptions, illustrations, and lists of synonyms of Beauveria spp.
B. bassiana forms white powdery conidial masses that cover the insect (Fig. 7A). Host cadavers
stick to a rice leaf or float on the water. Conidia, 2.5-3.5µm, are globose to broadly ellipsoid and are
produced on sympodial conidiogenous cells that are present in dense clusters on hyphae arising
from the mycelium mat (Fig. 7B,C). Conidiogenous cells have a globose to flask-shaped base, 2-3 ×
2-4µm, with a dented zig-zag shaped rachis up to 20 µm long (Fig. 8B). In culture conidiogenous
cells with an elongated base can occur.
B. bassiana is easily isolated on common mycological media. Malt extract (4%) agar (MEA) or
SDA are good for growth and sporulation. It grows well in liquid media, in which it produces hyphal
bodies (‘blastospores’) and mycelial pellets. The species can be produced by the marcescent
process, and over 10 9 conidia per mg dry mycelium can be produced (Rombach et al., 1988) (Fig.
5). On solid and in liquid media red or yellow oosporeins are excreted (E1 Basyouni et al., 1968).The
species can be mass produced on cheap solid media such as bran, boiled rice and soybeans with
processes similar to those used to produce M. anisopliae. In China B. bassiana is mass produced at
the community level; the starter cultures are grown in small flasks, but the actual mass is produced
in trays or even in open pits in the ground. Media are wheat bran, corn stalk powder, and compost
humus (Chiang and Huffaker 1976). Some B. bassiana strains have a strong antibiotic activity and
can grow in non-sterilized liquid media (Yevlakhova 1979). A high-yielding production method
Fig. 8. A. Paecilomyces lilacinus (Thom) Samson, (left) from the Malayan black bug Scotinophara coarctata (F.),
and (right) in liquid culture; B. Beauveria bassina (Bals.) Vuill., the white muscardine fungus, from plant
and leafhoppers; C. Gibellula pulchra (Sacc.) Cavara, from a spider in the Philippines; D. Nomuraea rileyi
(Farlow) Samson, a fungal pathogen of the hairy caterpillar, Rivula atimeta (Swinhoe), in the Philippines.
using stationary liquid cultures was described by Samsinakova et al., (1981). B. bassiania is a suitable
candidate for production and application where only low technology is available. Ferron (1981)
provides a review of ecology and field application of B. bassiana.
B. bassiana isolates were tested for their virulence against the green leafhopper N. virescens, the
whitebacked planthopper Sogatella furcifera Horvath, and brown planthopper on rice. Differences
in pathogenicity of the different strains were present (Aguda et al., 1984b). Rombach et al., (1986b)
field tested B. bassiana for brown planthopper control in the field. One conidial spray caused high
mortality levels after 3 weeks (Fig. 6B). Aguda et al., (1987) applied dry E. bassiana mycelium for
hopper control on rice in Korea (Fig. 9A, B). Long term control and significant reduction in
ROMBACH ET AL. 627
hopperburned rice plants was

achieved. Also, virulence of B.
bassiana isolates was compared
with other insect fungi (R.M.
Aguda and D. J. Im, un-
publ.,1986). Insects were dipped
in suspensions of conidia and
suspensions of formulated, dry
mycelium of B. bassiana. No dif-
ferences in virulence between
the fungi were detected, and
mortality levels only slowly in-
creased with increasing dosage
of the fungi (Fig. 10).
Hirsutella Species.-Various
Hirsutella species have been col-
lected from planthoppers and
leafhoppers (Soper 1985). Only
H. citriformis Speare and H.
strigosa Petch have been found
on rice hoppers (Rombach and
Roberts 1989). H. strigosa is
commonly found on the brown
planthopper and green leafhop-
per in the insectary and field at
the International Rice Research
Institute, Los Banos, Laguna,
Philippines (Fig. 11D, E). The
fungus covers the host in a light
brownish mycelium that forms
byssoid strands on the rice
leaves. Phialides, 15-35 × 4.5-
6µm, with a cylindrical base and
a stout neck, 12-25 × 1-2µm, are
scattered over the mycelium
strands. Base and neck are
warted. Polyphialides are Fig. 9. Entomogenous fungi for control of (A) the brown planthopper.
present (Fig. 12). Conidia, 9.5- Nilaparvata lugens (Stal), and (B) the whitebacked planthopper,
12.5 × 3.5-6µm, are cymbiform Sogatella furcifera (Horvath) in South Korea. (Bb2000 = 2000 g
formulated dry B. bassiana mycelium/ha; Bb100 = 100 g formu-
with rounded ends or ellipsoid, lated mycelium; BbLG = mycelium formulated the water absorp-
and are produced mostly with 1 tion; Bbcon = B. bassiana conidia, 5x10 12 /ha;, Mf = Metarhizium
flavoviride var. minus, conidia at 5 x1012/ha; Ma = M. anisopliae
conidia at 5x10 12/ha;, control = untreated control).
Fig. 10. Bioassay of fungi against brown planthopper, Nilaparvata lugens (Stal) from South Korea. Insects were
dipped in suspensions of either 5 × 10 7 conidia/ml, or in suspensions of 10 g formulated mycelium/liter. MA
= Metarhizium anisopliae; MF = Metarhizium flavoviride var. minus; BBE, BBPh, and BBel = Beauveria
bassiana, different strains; PF = Paecilomyces farinosus; BBmyc = B. bassiana mycelium.
or 2 in a mucus drop. In pure culture synnemata are produced (Fig. 11D), and the phialides and
conidia are markedly smaller.
H. citriformis (Speare 1920) is widely distributed on rice planthoppers and leafhoppers and can
be found on the brown planthopper in most of the rice growing areas of Asia (Fig. 13 A-D). Brady
(1979b) provides a description and discussion on H. citriformis. Gabriel (1968) illustrated the
species for the Philippines. During periods of high insect density the fungus is often the major
mortality factor; epizootics killing total populations are common and dozens of infected specimens
are clustered on a single stem of rice. H. citriformis has long, filamentous brownish grey to brownish
pink synnemata. The insect is covered in mycelia which also grow for short distances on the plant
surface, attaching the host to the substratum. Synnemata are 1-10 mm long, depending on the stage
of infection; they bear short and stumpy branches. Phialides, up to 45µm long, have a globose to
ellipsoid base, and long and slender needle-like necks. Conidia, 5-8.5 × 2-3µm, are cymbiform with
rounded ends, or ellipsoid, and enveloped in mucus (Fig. 12A). Synnemata are also produced in
culture.
ROMBACH ET AL. 629
Fig. 11. A. Paecilomyces lilacinus (Thorn) Samson infecting the black bug Scotinophara coarctata (F.); B. Gibellula
leiopus (Vuill.) Mains on a spider; C. Hirsutella jonesii (Speare) on the predator Cyrtorhinus lividipennis
Reuter; D. Hirsutella strigosa Petch, synnemata in culture; E. Hirsutella strigosa Petch infecting the green
leafhopper, Nephotettix virescens (Distant).
H. citriformis can be grown on media. SDA is best for isolation and growth. The species is
isolated by touching the synnemata with a sterile, preferably glass, needle to transfer conidia to the
agar. It can also be isolated by transfer of internal mycelia, or hyphal bodies, to the culture plate. H.
citriformis grows slow on solid media and colonies reach a diameter of 1-5 mm after 1-2 weeks of
incubation at room temperature. The competitive ability of H. citriformis in culture is low, and
Fig. 12. A. Hirsutella citriformis Speare, fungal pathogen of the brown planthopper. Note the conidia in mucus;
B. Hirsutella strigosa Petch, fungal pathogen of the green leafhopper Nephotettix virescens (Distant).
cultures are easily overgrown by bacteria, yeasts, and other fungi. Cultures are white, velvety and
wrinkled. After 4-8 weeks synnemata are produced in the center of the colonies (Fig. 3C). Light
does not affect production of synnemata. Phialides and conidia are present on the synnemata and
on the velvety mycelium. A medium sized colony of 5 cm diameter produces about 106 -107 conidia.
This is few compared to the conidia production of the muscardine fungi. Therefore, production of
conidia on solid media is only possible for laboratory studies and small field trials. For mass
production other methods with submerged fermentation have to be developed. H. citriformis can be
propagated in liquid yeast extract/molasses media; in this broth over 10 g dry weight of mycelium
per liter can be harvested from well aerated fermentors (Rombach et al., 1989b). In culture H.
citriformis produces a highly viscous metabolite similar in nature to the polysaccharides produced
by H. gigantea Petch (Loughheed and MacLeod 1958).The rate of excretion of the metabolites
differs among strains. Epizootics of H. citriformis were induced in populations of the brown
planthopper where single applications with a suspension of mycelium and conidia resulted in
mortality levels exceeding 80% at 3 weeks post treatment (Rombach et al., 1986b).
The mycoparasite C. ovalisporum Petch was collected on H. citriformis on the brown planthop-
per in the Solomon Islands (Rombach and Roberts 1987) (Fig. 13 E,F). C. calcarisporium is present
ROMBACH ET AL 631
Fig. 13. A-D. Hirsutella citriformis Speare, at different stages of development on the brown planthopper; E-F.
Hirsutella citriformis infected with the mycoparasite Calcarisporium ovalisporum Petch., early stage of
infection (E), and final stage of infection with the insect fungus completely overgrown with the parasitic
fungus (F).
as a diffuse, purely white mycelium and loosely bound, short and stumpy synnemata on brown
planthoppers infected with H. citriformis. C. ovalisporum grows in culture, but does not sporulate
on media lacking (heat-killed) mycelium of the host fungus. On virtually all H. citriformis infected
brown planthopper specimens collected on the Guadalcanal Plains (Solomon Islands) the
mycoparasite was also found. C. ovalisporum probably has an effect on H. citriformis epizootics in
brown planthopper populations.
2.1.2 Other Hyphomycetes

Morimoto (1957) and Aoki (1941) reported Paecilomyces farinosus (Holm ex S. F. Gray) Brown &
Smith (Fig. 3B) from rice planthoppers and leafhoppers in Japan. Kuruvilla and Jacob (1980b)
reported on its pathogenicity to several insect pests. P. farinosus is a common insect pathogen which
occurs in soil (Samson 1974). Müller-Kögler (1965) reviewed the literature on biological control
with P. farinosus.
Paecilomyces fumosoroseus (Wize) Brown & Smith has a wide host range including Lepidop-
tera and Diptera. The species was isolated from brown planthoppers collected in a field at the
International Rice Research Institute, Philippines. Samson (1974) provided detailed descriptions of
Paecilomyces spp. These fungi are suitable candidates for biological control studies on rice.
Fusarium species have been reported infecting rice insects, including Lepidoptera and Homop-
tera (Hongke 1983, Kuruvilla and Jacob 1979). Hongke reported 51-80% mortality of brown
planthopper populations in bioassay tests. Kuruvilla and Jacob (1979, 1980a) describe field cage
tests with F. oxysporum Schlecht. These tests resulted in total infection of brown planthopper
populations, even at a dose of 1-6 × 106 conidia/ml sprayed on rice plants. Nayak and Srivastava
(1978a) reported F. oxysporum infecting the rice greenhorned caterpillar Melanitis leda ismene
Cramer. Sinha and Prasad (1975) mention the occurrence of F. aleyrodis on larvae of the stem borer
Chilo partellus (Swinhoe).
Caution should be exercised in experimentation with Fusarium species. Only a very few
Fusarium species are considered insect pathogens sensu stricto. Other Fusarium species are not
insect pathogenic or only weakly so, and they include plant pathogens, food contaminants, and
mycotoxin producers. These are factors that limit the use of Fusarium species in biological control.
Extensive research on mass production and field application of Fusarium species is discouraged,
especially because virulent, and obligate fungi such as Beauveria and Metarhizium species that are
safe to humans are available.
2.13 Hyphomycetes on Stink Bugs and Weevils

Few pathogens have been found infecting rice black bugs, Scotinophara spp. (Pentatomidae:
Hemiptera) and the stink bug Leptocorisa oratorius (F.) (Alydidae: Hemiptera). These bugs feed by
sucking, and are free of diseases that infect through oral uptake.
Katsumata (1930) reported on M. anisopliae on Scotinophara lurida Burm. in Japan. Morimoto
(1957) reported on M. anisopliae and P. lilacinus and their use in biological control of S. lurida in
Japan. P. lilacinus was also collected from Scotinophara coarctata (F.) in Malaysia (Fig. 11A);
M. anisopliae (Fig. 3D) and B. bassiana were isolated from this host in Palawan, Philippines. In the
Philippines, B. bassiana is common in populations of L. oratorius (M. C. Rombach and R. M. Aguda,
ROMBACH ET AL. 633
unpubl.), (Fig. 3E). Meneses Carbonell et al., (1980) report on the successful control of the rice
water weevil, Lissorhoptrus brevirostris (Curculionidae) with M. anisopliae in Cuba.
Paecilomyces lilacinus.- P. lilacinus (Thom) Samson occurs on insects, nematodes, and in soil.
Spicaria rubido-purpurea Aoki (1941) is synonymous with P. lilacinus; it was reported on S. lurida on
rice in Japan (Morimoto 1957). P. lilacinus has promise for biological control of nematodes
(Candanedo et al., 1984), in addition to its possible value in biological control of insects. On
Scotinophara short synnemata (up to 1 mm long) are produced on the joints. The mycelium is white,
turning whitish purple at sporulation. Conidiophores have verticillate short branches with whorls of
2-4 phialides (Fig. 8A). The phialides, 7-9 × 2-2.5 µm, are flask shaped with a swollen basal part and
a short neck of 1-1.5 µm long. The conidia, 2-2.5 × 1.2-2µm, are broadly ellipsoidal and in short
chains.
The fungus is easily isolated on PDA, SDA, MA (4%), and YpSs agar. Isolation can be difficult,
because infected insects are often collected from wet paddy mud and are heavily contaminated.
With such specimens internal mycelium or hyphal bodies are used for isolation. P. lilacinus also
produces conidia in liquid culture (Fig. 8A); the phialides are elongated and the conidia vary in
shape and size.
A review by Morimoto (1957) summarizes his earlier work (1952-1954) on the microbial control
of S. lurida by P. lilacinus and M. anisopliae. These fungi infect S. lurida but are relatively safe for
the silkworm. Rice plants with field populations of S. lurida were sprayed and dusted with P.
lilacinus and M. anisopliae. Sprays were more effective than dusts. Also, black bug mortality was
higher in M. anisopliae plots than in P. lilacinus plots where mortality levels of 60-100% for up to 46
days were observed after application with M. anisopliae conidial dust. In one region with low insect
numbers the populations were completely eliminated at 14 days after application. The fungus
spread to untreated adjacent fields and caused high infection levels of up to 100%.
Morimoto (1957) distributed silkworm pupae infected with M. anisopliae in rice hills. These
cadavers generate a natural and constant inoculant to infect immigrating black bugs. This effect of
a more or less continuous conidia supply is also achieved by field application of mycelium particles
from the marcescent process (Rombach et al., 1986c). These particles adhere to the rice plants, and
produce conidia for at least one week in the hot tropics, and for several weeks in cooler temperate
regions such as Korea (Aguda et al., 1987).
Rombach et al., (1986c) reported results of experiments on the microbial control of the black
bug S. coarctata, in the Philippines in which B. bassiana, M. anisopliae, and P. lilacinus were tested
(Fig. 6D). In these experiments single conidia and dry mycelium applications significantly reduced
black bug populations for over a 2-month period.
It should be noted that handling P. lilacinus poses possible safety problems. This species has
also caused keratitis in human patients, but only in those that were under post operative treatment
with cortico-steroids and antibiotics (Kozarsky et al., 1984, Morgan et al., 1984). These “mycoses”
are saprophytic invasions of decomposing organic matter. Active invasion of healthy mammal
tissues by P. lilacinus has not been reported. Although, handling of P. lilacinus in the Course of
biological control programs is most probably safe, overexposure of eyes and the pulmonary system
should definitely be avoided, and simple safety precautions such as the use of dust masks and safety
glasses should be taken.
2.1.4 Hyphomycetes on Lepidoptera

Lepidopteran larvae on rice are infected by entomogenous fungi, but epizootics are rare. Only a few
fungi are of interest for the biological control of rice Lepidoptera.
Nomuraea rileyi (Farlow) Samson is often collected on rice lepidopteran larvae (Kish et al.,
1974). This fungus is well studied because it causes epizootics in populations of noctuid caterpillar
pests on cabbage, clover, soybean, velvetbeans and other crops (Ignoffo 1981, Allen et al., 1971,
Sprenkel and Brooks 1973), and at present there is interest in its commercial production. On these
crops early instar larvae are more susceptible to the fungus than later stages. Ignoffo et al., (1976a,
b) concluded that it is most practical to apply the fungus as a prophylactic agent before damaging
populations occur. In recent years N. rileyi was collected and isolated from rice pests such as
armyworms and cutworms, Mythimna spp, and Spodoptera spp. respectively, the hairy caterpillar,
Rivula atimeta (Swinhoe, Fig. 8D), and the green semilooper, Mocis frugalis (F.) (Fig. 28) and
Naranga aenescens Moore (Fig. 7D). N. rileyi was also collected on larvae of the leaffolders
Cnaphalocrocis medinalis Guenee and Marasmia sp.
Fig. 14. Nomuraea releyi (Farlow) Samson, suppression of hairy caterpillar, Rivula atimeta (Swinhoe.), in the
Philippines. This natural epidemic of the fungus effectively suppresses the second, most damaging, popula-
tion of the hairy caterpillar.
ROMBACH ET AL. 635
N. rileyi produces a compact, pale green to gray-green layer of conidiophores on a white basal
felt of mycelium on the insects. Conidiophores have branches in close whorls near septa with 2-5
phialides on each branch (Fig. 8D). Phialides, 4.7-6.5 × 2.3-3.0µm, have a cylindrical or swollen
base and the neck is absent or greatly reduced. Conidia, 3.5-4.5 × 2-3µm, are broadly ellipsoid and
in dry chains.
The fungus is easily isolated on agar media, but growth is slow; it sporulates well on YpSs and
OA. Information for the mass production of the fungus is limited (Ignoffo 1981).
To date no biological control experiments with the fungus have been carried out on rice in the
field. However, some information on its natural occurrence is available. A natural and dramatic
outbreak of the fungus prevented damage by an increasing population of a Spodoptera sp. (F.
Medrano, per. comm., 1985) on rice in the Philippines. Also, N. rileyi often suppresses outbreaks of
the second generation of the green hairy caterpillar, R. atimeta, in South Cotabato, Philippines (Fig.
14). Bioassay tests with N. rileyi and B. bassiana against larvae of Cnaphalocrocis medinalis Guenee
show an ED50 of 3.5 x 105 conidia/ml (Aguda and Rombach 1987), when the larvae are dipped in
the suspension and incubated on rice leaves. In these tests B. bassiana was more effective in
infecting this insect than N. rileyi. However, differences in susceptibility of insects and virulence of
Nomuraea strains exist (Boucias et al., 1982, Puttler et al., 1976).
P. fumosoroseus has been reported attacking various lepidopteran and dipteran hosts. It was
reported from the European corn borer Ostrinia nubilalis (Huebner) in China (Sheng et al., 1965,
In Samson 1974) and France (Maniania and Fargues 1984). P. fumosoroseus was also isolated from
the striped stern borer Chilo suppressalis (Walker) (Maniania and Fargues 1984). Studies of the
virulence of the isolate for Spodopfera spp. indicated that some insect strains were highly suscep-
tible. Riba et al., (1983) reported that eggs of O. nubilalis were highly susceptible to the fungus. This
is an interesting observation because the egg and neonate larvae of stem borers are the most
vulnerable stages; older larvae hide inside the rice stem.
P. farinosus is a cosmopolitan insect pathogen (Samson 1974) (Fig. 7E, 3B). Biological control
experiments with the fungus are listed by Steinhaus (1949) and Müller-Kögler (1965). P. farinosus
was reported from the striped stem borer C. suppressalis in Japan (Nickel 1964). P. fumosoroseus
and P. farinosus are easily mass produced and applied in the field. They should be further evaluated
for rice stem borer control.
Paecilomyces tenuipes (Peck) Samson (Fig. 3F) is rarely collected on lepidopteran pupae. The
species has been isolated on artificial media, but its value for biological control of lepidoptera is not
known.
B. bassiana is a pathogen of European corn borer, O. nubilalis eggs (Lynch and Lewis 1978),
larvae (Steinhaus 1949), and adults (Smith and York 1960). York (1958) and Riba (1984) reported
high mortalkty in field populations of O. nubilalis after B. bassiana applications. In China, microbial
control of this borer by B. bassiana is practiced over large areas (Hussey and Tinsley 1981). B.
bassiana infects rice stem borers (Rao 1975, Yadava et al., 1979, Math et al., 1966, Srivastava and
Mathur 1970), and the rice skipper, Pamara mathias F. (Nayak et al., 1978a). The closely related
fungus B. brongniarti was also tested for rice skipper infection (Nayak et al., 1978a). Experiments on
biological control of rice stem borers and other rice Lepidoptera with B. bassiana have been
restricted to laboratory bioassay tests (Nayak et al., 1978b, Nayak and Srivastava 1979b, Yadava et
al., 1979). N’Doye (1976) observed reduced oviposition rates and viability of C. suppressalis eggs in
survivors of a mycosis. Laboratory bioassay tests show that leaffolder Marasmia sp. larvae can easily
be infected with B. bassiana (M. C. Rombach and R. M. Aguda, unpubl.).
B. bassiana is relatively simple to mass produce on solid media as well as with the marcescent
process (Fig. 5). It is safe for humans and beneficial organisms. Therefore, this pathogen is a prime
candidate for experimentation of microbial control of rice stem borers and leaffolders.
2.2 ENTOMOPHTHORALES
Entomophthoralean fungi (Zygomycetes) commonly cause epizootics in insect populations (Wild-
ing 198l), and several of these fungi have potential for commercial development. In this group the
conidia are forcibly discharged by turgid cells and are launched over distances of several cen-
timeters. This results in an aureole or halo of conidia on the plant substratum surrounding dead
hosts (Fig. 2D). The forced discharge of the conidia can be exploited when culturing Entomoph-
thorales. Dead hosts are placed on the inside of a petri dish lid and the agar surface placed above
the cadaver, and conidia will be dispersed on to the agar. The most adequate medium for isolation
of entomophthoraleans is coagulated egg-yolk medium/SDA combination (Stevens 1974). A mycos-
tatic agent, Mycostatin®, and antibiotics such as a penicillin — streptomycin combination or Gen-
tomycin® can be added to inhibit contaminant growth.
Entomophthoralean fungi grow vegetatively as short and irregular hyphal segments or hyphal
bodies. The life cycle includes thick walled resting spores and zygospores which are produced in the
body cavity of the host; these structures are produced after conjugation of adjacent hyphal cells.
These resistant spores survive periods of adverse environmental conditions.
Entomophthoralean taxonomy became increasingly complicated after the basic studies of
Roland Thaxter (1888) a century ago. Humber (1981) reviewed the most recent taxonomic criteria
and taxonomic keys are provided by King and Humber (1981) and Waterhouse and Brady (1982).
At present there are about 150 species of entomophthoralean insect fungi. Of these six occur
on planthoppers and leafhoppers worldwide, with three species on rice planthoppers and leafhop-
pers.
2.2.1 Entomophthorales on Planthoppers and Leafhoppers

Erynia (Entomophthora) delphacis (Hori) is an obligate pathogen of planthoppers, leafhoppers and
spittlebugs (Latg et al., 1980). It is found on rice hoppers in Asia (Fig. 2C). E. delphacis conidia,
15-33µm long, are elliptical to ovoid with a papillate base. The conidiophores are branched and
sometimes simple. Resting spores and rhizoids are not produced.
E. delphacis closely resembles the aphid pathogen Erynia neoaphidis Remaudiere & Hen-
nebert. Entomophthora aphidis Hoffman is the older synonym of Erynia neoaphidis; E. delphacis
ROMBACH ET AL 637
was reported several times as E. aphidis on rice (e.g. Hinckley 1963). E. delphacis can be isolated
and cultured on egg-yolk medium and SDA. Growth is optimum at a temperature of 25°C and a pH
of 6.3-6.7 (Shimazu 1976). Optimum temperature for germination is 25°C, and a relative humidity
exceeding 93%, but preferably free water, is necessary for germination (Shimazu 1977).
E. delphacis has not yet been applied to control field populations of rice hoppers. But, this
pathogen is a suitable candidate for biological control. It can be isolated in pure culture and mass
produced by the marcescent process (R. S. Soper, pers. comm. 1986).
Erynia radicans (Brefeld) Humber, Ben-Ze'ev & Kenneth is often incorrectly referred to as
Entomophthora sphaerosperma Fres. or Zoophthora rudicans (Brefeld) Batko. Turian (1957)
proposed to institute E. rudicans cicadelliphaga for the fungus attacking leafhoppers. The fungus is
collected worldwide on plant and leafhoppers. It was collected on the brown planthopper in
southeast China (Shimazu 1979), and Samal et al. (1978) reported E. radicans as E. fumosa; the
latter fungus is synonymous with Neozygites fumosa (Speare) Remaudiere & Keller. Ben-Ze'ev &
Kenneth (1981) provided a detailed description of E. radicans.
In E. radicans rhizoids attach the insect to the plant. Conidiophores form a compact layer on
the host. Primary conidia, 16-24 × 5.5-8 µm, and secondary conidia, 13.5-18 × 6-9 µm, are ellip-
soidal; capilliconidia, 19-24 × 4.5-8 µm, are almond shaped. The fungus can produce globose
resting spores, 22-29 µm, inside the cadaver.
E. radicans causes devasting epizootics in populations of Empoasca kraemeri Ross & Moore on
cowpea in Brazil (Daoust et al., 1983) and brown planthopper in Asia (Shimazu 1979). The fungus
can be readily cultured on artificial media and mass production through the marcescent process is
feasible. Application against rice hoppers should be tested.
Conidiobolus coronatus (Cost.) Batko is common on rice planthoppers and leafhoppers (Fig.
2D). King (1976a,b; 1977) provided information on the taxonomy and biology of C. coronatus.
Primary conidia, 14-39 × 17-49µm, are spherical to pyriform with rounded apex, and multinucleate.
Microconidia, 7.5-2.5µm, are globose and attached to the outside of the original conidium. Villose
conidia are present on the host and in pure culture.
In certain C. coronatus strains the conidia can actively transverse the sclerites and the
membranes of insects (Yendol and Paschke 1965), and these strains are insect pathogens sensu
stricto. However, other strains are a saprophytic, grow in soil, or cause human and equine en-
tomophthoromycosis (King 1979).
The brown planthopper was controlled by one application of C. coronatus conidia (R. M.
Aguda, pers. comm. 1984). Padua and Gabriel (1975) developed a coconut-milk medium for its
mass-production. We consider this fungus hazardous for biological control experimentation be-
cause of the possibilities of mammalian infection. Its mass production and field application is
discouraged.
Zoophthora radicans (Brefeld) Batko was collected from the leaffolders. Marasmia sp. and
Cnaphalocrocis medinalis Guenee (M. C. Rombach and R. A. Humber, unpubl. 1988 (Fig. 15). The
fungus causes extensive epizootics in greenhouse leaffolder populations, but is rarely collected in
the field.
22.2 Entomopthorales on Grasshoppers

Entomophaga grylli (Fres.) Batko occurs in field populations of grasshoppers worldwide and
epizootics can significantly reduce populations (Pickford and Riegert 1964, McDaniel and Bohls
1984). E. grylli was occasionally reported from lepidopteran and dipteran hosts. E. grylli has globose
to pyriform multinucleate conidia with prominent papillae. Brady (1979a) provides a description.
Grasshoppers infected with E.grylli climb to the top of plants and grasp the plant with the front pair
of legs; large numbers of infected grasshoppers are mostly found attached to plants during
epizootics. In rice E. grylli was found on unidentified species of grasshoppers in Indonesia (R. A.
Humber, pers. comm, 1984) and Bhutan (B. M. Shepard, pers. comm. 1987). Weiser et al. (1985)
reported it from Oxya hyla intricata (Stahl) a rice pest in Vietnam, where the fungus infected 85-90%
of the population. Roffey (1965,1968) listed the fungus on a number of minor grasshopper pests of
rice from Thailand.
2.3 FUNGI ON PREDATORS

Several entomogenous fungi infect beneficial insects (predators) and spiders in rice fields. The
fungi infecting beneficials are all specific for certain insects or spiders, and always occur at low
levels; epizootics of these fungi have not been reported.
Gibellula leiopus (Vuill.) Mains and G. pulchra (Sacc.) Cavara were commonly found on
spiders in rice fields (Fig. 11B, 8C). G. leiopus has a compact appearance due to the short
synnemata and the short conidiophores (Fig. 11B). In recent collections the fungus was found on
small spiders (e.g., Callitrichia spp.); it has not been reported from lycosids, one of the most
important groups of predators in rice. Not much ecological information on Gibellula spp. is
available. Samson and Evans (1973) concluded that G. pulchra was an important mortality factor for
spiders on cacao trees in Ghana. In our experience these fungi are a minor mortality factor for
spider populations in rice, with infection levels never exceeding 1% of the population.
Hirsutella (Synnematium) jonesii (Speare) Evans and Samson was described from Mezira sp.
(Hemiptera). It was collected on rice in the Philippines on Anthicus sp. (Coleoptera), and on
Cyrtorhinus lividipennis (Reuter) (Hemiptera) (M. C. Rombach, unpubl.) (Fig. 11C). However, the
incidence of this fungus was always below 1%, even in dense Cyrtorhinus populations.
3. ENTOMOGENOUS BACTERIA
3.1 BACILLUS THURINGIENSIS
Bacillus rhutingiensis Berliner (B.t.) is the most widely used microbial for control of insects (Burges
1982, Samson et al., 1986). B.t. produces endogenous spores and toxins (Fig. 16). The resistant
spores enhance survival in the field and in storage, which facilitates formulation and application of
ROMBACH ET AL. 639
Fig. 15. Zoophthora radicans (Brefeld) Batko. Stages of infection of the rice leaffolder, Cnaphalocrosis medinalis
Guenee. A. A healthy larva; B. vegetative fungus growth inside the host; and C. sporulation on the cadaver.
the B.t. products. The spores, and endotoxin, have to be ingested by the insect and thus toxicity for
non-phytophagous fauna is negligible. Microbial control with B.t. products can safely be combined
with biological control by natural enemies. Also, nearly all B.t. entomotoxins are safe for man and
other mammals. A list of non-target fauna not susceptible to B.t. is provided by Burges (1982).
B.t. is the most commercially successful insect pathogen. At present about US $40 million worth
of B.t. products are sold worldwide. Most of these materials are used against lepidopteran pests on
a number of field crops, and for mosquito larva control. Recently B.t. strains isolated from
coleopteran hosts were reported from Germany and the USA. Products containing spores of these
strains will soon be available.
After ingestion of the spores and protein endotoxin crystals the insect gut and mouthparts are
paralyzed, feeding is arrested and no further crop damage occurs. The insect dies soon thereafter.
Reports of epizootics of B.t. in natural insect populations are rare (Burgerjon and Martouret 1971).
This is probably because the lethal bacterial structures, such as spores and toxic crystals, appear
only in small quantities in insect cadavers (Angus l968). Thus recycling of B.t. in insect populations
is practically absent and repeated application is necessary for control.
B.t. grows on simple artificial media and can be produced by solid and liquid fermentation
processes. Several commercial products with different B.t. varieties and serotypes have been
produced.
Fig. 16. Bacillus thuringiensis Berliner. Note the “footprint” appearance, i.e. the spore with the parasporal inclusion
body.
The prerequisite of oral uptake limits the host range of B.t. In rice primary targets are
lepidopteran larvae such as leaffolders, armyworms, hairy caterpillars and, to a lesser extent, stem
borers. B.t. was described from Anagasta (Ephestia) kuhniella Zell., a stored product pest (Berliner
1915). Other lepidopteran pests of stored products have been effectively controlled by B.t. (Burges
1964). Recently, B.t. products were approved for use in stored grains in the United States. It is very
effective against the Indian meal moth, Plodia interpunctella, and almond moth, Ephestia cautella
(McGaughey 1976,1978), however, few studies on stored rice insects are reported.
The leaffolder Cnaphalocrocis medinalis Guenee was controlled by four commercial B.t.
products (Srivastava and Nayak 1978a). The authors claim significantly higher mortality by similar
doses of Dipel® and Thuricide® as compared to Bactospeine® and Cajrab®
Bioassay techniques to test the effectiveness of B.t. materials for the striped stem borer Chilo
suppressalis (Walk.) were developed by Rombach et al., (1989a). B.t. products incorporated in
artificial diet of the stem borer larva greatly inhibited feeding, but mortality was relatively low.
Nayak et al., (1978b) reported control of the stem borers Scirpophaga incertulas (Wlk.), Sesamia
inferens W., and Chilo sp. by Thuricide®. All larval stages can be infected by the product by a cut
stem technique in which a small piece of rice stem is soaked in B.t. spore suspension prior to
feeding. Yadava (1978) obtained high larval mortality of the pink stem borer, S. inferens using this
technique.
Internal parts of rice stems are, in general, free of pathogens. Thus, in the field, only neonate
stemborer larvae are exposed to B.t. when they feed on the stem or on the leaves before tunnelling
ROMBACH ET AL. 641
into the stem. Nayak et al., (1978b) sprayed a 1% Thuricide suspension in the greenhouse during
egg hatching. Dead hearts and numbers of living larvae in tillers were reduced by 76%, and white
heads by 67% respectively. Product activity persisted for 15 days. However, under field conditions
the spores and crystals will rapidly deteriorate due to UV light and weekly sprays may be necessary
to kill neonate larvae before stem invasion. This would be prohibitively expensive for practical use,
unless a method of precise timing can be developed.
Adverse effect of sunlight on B.t. was tested in China (Anonymous 1981). A spore preparation
was applied to rice infested with second instar leaffolder larvae at 8:00 a.m. and at 4:00 p.m.
Morning treatments resulted in 22-39% mortality and afternoon treatments in 58-72% mortality.
Bounias and Guennelon (1974) tested B.t. for control of the striped stem borer C. suppressalis.
This stem borer was introduced into Camarque (France), and its populations were mostly free of
natural enemies. High mortality of larvae was obtained when fed artificial diet with B.t. spores. The
authors conclude that B.t. can be used for stem borer control only if a two-month period of plant
coverage is ensured during egg hatching. We estimate that, under tropical conditions, this will
require at least four to five B.t. applications.
B.t. research in Japan was delayed for more than a decade due to quarantine measures for
protection of the sericultural industry (Aizawa 1967). Recently B.t. strains with low toxicity for
silkworms and high toxicity for target pests were developed through selection (Aizawa et al., 1975).
Also, strains were selected that were capable of producing the endogenous entomotoxins but no
spores (Nishiitsutsuji-Uwo et al., 1975). These strains will not disperse from the treated crop to
mulberry trees with silkworm larvae. In Japan B.t. was effective against the rice plant skipper,
Pamara guttafa (Aizawa 1976).
In China B.t. is intensively researched for control of insect pests on rice (Chiu 1984, Hussey and
Tinsley 1981). Good control of rice skipper larvae (Pamara spp.) was reported after only one B.t.
application. Combinations of B.t. with low dosages of a chemical insecticide (e.g. trichlorfon at
10-20% of the recommended rate) were reported to be most effective.
China is one of the world’s largest B.t. producers, and production is mainly carried out on the
village level using solid fermentation on wheat bran, corn meal, soybean, and cotton seed cake.
However, recently large production facilities using liquid fermentation were installed.
Hussey and Tinsley (1981) estimated that in China about 2,500 metric tons of B.t. are produced
annually; most of this material is used for control of lepidopteran pests in forests.
3.2 OTHER ENTOMOGENOUS BACTERIA

Philip and Jacob (1982) reported Bacillus cereus Frankland & Frankland from the leaffolder C.
medinalis. Bacillus subtilis (Ehrenberg) Cohn was reported from the green horned caterpillar,
Melanitis leda ismene Cramer (Nayak and Srivastava 1978b). These bacteria produce spores but not
entomotoxic protein crystals. B. subtilis is a common soil bacterium (Schroth and Hancock 1981).
Its insect pathogenicity is doubtful; it is an opportunistic invader of weakened insects. The value of
E. cereus and B. subtilis in biological control of rice pests is probably limited.
Some non-sporulating bacteria are insect pathogens (Buchner 1963). Srivastava and Nayak
(1978b) reported the red pigmented Serratia marcescens Bizio from field collected larvae of the
green horned caterpillar Melanitis leda ismene Cramer and pink stem borer S. inferens. Isolations
infected three different stem borer species, a leafroller and the rice skipper in the laboratory. These
test insects were continuously fed with the pathogen (Srivastava and Nayak 1978b, Narayanan and
Jayaraj 1974). However, this bacterium does not produce spores or entomotoxins; therefore its
killing action is slow and mass-production and storage is virtually impossible. We consider S.
marcescens of doubtful insect pathogenicity and probably of no value in biological control of insects.
Occasionally planthoppers and leafhoppers are found infected by entomogenous bacteria such
as B. thuringiensis (Li 1982) and S. marcescens (Rao and Rao 1979). However, these bacteria are not
effective against sucking insects which feed in the vacular tissues where these pathogens do not
occur.
4. ENTOMOGENOUS VIRUSES
Over 1,200 virus-insect relationships in more than 800 insect and mite species were reported by
Martignoni and Iwai (1981) and Martignoni and Longston (1960), and numbers have increased
since. Many insect viruses possess promising properties for biological control, especially the
baculoviruses and reoviruses, which have stable inclusion bodies (Entwistle 1978). In some instan-
ces the viral pathogens can be introduced for insect control, and can recycle efficiently from one
insect generation to another. But in some annual cropping systems the insect viruses are applied
similar to a chemical insecticide (i.e. repeated applications when pest numbers increase). This type
of control is often employed in unstable agricultural systems where insect populations decline at the
end of cropping seasons. Examples of this method are control of cotton bollworm by Elcar® and
control of Heliothis zea Boddie on soybean (Ignoffo et al., 1978). Payne (1982) reported six insect
families with viruses and reviewed the ecology and insect control efforts using these microbials.
Insect viruses are classified into families according to morphology of the virus particle, chemi-
cal and physical nature of the nucleic acids, and presence or absence of inclusion bodies.
Transovum transmission of nuclear polyhydrosis viruses (NPVs; Baculoviridae) from genera-
tion to generation has been demostrated. However, transmission by oral uptake of the infective
particles by the host is more common, and thus the insect stages feeding on plants are most
vulnerable to infection. Sucking insects such as planthoppers and leafhoppers and hemipteran
insects are not infected by insect viruses.
The family Baculoviridae contains the subgroups of the nuclear polyhedrosis viruses (NPVs),
granulosis viruses (GVs), and Oryctes viruses. Both NPVs and a GVs are represented in rice insects.
Baculovirus species (Baculoviridae) are bacilliform, contain circular double stranded DNA
and can form proteinaceous inclusion bodies. No viruses with a similar morphology have been
observed in plants or in vertebrate hosts (Payne 1982).
The NPVs have inclusion bodies produced in the insect tissues at the terminal stage of infection
(Summers 1977). Viral particles embeded within the inclusion body are released in alkaline pH
ROMBACH ET AL. 643
(e.g. in insect guts). The main difference between the NPVs and the GVs is the presence in the
majority of GVs of only one single virus particle in each ‘capsule’ or inclusion body. In NPVs a
variable number of virus particles are embedded in the matrix of the polyhedra.
Reoviridae are represented by a cytoplasmic polyhedrosis virus (CPV) of the rice leaffolder C.
medinalis. Baculoviridae and Reoviridae are promising for microbial control because they produce
stable inclusion bodies.
Iridoviridae are represented by the Chilo iridescent virus (CIV). The virus does not produce
inclusion bodies; its utility in biological control is limited.
4.1 NUCLEAR POLYHEDROSIS VIRUSES (NPVs)

An NPV produced polyhedral inclusion bodies in tissues of laboratory reared pink stem borer
Sesamia inferens (Godse and Nayak 1983). The NPV infected larva has a whitish ventral side,
gradually progressing to the end of the body and most prominent at the intersegmental membranes.
A few days after infection the host dies. The dead larva is whitish brown. Cadavers lodge within the
plant or hang on the outside of the stem. The liquefied body contents contain numerous polyhedral
inclusion bodies with an average size of .84 µm. Rod shaped viral particles with a size of 35 ×
215 nm are present in the inclusion bodies. Virus particles are bundled in envelopes in groups of
2-3; bundles are randomly distributed in the inclusion body.
Other NPVs infected Spodoptera mauritia Boisduval (Jacob et al., 1973), grass webworm,
Herpetogramma licarsisalis (Walker), corn semilooper Chrysodeixis chalcites Esper (Im et al., 1989a)
and the rice moth Corcyra cephalonica St. (Rabindra 1973), the rice skipper Pelopidas (Pamara)
mathias (Fabr.) (Nayak and Srivastava 1979a) and the common cutworm Spodoptera litura (Fab.)
(Tsai et al., 1978, Im et al., 1989a). Pathogenicity and histopathology of the latter virus was reported
by Im et al. (1989a). No further information on these viruses is available.
Over 70 years ago Chapman and Glaser (1915) reported ‘wilt disease’, an NPV, in the ar-
myworm Pseuduletia unipuncta (Haworth), and natural epidemics still frequently occur (Diliwari et
al., 1981). Mythimna separata was reported to suffer from an NFV (Neelgund and Mathad 1972, Im
et al., 1989b) (Fig. 17). The virus was multiplied and effectively controlled M. separata in the field
(Neelgund 1975, Mathad and Neelgund 1973). The larvae die in a peculiar position, hanging down
with the prolegs clinging to the plant (Fig. 4D). Manjunath and Mathad (l978,1979,1981a, 1981b)
studied the effect of sunlight and temperature on Mythimna NPV in preparation for field applica-
tion. Bioassay tests to determine the virulence of the pathogens were reported by Neelgund and
Mathad (1974,1978) and Watanabe and Tanada (1972a, b). Dhaduti and Mathad (1978,1979,1981)
tested the effect of Mythimna NPV on silkworm. No significant infestation was obtained. These
NFVS are promising biological control agents, and should be pursued further for rice armyworm
control.
4.2 GRANULOSIS VIRUSES (GVs)

The GV of the rice brown semilooper, Mocis frugalis was first reported from Palawan Island in the
Philippines (Im et al., 1986a). Infected larvae were yellowish, but slowly darkened when the
Fig. 17. Mythimna separata (Wlk.) nuclear polyhedrosis virus (NPV): A. In 1st instar larva 2 days post infection.
The lower hypodermis cell is healthy, the upper one is infected with a virogenic stroma and NPV virions;
B. At 4 days post infection. Polyhedral inclusion bodies and virions.
infection progressed. Refractive granules were found in the fluid of the fat bodies. The virus was
purified and capsules were ellipsoidal but some were irregular and sickle shaped with varying
lengths. The capsules measured 433 × 243 nm, and contained a single virion of 301 × 83 nm.
Eighteen amino acids occurred in the capsules, including basic amino acids (lysine and arginine)
and acidic amino acids (aspartic acid and glutamic acid). However, low quantities of histidine,
cystine, methionine, alanine, tyrosine and tryptophan were found (Im et al., 1986b).
A GV of the rice leaffolder, Cnaphalocrocis medinalis (Guenee) was also reported from the
Philippines (Im et al., 1989b) and India (Jacob et al., 1971). The virus capsules were ellipsoidal in
shape and measured 413 × 234 nm. The size of the virions of this virus was 280-320 × 56-62 nm.
4.3 IRIDESCENT VIRUSES (IVs)

IVs are a heterogeneous group of larger viruses pathogenic to insects. IVs are described as
icosahedral cytoplasmic desoxyriboviruses (Kelly and Robertson 1973). The particles are isometric,
contain linear double stranded DNA, and no inclusion bodies are produced. The group derives its
name from a physical property of the virus particles — if light strikes the compact virus pellets in host
ROMBACH ET AL. 645
tissues a violet, green, or blue hue is produced. Tinsley and Kelley (1970) concluded that this is a
constant feature of all known viruses in the group and suggested that the group name be retained.
The Chilo IV (CIV, or iridescent virus type 6) was described from stem borers (Fukaya and
Nasu 1966). No information is available on the dispersion and infection of hosts under natural
conditions. Multiplication of the virus was carried out by contamination of artificial diet and feeding
of Chilo larvae. After 12 days the larvae became milky white although heavily infected larvae had a
bluish cast. Diseased larvae survived for a considerable time. Electron microscopic examination
revealed large enveloped virus particles (160µm). Ishikawa and Muroga (1976) tested the influence
of CIV on larval development of C. suppressalis, and the greater wax moth, Galleria mellonella L.
Both insects were easily infected by virus injection and severe abnormalities during pupation were
observed. Replication of CIV was studied by Kelly and Tinsley (1974a, 1974b), the activity of its
ribonucleic acid polymerase by Kelly and Tinsley (1973) and biochemical aspects of its non-struc-
tural proteins were reported by Kelly and Tinsley (1972).
5. ENTOMOGENOUS NEMATODES
Pathogenic nematodes can control some insect pests (Poinar 1979). Commercial products contain
Heterorhabditis sp. for control of the black vine weevil, Otiorhynchus sulcatus F. in greenhouses
(W. R. Simons, pers. comm. 1985), Neoaplectana carpocapsae Weiser for control of various ter-
restrial insects, and Romanomermis culicivorax Ross & Smith for mosquito control. For steiner-
nematid and heterorhabditid nematodes a novel mass rearing method is now available (Bedding
1984). Keys to insect nematode species and information on their ecology and applications is given
by Poinar (1979).
5.1 NEMATODES ON HOPPERS

In rice the nematode Agamermis unka Kaburaki & Imamura is a parasite of adults and nymphs of
leafhoppers (Nasu 1967), but its incidence in natural populations is low. The natural incidence of
parasitism by indigenous insect nematodes on brown planthopper, Nilaparvata lugens (Fig. 18),
whitebacked planthopper, Sogatella furcifera, and green leafhopper, Nephotettix virescens, was
monitored at twelve sites in Laguna Province, Philippines, during the wet and dry seasons from
1982-1984 (Pena and Shepard 1985). The highest incidences of parasitism were 36% and 50% for
the whitebacked and brown planthopper, respectively. Parasitism levels were highest in the wet
season. This is probably due to the higher relative humidity, which is important for nematode
survival. Mean levels of parasitism during this season across all the locations revealed that only 19%
of the brown planthoppers and 17% of the whitebacked planthoppers were parasitized. It is unlikely
that these low parasitization levels have a significant impact on hopper populations.
5.2 NEMATODES ON STEM BORERS

Species of Agamernis spp., Amphimermis, Cordius sp., and Mermis sp. were reported on rice stem
borers (Lippold 1972). Strain DD-136 of Steinernema feltiae Filipjev is the only nematode reported-
Fig. 18. Brown planthopper, Nilaparvata lugens (Stal) infected with an entomogenous nematode.
ly tested on rice insects. The bacterium Achromobacter nematophilus Poinar & Thomas is as-
sociated with this nematode. Ramakrishnan and Kumar (1981) review stem borer research with this
nematode in India and Meneses-Carbonell (l983) reports on experiments of water weevil control
with DD-136 in Cuba. Control of the cutworm, Platysentra (Cirphis) compta (Walker) with DD-136
was reported by Israel et al., (1969). However, this approach was not widely adopted, Other target
insect pest species for DD-136 were Scirpophaga incertulas (Walker) (Yadava and Rao 1970, Torii
1975, Rao et al., 1971), and Chilo suppressalis (Torii 1975, Torii and Yasumatsu 1973). Treatments
of small field plots with a nematode suspension failed to reduce S. incertulas populations, although
some infected larvae were observed (Rao et al., 1971). Numbers of ‘white-heads’ (panicles with
empty grains) were not significantly reduced in nematode (DD-136) treated plots. However, sprays
with nematodes were effective in reducing numbers of C. suppressalis larvae (Yadava and Rao
1970). The lack of prevention of whiteheads is attributed to the cryptic nature of stem borer larvae
and the fact that they enter the stem and cause damage even after they are infected with nematodes.
Insect nematodes can infect several insect pests of rice; they surely contribute to the overall
mortality levels of pest insects. Stem borer control is complicated and virtually no effective control
measures are available; the use of entomogenous nematodes, S. feltiae in particular, should receive
further research emphasis.
ROMBACH ET AL. 647
ACKNOWLEDGEMENTS
The preparation of this chapter was partly funded by grants of the Office of the Science Adviser, US
Agency for International Development, Washington DC, and the Nessie Smith Noyes Foundation,
New York, to the Boyce Thompson Institute (BTI) for Plant Research at Cornell, Ithaca, NY, and
by the International Rice Research Institute (IRRI), Los Banos, Philippines.
The encouragement of Dr. Donald W. Roberts (BTI, Insect Pathologist and Program Coor-
dinator, Insect Pathology Resource Center) and Dr. B. M. Shepard (IRRI, Entomologist and Head,
Entomology Department) is highly appreciated. G. Maureen Rombach is thanked for the editorial
help and the preparation of the illustrations. Dr. R. A. Samson (Centraalbureau voor Schimmelcul-
tures, Baarn, The Netherlands) is thanked for providing the EM scanning photographs of the fungi,
and Dr. P. F. Entwistle (Institute of Virology, Oxford, England) for the EM transmission
photographs of the NP virus.
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9
Insecticide Management in Rice
S. CHELLIAH AND M. BHARATHI

Tamil Nadu Rice Research Institute
Aduthurai, Tamil Nadu, India
1. INTRODUCTION
INSECTICIDES SERVE AS IMPORTANT components in the management of rice insect pests. Insec-
ticides are the only tool in the pest management strategy that is reliable for emergency action when
insect pest populations approach or exceed the economic threshold. In the words of the Committee
on Plant and Animal Pests, National Academy of Sciences, “a major technique such as the use of
pesticides can be the very heart and core of integrated systems. Chemical pesticides will continue
to be one of the most dependable weapons of entomologists for the foreseeable future” and this is
true for rice insect pest management also. Conversely, insecticide use results in several undesirable
effects. Frequent and continuous insecticide use leads to development of insecticide resistance in
pests, sudden outbreak of major and secondary pests after insecticide application, destruction of
beneficial insects, and human and animal poisoning. Added to these, the escalating cost of insec-
ticides is another factor which discourages the use of insecticides for crop protection. This paper
deals with the status and strategies of insecticide use in rice.
Insecticide use on traditional rice cropping systems was minimal. However, in the last two
decades, insecticide use on rice has greatly increased, as improved varieties with greater yields, and
profit potential have been planted.
The evolution of chemical control in rice since the later part of the middle period in Japan has
been recorded by Suenaga and Nakatsuka (1958) and Matsuo (l96l). The use of whale oil which
was found effective in 1670, had spread throughout the country by 1840. Kerosene began to replace
whale oil in 1897. Japan, Taiwan and China have used insecticides to control rice insect pests since
modern insecticides were introduced after World War II.
658 INSECTICIDE MANAGEMENT IN RICE
The introduction of DDT and BHC in the late 1940s and of parathion in the 1950s established
the technical basis for pest control with insecticides in Japan. Organophosphorus insecticides were
first used in 1952. Diazinon and malathion soon replaced parathion. Since 1964, carbamate insec-
ticides have been used in Japan. The insecticides were effective against stem borers, leafhoppers
and planthoppers. To increase rice production in the 1950s, the Japanese government subsidized
insecticides and application equipment and provided technical support to farmers which resulted
in wide scale use of insecticides for rice insect pest control. A similar revolution also occurred
throughout tropical Asia.
It was estimated in 1980 that $3.9 billion was spent on insecticides worldwide and $563 million
(14.4%) for rice alone (Lim and Heong 1984). Yield increases attributed to insecticide use have
been experienced under varied conditions in many rice growing countries. As early as 1967, it was
estimated that an average of 44% yield increase was obtained in rice in Taiwan as a result of
insecticidal protection for the rice crop (Tao and Yu 1967). Ku et al. (1980) reported that in Taiwan,
the increase in yield of insecticide treated crops over that of untreated ones ranged from 8.9 to
23.4% in the first crop and 13.8 to 54.2% in the second crop. At the International Rice Research
Institute (IRRI), yield increases of 2.7 t/ha due to insecticide use were reported in experiments
conducted during 1964-1971 (Pathak and Dyck 1974). Similar experiments on farmers’ fields in the
Philippines showed a 20-25% yield increase. In the Philippines, rice yield increases ranging from
0.44 to 3.19 t/ha due to insect pest control by insecticides have been reported (IRRI 1975, 1977;
Heinrichs 1978; Litsinger et al., 1980).
In Malaysia, Lim (1972) reported that insecticides increased rice yields ranging from 0.72 to
1.35 t/ha. Gunasena et al. (1977) observed a yield increase of 0.7 t/ha in farmers’ fields in Sri Lanka,
due to insecticide use. As high as 2.6 t/ha increase in rice yield occurred in Bangladesh (Ahsan and
Hoque 1977). In Japan, yield loss decreased from 419 kg/ha in 1949-1951 when insecticide use for
rice insect pest control just began to 153 kg/ha in 1979-1981 when chemical use had become
extensive (Ishikura 1984).
Insecticide use in rice has several advantages and disadvantages; the advantages are:
• Insecticides afford the only practical control measure for insect pest populations approach-
ing or at the economic threshold.
• Insecticides have rapid curative action in preventing economic damage.
• Insecticides offer a wide range of properties, uses and methods of application.
• Benefit/cost ratios for insecticide use are generally favorable.
• Insecticides are generally readily available for pest control.
Disadvantages are:
• Development of resistance in the pests to the insecticides used — both cross resistance and
multiple resistance. Resistance to pesticides has been reported in the brown planthopper,
green leafhopper, whitebacked planthopper, smaller brown planthopper, striped stem
borer, rice bug and rice water weevil.
• Resurgence of primary and secondary pests,
CHELLIAH AND BHARATHI 659
Adverse effects on non-target species (e.g., natural enemies, honey bees, other pollinators,
fish and wildlife).
Hazards of pesticide residues in food products and in the environment acting as potential
'hidden costs.'
Human poisoning.
Increased cost of chemicals and application equipment.
Phytotoxicity due to application of the wrong pesticide, improper pesticide mixtures, incor-
rect timing of application, wrong formulations, using insecticides at more than the recom-
mended dosage, inadverdent application of sediments from the spray tank and use of
brackish water for insecticide mixing.
Although there are no insecticides to be pronounced as completely safe chemicals, there are
safe ways to use even the most toxic agents. In this context, there should be a harmonious
management approach in using insecticides for pest control.
2. CHEMICAL CONTROL OF RICE INSECT PESTS
2.1 PLANTHOPPERS
Planthopper pests in Asian rice include the brown planthopper (BPH), (Nilaparvata lugens),
whitebacked planthopper (WBPH), (Sogatella furcifera) and smaller brown planthopper (SBPH),
(Laodelphax striatellus).
2.1.1 Effective Insecticides

Several insecticides of various groups have been reported to be effective against the BPH. Chlor-
dimeform and isoprocarb at 0.5 kg ai/ha (Velusamy et al., 1978) and carbofuran, quinalphos,
phosphamidon and chlorpyriphos (Rao and Rao 1979a) were effective in BPH control. Mixtures of
HCH (BHC) and parathion when applied in rice fields, were very efficient in controlling BPH but
adversely affected predator populations in China (Li et al., 1983). Riddell (1982) stated that
carbosulfan gave maximum control of BPH both in greenhouse and field tests in Malaysia. In India,
the field tests conducted at Kerala revealed that carbosulfan effected quick knock down effect of
BPH (Pillai et al., 1983). Granular insecticides such as carbofuran, phorate and isofenphos yielded
a high degree of BPH control (Koshaiya et al., 1981). Patnaik et al., (1986) reported that the granular
formulations of BPMC and carbofuran consistently gave high control of BPH.
Growth regulating compounds having juvenile hormone and molting hormone effects have also
been used for rice pest control. Kajihara et al. (1982) reported that buprofezin (NNI-750) possessed
good insecticidal activity against BPH nymphs and adults. Mortality generally occurred as a result
of the nymphs being unable to emerge fully from the exuviae during molting (Asai et al., 1983). The
compound at 250 ppm when sprayed reduced the egg hatch from 100% to 43.3% and when the
compound was used at l000 ppm, egg hatch was only 9.7%. It was also known to have sterilizing
effects on the adults.
Hirao et al. (1983) stated that a mixture of 1% buprofezin + 2% BPMC formulated as a
driftless dust, significantly reduced populations of BPH for an extended period if the application
was made immediately after the main wave of immigration into Japan. Also, application of 1.5%
buprofezin as a driftless dust formulation was very effective in limiting the incidence of the grassy
stunt disease transmitted by BPH and stripe disease transmitted by the SBPH.
Third instar nymphs of BPH, GLH (Nephotettix sp.) and WBPH died at molting when they had
been sprayed with 0.075% buprofezin (Valencia et al., 1983). Heinrichs et al. (1984) reported that
buprofezin had a longer residual effect against the major hopper pests of rice than currently used
carbamate insecticides, and was safe to their predators, Lycosa pseudoannulata, Cyrtorhinus
lividipennis and Microvelia atrolineata.
Plant extracts from the roots of Eclipta alba and leaves of Azadirachta indica cause high BPH
mortality (Rao and Rao 1979b). Seed oil of chinaberry, Melia azadirach, at concentrations of
0.01-0.04% had marked antifeedant and some systemic effect against BPH and WBPH (Jianzhang
et al., 1983). Application of a mixture of neem cake and urea at a rate of 120 kg N/ha reduced the
food intake of newly emerged BPH females and also reduced the build up of nymphs and adult
populations to a certain extent (Saxena et al., 1984).
2.1.2 Factors Affecting the Efficacy of Insecticides

Proper placement of foliar sprays is important in obtaining effective control. BPH feed primarily at
the base of rice tillers and hence insecticidal sprays applied above the canopy often provide poor
control of the pest. With a non-systemic insecticide (e.g., ethylan) increased control of BPH was
achieved when the spray was directed to the base of the plant rather than on the canopy (Aquino
and Heinrichs 1978).
High reproductive capacity and oviposition behavior renders the insecticidal control of BPH
and WBPH difficult. The eggs of these pests are inserted into the stem tissues at the base of the
plant, where sprays and dusts do not penetrate. Hence, insecticides with ovicidal action can be used
to kill eggs. Heinrichs and Valencia (1978a and 1981) found that carbofuran, triazophos, methyl
parathion and bufencarb reduced BPH egg hatch significantly. Rao and Rao (1979c) reported that
carbofuran applied on paddy water, and BPMC, carbaryl and isoprocarb (MIPC) applied to water
or soil inhibited BPH egg hatching.
Additional factors that complicate the use of insecticides in the control of BPH are the
following:
• The BPH is a phloem feeder and systemic insecticides move primarily through the xylem.
Insecticide accumulates in the leaf tips and little of it accumulates in the leaf sheath area
where BPH feed.
• It is extremely difficult to control adults migrating from adjacent outbreak areas. Even when
fields are sprayed, the insects are capable of laying eggs before they are killed. Many
insecticides do not kill the eggs and when they hatch, the residual activity of the insecticide
is not sufficient to kill the hatching nymphs and, thus additional applications are required.
Timing of application is critical.
• Because of the high reproductive rate, BPH rapidly develops resistance to insecticides.
• Many insecticides applied at sublethal rates cause BPH resurgence. Because many farmers
use sublethal rates, these insecticides have caused serious problems.
• Physiologically different BPH populations or biotypes have been identified. Biotypes have
differential susceptibility to the commonly used insecticides such as carbofuran, metalkamate,
diazinon and methyl parathion in contact toxicity tests (Heinrick and Valencia 1978b).
2.1.3 Timing of Insecticide Application

Irrespective of chemical type, formulation or application method, careful and appropriate timing of
application is the simplest way to enhance insecticide effectiveness. Kiritani (1972) suggested that
appropriate timing will achieve insecticide specificity. In the case of BPH, where eggs are difficult
to kill, it is best to apply insecticides when the majority of nymphs are in the 3rd or 4th instar stages.
In the People’s Republic of China, three BPH generations occur on the first crop and damage is
caused by the third generation. Most effective control was obtained by treating the second genera-
tion (Anonymous 1977).
Timing of insecticide application should take into consideration the natural enemy population in
the field. Cheng et al. (1973-1975 unpubl.) estimated the rice yield loss caused by insecticide application
on natural enemies. Their results showed more predacious spiders and more egg predation in un-
protected plots than in plots where insecticides were applied as needed, and fewer spiders and less
parasitism in plots where insecticides were applied at 15 day intervals. Heinrichs et al. (1981) suggested
that, based on light trap catches and field surveys, the best time to spray for BPH is about two weeks after
peak trap catches, provided the economic threshold has been reached.
Methyl parathion and deltamethrin caused a population of BPH to resurge when these com-
pounds were applied as sprays 50 or 60 days after transplanting (DT), but earlier applications
caused no resurgence (Heinrichs et al., 1982a).
The use of an economic threshold (ET) is critical to the minimum but effective use of
pesticides. Use of the appropriate insecticide at the economic threshold level (ETL) places pest
control on a sound economic basis with minimum ecosystem disruption. In different countries,
various ETLs have been reported. Dyck and Orlido (1977) suggested an economic injury level of
one BPH/tiller or about 25 BPH/hill as a rough guide for practical BPH control operations
throughout a crop period. In Taiwan, Yen and Chen (1977) showed that 25 BPH/hill was a practical
control threshold, based on insecticide application experiments.
Yokoyama (1979) recommended insecticide application at a population of more than 20
adults/100 hills at about one month after the peak immigration of adults at tillering stage and when
more than 10 adults/100 hills occurred during the post-booting to heading stage. Ooi (1982)
suggested the economic threshold for the BPH to be 7 adults or 15 nymphs per hill in Malaysia.
Sellammal Murugesan and Chelliah (1982b) reported that an average of 2.5 insects/tiller
resulted in severe economic damage in a susceptible variety, TN 1, while even a mean of 2.6
insects/tiller did not cause economic damage in a moderately resistant variety, CO 42. They further
reported that the damage in the resistant variety ASD 11 was low at the above population levels.
Thus it is necessary to develop ETLs depending on the level of resistance a variety possesses.
Sellammal Murugesan and Chelliah (1982a) further observed that in moderately resistant varieties,
CO 42 and Triveni, at the same level of BPH population, the yield loss was relatively lower in
60-day-old plants than that of the 45 day-old plants. Thus it is suggested that the ETL should be
assessed taking also into consideration the level of insect resistance of the variety, stage of the crop,
the natural enemy population and the climatic conditions that prevail in a region.
2.1.4 Insecticide Application Methods

Foliar sprays. Efficient application methods are necessary to make insect control an effective and
economical proposition.
With a non-systematic insecticide (e.g., ethylan), increased control of BPH was achieved when
the spray was directed to the base of the plant rather than on the canopy (Aquino and Heinrichs
1978) (Table 1). Application of high volumes (300-1000 l/ha) of water as required in the commonly
used knapsack sprayer is a drudgery. However, when the canopy has closed, high spray volumes of
1000 1/ha are recommended for BPH control in the Philippines (Mochida and Heinrichs 1980). A
greenhouse test comparing the effect of spray volumes of several insecticides on BPH control
indicated that for most insecticides all the tested spray volumes (i.e., 250, 500 and 1000 l/ha)
provided similar initial control, but the highest spray volume provided the longest residual toxicity
(Fig. 1) (Heinrichs and Valencia 1979).
TABLE1
Placement of foliar sprays in brown planthopper (BPH) control. IRRI, 1977 wet season. (Aquino and Heinrichs 1978)
BPH/4 linear m row sampled with

a D-vac suction machine 2
Treatment 1 Before application 2 days after application Percent Control 3
Applied above the rice canopy
Metalkamate 4,766a 1,183a 74
Monocrotophos 8,213 a 3,168b 60
Ethylan 7,342a 2,589b 63
Control 6,190 a 5,893b 5
Applied below the rice canopy
Metalkamate 11,241a 947a 92
Monocrotophos 6,723 a 2,074 b 69
Ethylan 7,521a 335 a 96
Control 7,805a 8,811c 0
1 Hopper resurgence was induced by spraying twice with 30 g a.i/ha decamethrin, starting 15 days after planting.
Insecticides tested were applied at 750 g a.i/ha at 70 days after planting.
2 In a column, means followed by a common letter are not significantly different at 5% level (DMRT).
3 Values adjusted using Abbott's formula.
Fig. 1. Effect of three spray volumes on mortality of the green leafhopper, Nephotettix virescens and the brown
planthopper, Nilaparvata lugens. Mortality readings taken at 24 hours after caging insects on treated plants.
Other groups of insects were caged at 1, 7 and 14 days after treatment. All insecticide was applied at the
rate of 0.75 kg a.i./ha except for carbofuran which was applied at 0.25 kg a.i./ha. IRRI greenhouse. 1979.
(Heinrichs and Valencia 1979).
Studies at IRRI have indicated that the deposition of insecticide at the BPH feeding site with
an ultra low volume (ULV) applicator is similar to the application with a knapsack sprayer (Pick
et al., 1981). High wind speeds, however, cause drifting of the small droplets emitted by the ULV
sprayer. Electrostatic sprayers that add an electrical charge to the droplets, that is opposite to the
charge of the rice plant, show promise in overcoming the drift problem because the droplets are
attracted to the plant. Insecticides applied as foliar sprays, when exposed in the field, have
extremely short residual activity. Micro-encapsulation of insecticides provides a means of slow
release, thus increasing residual activity and also decreasing the hazard to the applicator (Mochida
and Heinrichs 1980).
Dusts: Dust formulations are easy to apply. However, because dusts act mainly through contact
toxicity, the natural enemy population is likely to be affected. Mixtures of insecticidal dusts
containing 2% MTMC and 2% phenthoate applied at 33.6 kg/ha from a knapsack duster fitted with
a long vinyl tube with regularly spaced openings, caused heavy mortality of BPH nymphs. However,
predators were adversely affected by the treatment (Lim et al., 1978).
Paddy Water Application of Granules: The use of a granular formulation, that can easily and
safely be broadcast by hand, eliminates many of the application problems encountered with foliar
sprays. Granules, however, are expensive and the number of effective insecticides in granular
formulation is limited. Carbofuran and diazinon granules have been commonly used for BPH
control.
Root-zone Application. Root-zone application provides much longer residual activity than foliar
sprays and broadcast of granules (Fig. 2). Root-zone application is more effective against the GLH,
Nephotettix virescens, than against the BPH. Another disadvantage of the root-zone method is that
application is most convenient at or shortly after transplanting and the BPH does not infest the field
until about 30 days after transplanting when the residual activity of the insecticide has substantially
decreased.
Fig. 2. Field evaluation of three methods of isazophos application at 0.5 kg a.i./ha. In the root-zone treatment,
insecticide was placed in a gelatin capsule and inserted into the soil near the root-zone and in the broadcast
method granules were broadcast into the paddy water. Mortality readings were taken 48 hours after caging
hoppers on treated plants and adjusted using Abbott’s formula. Variety IR22. IRRI, 1979. (Heinrichs and
Arceo 1979).
2.1.5 Selective Uses of Insecticides

Populations of planthoppers, as well as the natural enemies, are to be taken into consideration for
effective and judicious use of insecticides. Insecticide selectivity results from the intrinsic toxicity of
the materials and from various operational factors. Insecticides moderately toxic to BPH predators
and at the same time very effective against the BPH have been reported (IRRI 1977). Negligible
toxicity to the predatory spider Lycosa pseudoannulata was observed when applying insecticides
through improved application techniques (Heinrichs et al., 1979a). Such techniques also made it
possible to culture fish in rice fields treated with insecticides (Heinrichs et al., 1978a). Chang et al.
(1979) reported that two insecticides, isoprocarb and pyridafenthion, have the desired selective
toxicity for BPH and relative safety for the spider, Pirata subpiraticus.
Root-zone application of carbofuran at 1.5 kg ai/ha effectively controlled GLH and BPH.
Granules were effective against BPH for 30 DT without adversely affecting the stem borer egg-lar-
val parasite, Platygaster sp. (Chiu et al., 1980). Root-zone application of carbofuran and BPMC had
an inhibitory effect on hatching of BPH eggs (Rao and Rao 1980a).
2.1.6 Resistance to Insecticides

Continuous use of insecticides has resulted in BPH resistance to insecticides in Taiwan (Lin et al.,
1979), Japan (Nagata 1979) and the Philippines (Heinrichs 1979). Resistance to insecticides in
WBPH and SBPH has also been reported (Nagata 1979). BPH resistance to HCH dusts was
reported in Japan as early as 1967 (Nagata and Moriya 1969). Topical doses of HCH in the
laboratory increased resistance sevenfold within five generations. Studies by Ozaki indicated that
BPH had developed resistance to fenthion, fenitrothion, diazinon and malathion (Heinrichs 1979).
At IRRI, the effectiveness of diazinon markedly declined in 1969 (IRRI 1970) after it had been
used continuously for ten successive crop seasons. Studies indicated that the loss of effectiveness
was due to increase in resistance to the insecticide and to microbial degradation (IRRI 1971).
After several years of use, carbofuran’s effectiveness against BPH declined at IRRI. Applica-
tion of 2 kg ai/ha at 14 day intervals failed to control the pest. The responses of field-collected
insects to paddy water and to root-zone application of carbofuran at 1 kg ai/ha were compared with
the response of a susceptible greenhouse culture. The field collected hoppers showed an extremely
low mortality thus indicating that they were resistant compared to greenhouse cultured insects
(Heinrichs 1979). Contact toxicity tests also indicated that the field population was distinctly more
resistant to carbofuran than the susceptible greenhouse culture (Heinrichs and Valencia 1978c) and
it was reported that the field strain was seven times as resistant as the greenhouse strain.
The BPH was reported to have developed resistance to various other insecticides (i.e.,
monocrotophos and isoprocarb) (Heinrichs and Tetangco 1978). Chung et al. (1982) reported that
by continuous selection of a field strain, BPH developed 1183-fold resistance to malathion in 9
generations and 41-fold resistance to isoprocarb in 16 generations, and both of these selected
strains were equally resistant to propoxur and permethrin but remained susceptible to fenvalerate.
Ozaki and Kassai (1982) found that under field conditions, BPH developed resistance to fenthion
(30- to 32-fold), fenitrothion (22- to 23-fold), cyanofenphos (145- 423-fold) and malathion (20- to
31-fold) in a period of 6-7 years. Development of carbamate-resistance was also reported but
occurred at a slower rate.
666 INSECTICIDE MANAGEMENT IN R I C E
Kassai and Ozaki (1984) reported that a malathion-resistant strain developed 5- to 26-fold cross
resistance to naled, tetrachlorvinphos, monocrotophos, propaphos, fenthion, fenitrothion,
diazinon, isoxathion, pyridafenthion and carbaryl. They further stated that a fenitrothion-resistant
strain was found to have developed various increased levels of cross resistance to the above
insecticides and also malathion. Sun and Dai (1984) observed that BPH developed 1000-fold
resistance in Taiwan to permethrin while the resistance levels to other pyrethroids (i.e. cyper-
methrin, deltamethrin, fenvalerate) were much lower, ranging from 10- to 50-fold.
Development of insecticide resistance has also been reported in case of SBPH to malathion in
Korea (Choi et al., 1975) and Japan (Nagata and Mochida 1984), and in WBPH in Sri Lanka
(Wickremasinghe and Elikawela 1982) and in Japan (Nagata 1979).
2.1.7 Insecticide-Induced Brown Planthopper Resurgence
A major factor that has contributed to the number and severity of BPH outbreaks in the last two
decades has been widespread and indiscriminate use of insecticides in the rice ecosystem. The
degree of resurgence is dependent on an interaction of factors consisting of the insecticide and its
effects on the rice plant, the BPH populations, and the natural enemies of the BPH (Heinrichs and
Mochida 1984) (Fig. 3).
Fig. 3. Diagrammatic explanation of how insecticide management determines the degree of brown planthopper,
N. lugens resurgence through a direct effect on the BPH populations and via the host plant and natural
enemies of the BPH.
Numerous BPH resurgence-inducing insecticides have been identified in field tests at IRRI
(Heinrichs et al., 1982b; Reissig et al., 1982a, b) and in national programs. Studies have shown that
resurgence-inducing insecticides include representatives of the phosphate, carbamate, and
pyrethroid classes (Heinrichs and Mochida 1984). Insecticide type and rate, and the number and
method of insecticide applications all influence the degree of BPH resurgence (Fig. 3). Factors
contributing to BPH resurgence consist of the direct effects of the insecticide on the BPH popula-
tion and the indirect effects on the BPH population via the host plant and natural enemies.
The success of the ‘green revolution’ in rice in Asia has been limited by the problems caused by
insecticides that induce the resurgence of BPH populations. Because of the severe economic effect
of BPH resurgence, it is important that rational insecticide management programs be established
and integrated with other BPH control tactics in an integrated pest management approach.
2.2 GREEN LEAFHOPPER SPECIES

There are three important green leafhopper species; Nephotettix virescens, N. nigropictus and N.
cincticeps. Because these species are vectors of rice virus diseases, insecticides should have imme-
diate knockdown effects.

Insecticides are not necessary unless the virus diseases transmitted by GLH are prevalent. In India,
10 GLH/4 net strokes or 5/hill at the vegetative stage and 10/hill at the post flowering stage were
listed as thresholds (Heinrichs et al. 1979b). A threshold of 2 GLH/hill has been suggested in tungro
endemic areas, in Tamil Nadu, India (Anonymous 1985).
22.2 Insecticide Application Methods

A liquid band applicator developed at IRRI facilitated root-zone application and provided effective
control of early season pests such as whorl maggot and GLH (Heinrichs et al., 1979b). The
incorporation of carbofuran granules into the root-zone at the time rice is planted controlled tungro
virus disease by preventing feeding by the vector N. virescens (Basilio and Heinrichs 1981). Root-
zone application was more effective for the control of GLH than BPH (Mochida and Heinrichs
1980).
Seedling root coating and root soaking with carbofuran just prior to planting gave 100% GLH
mortality even at 50 DT (Seiber et al., 1977). They also reported that the root-coat treatment
technique provided excellent control of tungro virus incidence in the field up to 40 DT.
2.23 Resistance to Insecticides

Insecticide resistance in GLH is a serious problem in Japan (Nagata and Mochida 1984). Malathion
resistance in N. cincticeps was reported in 1960-61 and methyl parathion resistance in 1962.
‘Malathion and parathion were replaced by carbamate insecticides.
Ku and Wang (1978) observed development of insecticide resistance in N. cincticeps to the

carbamates BPMC, carbaryl, CPMC, carbofuran and isoprocarb. Fabellar et al. (1981) reported low
levels of insecticide resistance in the GLH to acephate and methyl parathion in the Philippines.
Resistance-ratios ranging from 27 to 451 for malathion, 32 to 875 for methyl parathion and 12 to 79
for carbaryl have been reported in the GLH population in Taiwan (Kao et a1.,1981). Resistance-
ratios for the same populations were found to be 1-55 for permethrin and 1-2.4 for fenvalerate.
Miyata et al. (1981) reported that the major factors involved in resistance to malathion in the
GLH were increased malathion-degrading carboxylesterase activity and reduced sensitivity of
AchE to malaoxon. Glutathione -S transferase was thought to be involved in resistance to
fenitrothion and for the carbamates the main resistance factor was reduced sensitivity of AchE
rather than carbamate metabolism. The major resistance mechanism in the GLH to malathion was
enhanced degradation by carboxylesterases while for BPMC, it was reduced sensitivity of AchE
(Saito and Miyata 1982).
The most common method of managing insecticide resistant insect populations is to use new
insecticides with no cross resistance to the old insecticide. Malathion and parathion were first used
for GLH control, then replaced with carbamates. Carbamates were again replaced with OP
chemicals such as diazinon and propaphos to which malathion- or parathion-resistant GLH showed
no cross resistance. Mixed formulations of organophosphates and carbamates also have been used
to overcome carbamate resistance (Hama and Iwata 1973). All of these insecticides are losing effect
because GLH is developing multiple resistance. Few promising chemicals are now available to
destroy resistant GLH populations except some pyrethroids.
2.3 STEM BORERS

Important species of stem borers of rice are the yellow stem borer (YSB), Scirpophaga incertulas,
and the striped stem borer (SSB), Chilo suppressalis, among which the former is a major pest of rice
in South Asia and the latter is important in Southeast Asia.

The yellow stem borer lays eggs covered in a hairy matrix that prevents insecticide penetration and
hence the ovicidal action of the toxicant is nullified. After hatching, the larvae bore into the stem
rendering the action of contact insecticides ineffective. Hence, systemic insecticides that move
through the plant system are most effective against the borers.

Careful timing of insecticide application can significantly reduce the amount of material used and
increase selectivity. In China, foliar sprays for the control of YSB were more effective during egg
hatch than during peak adult emergence (Chiu 1979). In areas where stem borers occur in distinct
broods, application of insecticides in relation to brood emergence was reported to be more effective
than calendar based schedules (Heinrichs et al., 1979b). The insecticides can be applied when the
stem borer damage reaches the ETL. An ETL of 5-10% deadhearts has been suggested for stem
borers (Heinrichs et al. 1979b).
2.3.3 Selective Uses of Insecticides

Insecticides that are safe to natural enemies may lead to an effective and durable suppression of the
pest. Spraying the rice crop with metalkamate did not hamper parasitization of the stem borer egg
masses (IRRI 1976). Application of carbofuran granules was very effective in controlling the stem
borer and sparing hymenopterous parasites, but it severely reduced the population of predatory
spiders and other predators (e.g. Cyrtorhinus lividipennis and damselflies) (Khusakul et al., 1979).
Miyata and Saito (1982), reported that malathion and pyridafenthion were less toxic to predators of
Chilo suppressalis (i.e., Paradosa astrigera, Conocephalus maculatus) because of low cuticular
penetration.
2.3.4 Resistance to Insecticides

Stem borers have developed resistance to insecticides. Resistance of the striped stem borer to BHC
and parathion in Japan has been reported (Nagata and Mochida 1984).
2.4 DEFOLIATORS
Major defoliator pests of rice include the leaffolder, Cnaphalocrocis medinalis, rice caseworm,
Nymphula depunctalis and rice armyworms, Spodoptera spp.

The larvae of leaffolders remain in leaf folds and feed on the tissue by scraping. Hence, insecticides
applied on the foliage often do not contact the larvae. Along with C. medinalis, other species of
leaffolders (i.e, Marasmia patnalis and M. ruralis) have been reported to occur in rice fields in
Madurai (Gunathilagaraj and Gopalan 1986) and at Aduthurai, Tamil Nadu, India (Personal
unpublished data). Because of the mixed occurrence of these leaffolder species, insecticides should
be screened against each species in determining their efficacy and making recommendations.
Caseworm larvae shelter inside leaf cases into which insecticide penetration is limited. The
insect migrates by falling on to the water surface and moving along with irrigation water. Thus
insecticide applied on the foliage has limited influence on caseworm larvae.
The armyworm feeds on foliage during night but hides in soil cracks and crevices in the day
time. Mature larvae are less susceptible to insecticides than early instars thus necesitating insec-
ticide application when the insects are in early instar stages.

The economic threshold for the leaffolder was reported in India as 10-15% damaged leaves. In
China, 10 eggs/100 plants or 20 damaged leaves per 100 plants was considered as the threshold
(Heinrichs et al. 1979b). Threshold levels of 10% damaged leaves in the vegetative phase and 5% of
flag leaf damage at the flowering stage have been reported in Tamil Nadu, India (Anonymous 1985).
Feeding by leaffolder and armyworm is extensive during night and hence, to achieve better control,
insecticides should be sprayed during the evening hours.

To control leaffolder larvae that are present inside the leaf folds, insecticide should be applied in
such a way that the entire foliage is covered by the insecticide spray. For armyworm control,
insecticide should be applied on to the base of the rice plant and soil crevices where the larvae
remain during day time. Heavy damage by leaffolder is often seen in patches in shady areas,
particularly under trees. Insecticides can be spot applied in these patches and thus further spread-
ing of damage can be avoided.
2.5 GALL MIDGE

The rice gall midge, Orseolia oryzae, is one of the major rice pests in India, Thailand and Sri Lanka.
Maggots of the gall midge feed on the shoot apex resulting in the suppression of the apical meristem
and formation of radial ridges from the inner most of the leaf primordium which is followed by the
elongation of the leaf sheath leading to the formation of an onion leaf-like structure.

The presence of the gall midge is only evident after damage has been done. Insecticides applied
after the appearance of the gall-like symptoms are evident will only check further incidence of the
damage. Because the gall midge maggot remains inside the gall, insecticides having contact action
have little effect on the maggots. Systemic insecticides are most effective.
2.5.2 Timing of Insecticide Applications

Maximum damage by this pest is caused up to 60 days after transplanting and hence, chemical
control should be applied at an early growth stage. Insecticide timing can be determined as based
on catches of adults in a light trap. Insecticide should be applied a few days after the peak periods
of adult catches in the trap. In areas where the gall midge is a chronic pest, applications in the
nursery and applications at transplanting may be warranted.

Granular insecticides applied on the water surface and seedling root dip in insecticidal solution are
effective application methods for the control of the gall midge. Root dipping in isofenphos 0.02%
solution for 12 h yielded relatively better control than dipping in 1 h (Misra et al. 1981). Samalo et
al. (1983) reported that seedling root dip for 12 h with 0.02% chlorpyriphos or isofenphos followed
by one application of ethoprophos granules at 1 kg ai/ha after 30 DT gave good control and allowed
a fairly high level of parasitization by Platygaster oryzae.
2.6 RICE BUGS ON PANICLES

The rice bugs Leptocorisa spp. cause damage by feeding on the sap of the milky grains and turn them
chaffy. In controlling this pest, care should be taken that the insecticide residues do not remain on
the grains after harvest.

An economic threshold of 5 bugs/100 panicles at flowering and 16 bugs/100 panicles at the milky
stage to grain maturity is recommended in Tamil Nadu, India (Anonymous 1985). Insecticides
should be applied when the bugs are in their early instars as they are more susceptible than later
instars.
3. ENVIRONMENTAL IMPACT OF INSECTICIDE USE IN RICE
3.1 EFFECT ON BENEFICIAL INSECTS

Some insecticides used in rice are more toxic to beneficial insects (e.g., natural enemies and
pollinators), fish species, and other non-target organisms than to the targeted pest species. Adverse
effects of insecticides on beneficial insects are due to various factors such as the wrong choice of
insecticides that are inherently more toxic to natural enemies than to pest insects, improper method
of insecticide application and incorrect timing of application.
Choi et al. (1978) observed that among three methods of carbofuran application (root-zone
application with capsules, soil injection as a liquid formulation and 2-4 broadcasts in the irrigation
water) for control of leaf- and planthoppers, the predaceous spider population was reduced in all
the three methods of application, but the reduction was greatest in the capsule placement.
Choudhury and Alam (1979) found that 0.2% diazinon spray controlled rice pests effectively
but was very toxic to the mirid predator Cyrtorhinus lividipennis and the predatory spiders. The dust
formulations of propoxur and MTMC were toxic to C. lividipennis (Ooi 1979). Carbofuran was very
effective in controlling the stem borer and spared the hymenopteran parasites. However, it severely
reduced populations of predatory spiders (eg., Tetragnatha sp., Oxyopes sp., Lycosa pseudoan-
nulata), C. lividipennis, and damselflies (Khusakul et al., 1979). The insecticide pyrimioxythion was
reported to be highly toxic to the parasite of rice gall midge Platygaster oryzae, whereas chlor-
dimeform was less toxic. Joshi and Venugopal (1983) indicated endosulfan spray to be very toxic to
the gall midge parasite, P. oryzae, while phosalone was relatively safe.
Fabellar and Heinrichs (1984) reported that cypermethrin and deltamethrin were highly toxic
to BPH predators Microvelia atrolineata, Lycosa pseudoannulata and C. lividipennis. Chelliah and
Rajendran (1984) found that endosulfan was least toxic to C. lividipennis while quinalphos, chlor-
pyriphos, phosalone, and monocrotophos were highly toxic.
Aguda et al. (1984) observed that the insecticides causing BPH resurgence (i.e., methyl
parathion and azinphos ethyl) greatly reduced spore germination of the entomogenous fungi
Metarhizium anisopliae and Beauveria bassiana which commonly attack BPH. Hence, it is advisable
to use whenever possible insecticides that are selective, effective, and safe to insect pathogens,
parasites and predators.
3.2 INSECTICIDE RESIDUES AND TOLERANCE LIMITS

Insecticides used for rice pest control may remain in the environment as residues in food, cattle
feed, water, or in the atmosphere and thus, pose multivarious health hazards, both long and short
term. Although few studies have been conducted to determine residues of insecticides remaining in
rice plants and the rice ecosystem after insecticide application, residues do not appear to be a major
problem.
Teimoory and Shekarabi (1979) estimated the residues of diazinon 10% granules applied at 20
kg/ha. The residues in water ranged from 0.062 ppm to 0 ppm from the day of application to the 10th
day after treatment, respectively. When phosphamidon was applied at 0.75 l/ha, residues in the field
water ranged from 0.114 pprn to 0 ppm and that of lindane from 0.79 to 1.05 ppm from 0 to 10th day
after treatment, respectively.
Out of five treatments with carbofuran only the high rate of 6 broadcast applications of 1 kg
ai/ha at 14-day intervals caused grain residues above the allowable level. The U. S. environmental
Protection Agency (EPA) has set a tolerance limit of 0.20 ppm in grain for carbofuran plus its
metabolite 3 hydroxy carbofuran (Heinrichs et al., 1978b). Studies conducted by Rao and Rao
(1980b) showed that residues of carbofuran were highest in grains, straw and soil when three
applications were made at 10, 40 and 70 DT but were below the tolerance limit fixed by the EPA.
Lindane at 0.75 kg ai/ha was found to be effective against the rice bug, Leptocorisa oratorius.
Residues in the harvested grains were well below the tolerance limit (Argente and Heinrichs 1983).
It is necessary to analyze for residue levels for various insecticides at various rates of usage and
methods of application and to establish the tolerance limits to avoid/minimize the toxic hazards of
chemical pesticides.
3.3 HUMAN HAZARDS

Hazards to humans by insecticide use occur in two ways; (1) direct hazards that occur during
handling and application and poisoning due to internal consumption and (2) indirect hazards of
pesticide residues in the usable products and the environment.
Granular insecticides (e.g., carbofuran and phorate) have high dermal toxicity and cause skin
irritation and allergenic symptoms. Some of the sprayable insecticides (eg., ethyl parathion) are
extremely toxic to humans during spray operations.
Although few chlorinated hydrocarbon insecticides are now used, it is important to mention
that some of them have high persistence in the environment and cause chronic toxicity. They
accumulate in the fat tissue of animals and humans and are found in milk and dairy products.
Organochlorines can be transferred from mother to fetus and consequently, babies may be born
with pesticide residues in their bodies. Milk from such mothers will also contain chemicals. These
pesticides accumulate in smaller quantities in organisms at lower trophic levels and because humans
are at higher trophic levels, accumulation of these insecticides in human bodies increases at an
exponential rate.
4. INTEGRATION OF INSECTICIDES WITH

OTHER CONTROL METHODS
In rice IPM, insecticides can be integrated with biological control, resistant cultivars and cultural
controls.
4.1 INSECTICIDES AND BIOLOGICAL CONTROL

To date, successes of classical biological control have been few. Numerous failures have been
reported (Nickel 1967, Yasumatsu and Torri 1968, Lim 1974). However, the use of biocontrol
methods, when integrated with insecticides, can be highly effective. In South China, Trichogramma
parasites are being mass reared and released to control leaffolders. Insecticides are used only when
necessary and usually for controlling other pests such as stem borers, thrips and planthoppers (Chiu
1980). Bacillus thuringiensis has also been integrated with low dosages of insecticides (0.1-0.2 of the
normal strength) to provide effective control of lepidopterous pests in Kwangtung Province (Chiu
1980).
4.2 INSECTICIDES AND RESISTANT CULTIVARS

Insecticides are highly complementary to resistant varieties. Often, rice varieties are resistant to a
few insect species and need insecticides for control of other pests. In the Philippines, IR 26 was
resistant to BPH, but insecticide protection was necessary against other insects (Heinrichs et al.,
1979b).
On moderately resistant varieties, developing insects may be less vigorous, smaller and less
fecund and may succumb more readily to insecticidal treatments. For example, stem borer infesta-
tions on moderately resistant varieties receiving only one or two insecticidal treatments were similar
to or lower than those observed on susceptible varieties receiving twice the number of insecticide
treatments (Saxena 1980).
Biotype selection is a major problem in the use of resistant varieties, especially in BPH control.
However, Panda and Heinrichs (1982) have identified tolerant rice varieties with no major genes for
BPH resistance (IR 46, Kencana, Triveni and Utri Rajapan). Though only moderately resistance, they are
believed to be stable to BPH stress. Such tolerant varieties, when supplemented with insecticides on a
need basis, can be expected to produce stable yield and a profitable economic return.
4.3 INSECTICIDES AND CULTURAL CONTROL

Close planting in rice not only contributes to rapid increases of pests such as BPH (Oka 1979), but
it also prevents foliar sprays from reaching target pests under the canopy. However, wide spacing
~~~ ~~ ~ ~~~ ~ ~
can significantly reduce yields. If IPM programs are to be successful, appropriate integration of
good cultural practices with control methods such as insecticides are necessary (Lim and Heong
1984).
Cultural practices along with insecticides can play a significant role in a complementary way.
The spraying of susceptible trap crops has proved successful (IRRI 1978). Because a cultural
practice may have differential effects on various pests (Kiritani 1972, Lim and Heong 1976),
insecticides could be integrated into such circumstances. The cultural practice could be used to
control key pests while chemicals are used to control less important species.
5. CONCLUSION
Insecticides will continue to serve as an essential component in the management of rice insects for
several years. However, their use should be based on sound economical principles. It is justifiable
to use insecticides only when benefit-risk ratios are in favor of insecticide use. Development of
insecticide resistant populations, resurgence and adverse effects on non-target organisms are to be
understood in greater depth in order to use insecticides most effectively in rice insect control. Thus,
the insecticides are double-edged swords and using them in an unscientific manner may bring
catastrophic results.
Worldwide regulations on the release of new insecticides are becoming increasingly more
stringent. It is thus obvious that we cannot afford to lose the currently available compounds. It is
imperative therefore, to develop appropriate insecticide management strategies to extend the
commercial life of the existing insecticides, as well as to protect human health and the environment.
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Pillai, K. S., K. Saradamma and N. M. Das. 1983. Field evaluation of two newer insecticides against some rice pests in
Kerala. Pesricides 17(2): 15-18.
Rao, P. R. M. and P. S. P. Rao. 1979a. Evaluation of systemic effect of insecticides against brown planthopper. Intl. Rice
Res. Newsl. 4(4): 16.
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and Azadirachta indica A. Juss. for the control of rice brown planthopper. Ind J. Agric. Sci. 49(11): 905-906.
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Noyaku. 26(3): 3-15.
10
Integration of Rice Insect Control

Strategies and Tactics
C. MICHAEL SMITH
Department of Plant, Soil and Entomological Science
University of Idaho
Moscow, Idaho
1. INTRODUCTION
DURING THE PAST THREE decades, significant shifts in the insect pest complexes of Asian rice
producing nations have occurred. Agricultural scientists in these nations have gained the
knowledge necessary to manage these insects, and have synthesized various control tactics that are
now in use in the form of integrated pest management (IPM) programs. These programs are usually
carried out through the efforts of each nation’s national program staff for agriculture. Examples of
such program activities exist in India, Indonesia, Japan, Malaysia and the Philippines. In these
countries, IPM programs utilize traditional chemical, biological and cultural control measures to
varying degrees, depending on the nature of the pest insect complex and the socioeconomic
conditions existing in the areas where IPM is carried out. In the Peoples’ Republic of China, IPM
programs are quite comprehensive, utilizing biological, chemical and cultural controls linked to
extensive surveillance programs.
One form of cultural control, the development and cultivation of insect resistant rice cultivars,
has been tremendously successful, due to the cooperative efforts of rice researchers throughout the
world. More than 20 million ha of rice in Asia are now planted to cultivars with resistance to the
brown planthopper, Nilaparvata lugens Stal, and an additional 10 million ha of cultivars with
resistance to various other insect pests are planted throughout the world (IRRI 1982). Sources of
682 INTEGRATION OF RICE INSECT CONTROL STRATEGIES
resistance to several of the major rice diseases and insects have been identified, cataloged, and are
currently in various stages of varietal development (Khush 1980,1984).
The purpose of this chapter is to examine instances where the different types of IPM tactics
have been successfully integrated and to suggest requirements for the continued development of
IPM programs for control of rice insects.
2. CULTIVAR RESISTANCE AND CHEMICAL CONTROL

With the advent of the Green Revolution, insect pest problems in rice have generally intensified,
and as a result, insecticide use patterns have increased. In some instances, insecticides significantly
enhance the yield of rice cultivars with insect resistance (Heinrichs et al., 1986b). Heinrichs et al.
(1979) demonstrated that carbofuran treatment of IR26, a cultivar with brown planthopper biotype
1 resistance, gave control of the rice whorl maggot, Hydrellia philippina, and increased yields by
15%. Kulshreshtha (1979) demonstrated 35-71% yield increase in IR 26 rice after insecticidal
treatment for the yellow stem borer, Scirpophaga incertulas (Walker), in India. Similar results were
reported by Heinrichs et al. (1984,1986b) with insecticides applied to moderately resistant cultivars
for control of the whitebacked planthopper, Sogatella furcifera (Horvath), and the green leafhop-
per, Nephotettix virescens (Distant).
However, insecticides may be of no practical value in the cultivation of some resistant cultivars.
Research in India (Prakasa Rao 1974) and Thailand (Tirawat 1975), indicates that cultivars with
high levels of resistance to the gall midge, Orseolia oryzae (Wood-Mason), received no benefit from
insecticide treatment. Kalode (1980) indicated that cultivars with O. oryzae resistance in India
yielded similarly, regardless of whether or not insecticides were applied prophylactically, as deter-
mined by surveillance, or not at all (Table 1). Studies by Reissig et al. (1981) in the Philippines with
N. lugens and N. virescens also indicate that insecticide treatments of IR36, a multiple insect and
disease resistant cultivar, do not increase rice yields or farmers' net profits.
TABLE 1
Control of O. oryzae and yields of Shakti (resistant) and Jaya (susceptible) rice cultivars under different
pest management practices. Sambalpur, Cuttack and Bhubaneswar, India 1977-1978
Cultivar
Shakti (R) Jaya (S)
Insecticide treatment Silver shoots Yield Silver shoots Yield
method (%) (t/ha) (%) (t/ha)
Surveillance-based 2.3 3.5 12.7 2.2
Prophylactic 1.0 3.3 9.8 2.3
None 2.7 3.3 21.7 1.8
1 From Kalode (1980).
Increased insecticide use has also caused increased misuse, either from improper insecticide
application, formulation, timing or rate. In rice production, insecticides can have several negative
effects. Insecticides may kill beneficial insects, causing resurgence of pest insect populations, may
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cause the development of insecticide-resistant pest populations, or kill non-target vertebrates

(Litsinger 1983). Reissig et al. (1982) demonstrated that applications of insecticides for outbreaks
of non-target insect pests on rice cultivars with resistance to a key insect pest cause resurgence of
the key pest. This practice also creates a greater possibility of key pest biotype development and
destroys beneficial predators and parasites.
Insecticides also induce changes in pest insect behavior that alter the levels of cultivar resis-
tance of pests. Salim and Heinrichs (1987) demonstrated that more S. furcifera adults are attracted
to plants of resistant, moderately resistant, and susceptible rice cultivars treated with deltamethrin
insecticide than to untreated plants (Fig. 1). As a result, S. furcifera nymphal growth, survival, and
population development were all greater on treated resistant plants than on untreated resistant
plants.
Fig. 1. Effect of deltamethrin insecticide-treated (I) and untreated (C) rice cultivars (R = resistant, MR =
moderately resistant, S = susceptible) on Sogatella furcifera behavior (from Salim and Heinrichs 1987).
The rice cultivars described above have multiple component resistance based on two or more
monogenic sources. The ease with which biotypes of N. lugens have developed virulence to
monogenic resistant cultivars has shown the need for the development of more moderately resistant
cultivars with a broader genetic base of resistance. Recent research at the International Rice
Research Institute (Heinrichs et al., 1984) has shown that N. lugens reared on cultivars with only
moderate resistance suffer higher mortality when treated with insecticides than those reared on
susceptible cultivars (Table 2). Mortality is not different between hoppers reared on resistant and
moderately resistant cultivars when treated with insecticides. Thus, the use of moderate resistance
is of value in reducing the degree of selection pressure causing biotype development.
TABLE2
Insecticide-induced mortality of biotype 2 N. lugen reared on moderately resistant, and
susceptible, rice cultivars, IRRI, 19801
Mean mortality(%)2
Insecticide TNl ASD7 Sinna Sivappu
(S) (MR) (R)
Carbofuran 12F 64b 94a 95a
Carbosulfan 20EC 66b 93a 94a
BPMC 50EC 46b 79a 90a
Control 10b 18a 23a
1From Heinrichs et al. 1984.
224 h mortality; means in a row with the same letter are not significantly different at the P = 0.05 level by Duncan's
multiple range test.
3. CULTIVAR RESISTANCE AND BIOLOGICAL CONTROL

Complexes of biological control agents have been identified for almost all major insect pests of rice
(Chiu 1979). Many of the these predators, parasites and pathogens work effectively to regulate rice
insect pest populations (Kenmore 1980). Because insecticides have detrimental effects on predators
and parasites, IPM research is beginning to determine the interaction of cultivar resistance with
biological control. Early research (IRRI 1975) demonstrated that predation of N. lugens by
predators on hopper-resistant cultivars gave better N. lugens control than either the predator or the
resistant cultivars alone. The effects of moderately resistant cultivars combined with those of the
spider predator Lycosa pseudoannulata (Boes et al. Str.) and the mirid bug, Cyrtorhinus lividipennis
Reuter keep N. virescens (Fig. 2), N. lugens (Fig. 3), and S. furcifera populations below economic
threshold levels (Myint et al., 1986, Kartohardjono and Heinrichs 1984, Salim and Heinrichs 1986).
These results demonstrate how two major IPM tactics can be combined for more efficient insect
control in rice.
4. CULTIVAR RESISTANCE AND CULTURAL CONTROL

Varying the cultural practices in a rice production system is a management tactic that was first used
by Chinese farmers, over 2,000 years ago, when adjusted planting dates and crop stubble burning
were used to reduce the buildup of insect pest populations (Flint and van den Bosch 1981). There
are few documented examples of the integration of cultural practices with plant resistance, but most
cultural practices integrate well in IPM systems.
4.1 PEST AVOIDANCE

Planting the rice crop in a manner that avoids the peak pest insect population generally gives some
degree of pest control. Heinrichs et al. (1986a) found that N. lugens populations and hopper-
predator ratios on early-maturing rice cultivars were significantly lower than those on later (mid-
SMITH 685
Fig. 2. Mortality of Nephotettix virescens due to rice cultivar (R = resistant, S = susceptible) and predation by Cyrtorhinus
lividipennis (minds) and Lycosa pseudoannulata (spiders) (from Myint et al., 1986).
Fig. 3. Mortality of Nilaparvata lugens due to rice cultivar (R = resistant, S = susceptible) and predation by
Cyrtorhinus lividipennis (minds) and Lycosa pseudoannulata (spiders) (from Kartohardjono and Heinrichs
1984).
season) maturing cultivars. These results indicate that the incorporation of N. lugens resistance into
early maturing cultivars would serve to increase rice crop protection. The same relationship exists
with cultivars resistant to O. oryzae. Sen et al. (1984) determined that cultivars with a maturity of
medium duration had less O. oryzae damage than cultivars with maturities of long duration.
Oka (1983) suggested that the incorporation of cultivars into a synchronized rice planting
scheme would further reduce the buildup of economically damaging N. lugens populations. Planting
of a rapid-maturing, secondary (legume) crop between the two rice crops was suggested as an
additional means of disrupting and suppressing N. lugens populations (Fig. 4). The planting of
Fig. 4. Current (upper) and suggested (lower) irrigated rice cropping systems using resistant (R) and susceptible
(S) rice cultivars and second crops to reduce Nilaparvata lugens populations (from Oka 1983).
SMITH 687
insect-resistant rice cultivars was further suggested as a means of reducing pest insect incidence in
this cropping system.
Delaying rice planting until after emergence of the first generation of S. incertulas allows growth
of rice at a time of low borer incidence. In Pakistan, farmers are prohibited by law from planting
until after emergence of first generation moths (FAO 1979). Shifting planting dates has also helped
minimize damage from various borer species in India, Indonesia and Malaysia (Khan 1967). In
Japan, however, the striped rice borer, Chilo suppressalis (Walker), has become adapted to a
modified schedule of late planting (FAO 1982). This insect-crop interaction indicates the need for
close monitoring of insect populations even when planting dates are changed to avoid damage. If
changes in insect emergence do not occur, populations may be further depressed if resistant
cultivars are used in delayed, synchronous plantings.
4.2 FERTILIZER MANAGEMENT

Populations of rice insect pests increase with increased rates of nitrogen fertilization, regardless of
the level of and type of insect resistance (Heinrichs and Medrano 1985). However, the use of
increased amounts of nitrogen has been one of the major factors contributing to the success of
high-yielding rice cultivars. Reissig et al. (1985) state that split applications of nitrogen are more
beneficial than single nitrogen applications. The first application is used more efficiently by the rice
plant and keeps weed growth to a minimum until a second application is made. A reduction in the
amount of nitrogen applied at each application also prevents the buildup of insect populations
during rice vegetative growth, a factor that will synergize the performance of moderately resistant
cultivars (Reissig et al. 1985). These findings indicate that combining insect resistance with fertilizer
responsiveness is best accomplished by selecting cultivars with moderate levels of insect resistance
that are fertilizer responsive.
43 SANITATION
Removal of rice stubble and weeds from rice fields and surrounding areas are practices of crop
sanitation which can work synergistically with the planting of insect resistant cultivars. Cutting,
burning, and plowing rice stubble during the post-harvest period are methods which have been
shown to provide varying degrees of control (60-80%) of larvae of the white stem borer, Scirpophaga
innotata (Walker) and C. suppressalis (FAO 1979, Grist 1953, Khan 1967, Shin-Foon 1980). All
methods of stubble removal are labor intensive, however. A detrimental side effect of both stubble
destruction and weed removal from in and around fields is the destruction of natural enemies and
their habitats (Oka 1979). Though sanitation and insect resistance may be effective in pest popula-
tion reduction, field sanitation as an IPM tactic requires further study to determine if benefits
outweigh detriments.
4.4 TRAP CROPS

Planting a small portion of the rice field earlier than normal to an insect-susceptible cultivar to trap
incoming pest populations is a practice that has potential for integration with planting of resistant
cultivars. Saxena (1982) demonstrated that fields with a trap crop planted 20 days ahead of the main
crop, attracted more N. lugens and N. virescens, gave significantly higher yields, and preserved more
natural enemies than control fields without trap crops (Table 3). Crop recommendations in China
suggest early planting of a small area of rice with a susceptible cultivar, followed by treatment with
insecticide before planting of the main crop (IRRI 1978).
TABLE 3
Pest incidence and yields of rice fields with and without trap crops. IRRI, 19771
Mean virus incidence (%)

Treatment Grassy stunt Ragged stunt Tungro virus Total yield (t/ha)2
Trap crop3 1.9 13.8 1.3 –
Main Crop 1.2 13.8 0.5 2.8a
Control 2.5 22.1 1.7 1.9b
1 From Saxena 1982.
2 Duncan’s multiple range test, P = 0.05.
3 Planted 20 days before main crop.
4.5 WATER MANAGEMENT

Flooding of infested fields reduces populations of several rice insects and aids the adverse effects
of resistant cultivars on insect populations. In China, early spring flooding of rice stubble has
effectively been used to control overwintering C. suppressalis larvae. Mortality is very high (95-
100%) due to increased metabolic activity of larvae at that time of year and greater penetration of
stubble by flood water (Shin-Foon 1980).
Conversely, draining water from rice fields decreases populations of N. lugens (Dyck 1974a)
and the rice water weevil, Lissorhoptrus oryzophilus Kuschel (Isely and Schwardt 1934). For N.
lugens control, removal of water when the crop is near maturity is practical. However, drainage of
fields for L. oryzophilus control is expensive, due to loss of paddy water and fertilizer. The primary
consideration in determining whether water management practices integrate with plant resistance
should be the effect on growth patterns and yields of resistant cultivars when water management
practices are altered.
5. CHEMICAL CONTROL AND BIOLOGICAL CONTROL

The nature of these two control tactics makes them, by definition, incompatible. A limited amount
of research has shown that under certain conditions, however, this is not necessarily true. Dyck
(1974b) found that population levels of the N. lugens mirid predator, C. lividipennis, were adversely
affected by spray application of methyl parathion, but relatively unaffected by applications of
granular insecticides. Shin-Foon (1980) demonstrated a similar response of spider populations in
China to foliar spray and root-zone insecticide application. Low rates of granular insecticides
applied to rice in Japan have little toxicity to spider predators that help regulate leafhopper
populations (Kiritani et al., 1972).
SMITH 689
Insecticides such as ethylan, endosulfan, and BPMC are more selective and have less toxicity to
the predators C. lividipennis, L. pseudoannulata, and Microvelia atrolineata (Bergoth) than insec-
ticides such as deltamethrin (Fabellar and Heinrichs 1984, 1986; Heinrichs et al., 1982; Salim and
Heinrichs 1985). In China, Li-Ying (1982) noted differential mortality of the S. incertulas parasites
Trichogramma japonicum and T. dendrolimi in response to treatment with various insecticides. The
obvious way to combine the maximum possible effect of chemical and biological control agents on
rice insect pests is to use selective insecticides, and to use them only when action thresholds of pest
population levels have been exceeded.
6. PROBLEMS IN INTEGRATING CONTROL STRATEGIES

As evidenced by this review, various insect control methods are beginning to be integrated in the
rice producing areas of the world. However, there are both social and technical requirements
necessary for this trend to continue before true IPM programs can be established. From a social
standpoint, there is great need for continued government support of both research and extension
programs at the international, national and local levels. Support to extension is necessary for
continued development of local farmer IPM teams, trained to grow rice using proven IPM tactics
(FA0 1982, Oka and Soenardi 1978, Reissig et al., 1985).
From a technical standpoint, plant resistance as a control tactic itself, will require further
development. Sources of resistance to the rice bug Leptocorisa oratorius F., the rice caseworm
Nymphula depunctalis Guenee, the armyworm Mythimna separata (Walker), and certain other
insects are yet to be identified. Higher levels of resistance to S. incertulas, C. suppressalis, L.
oryzophilus, and several other species also need to be identified. The need for S. incertulas resis-
tance is especially critical due to the key pest status of this insect. For most insects, the need remains
for the development of cultivars with horizontal, multigenic resistance, to both slow the rate of
biotype development and broaden the genetic base of resistance.
Though significant accomplishments have been made in biological control, additional research
is necessary to demonstrate the function of biological control agents alone (Lowe 1982) or in an
integrated role with cultivars, cultural practices (Pathak and Dhaliwal 1981) and insecticides.
Certain cultural practices have been observed to contribute positively to rice insect pest control but
few have been verified to do so experimentally (FAO 1982). Due to the possibility of negative effects
of cultural practices on biological control, more research is necessary to define the interaction of
cultural and biological control when combined with resistant cultivars.
7. FUTURE STRATEGIES
Several strategies for the future improvement of rice integrated insect pest management emerge
from the discussion in the preceding section. A major strategy in future rice IPM programs should
revolve around the combined use of existing biological control agents and insect resistant rice
cultivars that possess moderate levels of resistance. Moderate antibiosis or antixenosis rice resis-
tance is effective, because it places less stress from plant allelochemical or morphological factors on
beneficial insect predators and parasites. Tolerance resistance in rice plants also has the same type
of beneficial effect on natural enemies, because it involves only plant growth characteristics, and
does not rely on increased plant chemical or morphological factors. An added benefit of all three
forms of resistance results from the fact that they place less selection pressure on pest insect
populations to develop resistance-breaking insect biotypes.
In addition to existing biological control efforts, new biocontrol agents may also prove to be the
basis of an additional strategy for future rice insect pest management. Several new fungal pathogens
have been shown to cause mortality in rice insect pests (see chapter 8), however, none of these
agents have been evaluated for their effectiveness of integration with other insect management
tactics such as insect resistant rice cultivars, insecticidal controls, or other biological control agents.
A new type of rice insect pest management strategy involving the integration of plant resistance
and an insecticide may also soon be possible, due to the adaptation of rice plant molecular gene
transfer techniques. These techniques allow integration of the two control tactics by insertion of the
toxin from the crystal protein gene produced by Bacillus thuringiensis (BT) into crop plants. BT
toxin strains with specificity for both Coleoptera (BT san diegoensis) and Lepidoptera (BT kurstaki)
have been successfully incorporated into tomato and potato plants. Although this technique is now
only possible in these two Solanaceous crops, rapid technological advances may soon allow the same
type of combination of management tactics in rice cultivars. As with all types of insecticides,
however, the development of cultivars that cause extremely high levels of mortality should be
avoided, in order to conserve natural enemies of rice insect pests and to avoid the rapid onset of BT
resistance. Resistance to BT has been documented to occur in the Indian meal moth in a single
season (McGaughey and Beeman 1988).
Finally, the development of an additional new strategy in rice insect pest management is also
possible by integrating altered cultural practices with other insecticidal, biological, or cultivar
management tactics. As mentioned previously in this chapter, alterations in existing cultural prac-
tices may have a real impact on lessening rice insect pest management problems and prolonging the
life of rice cultivars. The practice of synchronous cultivation of rice to disrupt pest insect population
levels and the production of legume or vegetable crops in between rice crops to conserve beneficial
natural enemy levels warrant continued evaluation as possible key components of an integrated
program of rice insect management. The acceptance and implementation of these tactics and others
discussed in this chapter will be dependent on the efforts of national agricultural programs to effect
change in both sociological and technological practices of rice production.
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11
The Brown Planthopper: Promises,

Problems, and Prospects*
MICHIEL C. ROMBACH* *
Boyce Thompson Institute for Plant Research at Cornell
Tower Road, Ithaca, NY, USA
KEVIN D. GALLAGHER**
Department of Entomology, Division of Biological Control
University of California at Berkeley
Albany, California, USA
Planting rice is lots of fun,

Rice for all and everyone,
Let’s all join and till the field,
There’s good life for all your yield
It’s so nice, planting rice
(Philippine song to make school children rice-conscious, 1967)
1. INTRODUCTION
RICE FIELDS DESTROYED BY the brown planthopper (BPH), Nilaparvata lugens (Stal) (Fig. 1A) are
a shocking sight — brown, dry stalks, and yellow lifeless straw remain where healthy stems and lush
greenery were before. At the start of the rice growing season small rice seedlings are carefully
transferred from the nurseries into the field — in perfectly straight rows, by hand, and one by one.
* This chapter reviews results of research only. Mention of any proprietary product does not constitute an endorsement
or recommendation for its use by the International Rice Research Institute, the Boyce Thompson Institute for Plant
Research, nor by the University of California at Berkeley.
* Present address: Department of Entomology, International Rice Research Institute, Los Banos, Laguna, P.O. Box 933.
Manila, The Philippines.
694 THE BROWN PLANTHOPPER
Fig. 1. (A) Brown planthopper (BPH) nymphs feeding on a rice stem and
(B), a spider (Lycosa sp.) feeding on BPH nymphs.
After transplanting financial burdens for the farmer remain high, regular applications of fertilizers
and insecticides are made and often hand weeding is necessary.
Destruction of the field by BPH, close to harvest, is a tragic occurrence - and has repercussions
for the farmer, his family, and his country. The large scale resurgence of the BPH is one of the most
important effects accompanying the introduction of the successful “green revolution” practices in
tropical Asian rice.
First, it should be noted that it is beyond doubt that the “green revolution” programs have filled
millions of empty rice bowls, prevented large scale famine, and saved the Asian region from
disrupting political upheavals. However, a less desirable effect of the modern practices was the
emergence of several new insect pests. Of these BPH was the most visible protagonist. We believe
that BPH can be suppressed, without losing any of the advantages gained by the green revolution.
The green revolution, some of its gains and drawbacks, and a strategy to solve the BPH impasse will
be discussed under headings of Promises, Problems, and Prospects.
The world stage. – From the heavy dependence on the new broad spectrum insecticides in the
40s and the 50s the Western world moved into the decades of integrated pest management (IPM)
(Perkins 1982, Wheatley 1987). Reduction of pesticide pressures while maintaining yield levels was
ROMBACH AND GALLAGHER 695
the most important task. However, in recent years most Western crop production has moved from
occasional shortages to huge surpluses. Therefore, gross national yields have ceased to be the
primary target for Western agricultural research; achieving optimum yield with minimum insec-
ticide inputs are no longer primary research objectives.
This is not the case in the developing world. Most tropical countries, from cotton in the Sudan
to rice in Southeast Asia should adopt a pest control philosophy typical of the 1955-1965 decade of
the Western world (Brader 1986): reliance on broad spectrum persistent chemicals is being
discredited, and a change to integrated control with less persistent and selective insecticides
encouraged. Throughout the third world serious problems with broad spectrum chemical insec-
ticides have arisen – and one of these problems is BPH resurgence on rice.
2. PROMISES
The Philippines has been a rice importing country for decades. Large quantities of rice were bought
from neighboring countries, particularly Indonesia, China, and Thailand.
However, economic and social problems intensified with continuing dependence on rice
imports. The combination of an increasing population of 32 million by 1965 and nationwide low and
unstable food production lead to growing unemployment in the rural areas, social unrest, and
massive migrations to the larger cities. To counteract these unfavorable social trends a national
program for self-sufficiency in rice was given maximum government priority with the change in
presidency and government in 1966.
2.1 THE ASIAN RICE REVOLUTION

The Philippine Rice and Corn Administration initiated the national Rice Sufficiency Program in
1966 (Barker 1984, Salas 1985) (Table 1). This program aimed to create a rice surplus by 1969 by
increasing irrigation and yields by at least 50%. Farmers were recommended to plant only the
modern high yielding varieties (HYVs) such as IRRI’s “miracle” rice variety IR8. This variety was
followed by other IRRI and national rice varieties. Farmers were also encouraged to use large
quantities of fertilizers, and to apply insecticide on a routine basis. To assist farmers financially with
these high input production schemes the Economic Development Program and national banks
arranged for low interest credit structures without collateral. The high fertilizer and insecticide
inputs caused high pre-harvest expenses. On a national level several international development
organizations and development banks arranged for large loans to the Philippine government to pay
for the program. Also, donations of equipment (e.g., irrigation pumps), helped in the success of the
development program.
The HYV seeds were distributed through government channels such as the Social Security
System to reach private farmers, and the Government Service Insurance System for government
employees. After harvest the government assured a solid price floor by procurement of all surpluses
of the 1967 wet season crop. Farmers’ acceptance of the new HYVs and the accompanying cultural
practices was virtually immediate. This was not in the least due to nationwide on-farm demonstra-
tions, massive advertisement campaigns, and rice yield contests, in which one farmer proudly
proclaimed the unlikely harvest of over 12 tons of rice from his 1 ha farm.
TABLE 1
Important events in BPH history in the Philippines
Date Event
PIX-1967 BPH rare rice insect - damage negligible.
1967 Rice Sufficiency Program initiated. For the first time the wet season harvest exceeds the demand.
Screening for resistant varieties - Mudgo is found BPH resistant.
1963-73 First large scale BPH outbreaks reported.
1970 Bph1 and bph2 genes found.
1973 Masagana 99 Program initiated. Technology packages transferred to the farmers.
1974 Introduction of IR26 (Bph1 gene) - immediate BPH suppression over large areas.
1976 Outbreaks on IR26.
1976 Introduction of IR36 and IR42 (bph2 gene) - immediate BPH suppression. Masagana 99 govern-
ment support decreases.
1977 Bph3 and bph1 genes found.
1980 Recognition of natural enemies in BPH suppression.
1982 Outbreaks on IR42.
1982 Introduction of new varieties with Bph3 gene - immediate suppression.
1987 Introduction of varieties with bph4 gene.
1988 BPH multiplies on IR66 (bph4 gene) in Negros, the Philippines - outbreaks expected.
The Philippine rice production drive under the Rice Sufficiency Program (1967-1969) and
Masagana 99 Program (Tagalog for ‘plentiful’, and 99 refers to ‘cavans’, unit of weight of rice per
ha) started in 1973 and is a true ‘green revolution’ success story. The Philippines became the main
protagonist for Asia’s rice revolution. Rice harvest nearly doubled, and the irrigated area increased
from 250,000 ha in 1968 to over 450,000 ha in 1969 (Salas 1985). The Manilan Evening News (17
Nov., 1967) reports that “a new prosperous-looking rice farmer is emerging on the scene: the high
yielding varieties farmer”. The 1967 wet season harvest exceeded the Philippine demand and for the
first time in decades the Philippines exported a small tonnage of rice. In 1968 an impressive 77,000
MT of rice was exported to Indonesia. Due to these national programs the rice harvest rose from a
meager 4.1 million MT in 1965 to an impressive 7.8 million MT in 1979. According to Dr. Norman
Borlaug, a driving force behind the green revolution: “Never before in the history of agriculture has
a transplantation of high yielding varieties coupled with an entirely new technology and strategy
been achieved on such a massive scale in so short a period of time and with so much success”.
2.2 CHEMICAL INSECTICIDES

Crucial measures were taken in the context of insect control and the subsequent BPH problems that
emerged during the Philippine rice revolution (Tables 1,2). Farmers purchased broad spectrum
organophosphate and carbamate insecticides at tax free rates with their low interest loans - an
insecticide subsidy regulation in disguise. Insecticides were also supplied by international aid
programs; in 1967 alone the Japanese government supplied 5,000 MT of broad spectrum insec-
ticides for use on rice. Most important, insecticide use was artificially increased by forcing farmers
to purchase insecticides along with the heavily subsidized fertilizers in their loan package — even if
insecticides were not required.
At the same time Indonesia experimented with management of large areas of rice culture by
western style corporations under government contract (for social consequences see also Feder,
1983). From 1965 to 1970 companies like Ciba-Geigy, Hoechst, and Mitsubishi provided manage-
ment, credit, seeds, fertilizers, and insecticide applications for over 2.5 million ha of irrigated rice.
In some areas farmers were forced to conform with synchronous planting schemes so that insec-
ticides could be applied by airplane at set schedules and insecticide use became extreme. In most
other Asian countries insecticide subsidies were installed, because insecticides were, with HYV
seeds, regarded as absolute pre-requisites for a successful rice revolution. Routine insecticide
treatments were often made to control insect populations that were far too low to cause economic
damage.
To control rice insect pests various insecticides were recommended under the technical advice
schedules of the Philippine Masagana 99 program; a compilation of this pesticide regime is given in
Table 2. The package of technology recommended calendar based insecticide treatments. For a few
pests crude thresholds were suggested, such as for stem borers at 10 or more deadhearts/m 2, 2 rice
bugs (Leptocorisa spp.)/m 2 and 25 hoppers/rice hill. The list of recommended insecticides con-
tained carbamate, organophosphate, and chlorinated hydrocarbon insecticides (Table 2). These
were the government recommended products. Field observations, however, show that the following
insecticides were most widely used (pers. comm. Entomology Dept. Staff, IRRI; see also Goodell
1979): monocrotophos (all around insecticide), chlorpyrophos + BPMC (mix for stem borers and
plant- and leafhoppers), carbofuran (all around insecticide), BPMC (for hoppers), MIPC (for
hoppers), azinphosethyl + MTMC (for stem borers and hoppers), and MTMC (for hoppers).
Heavy use of insecticides gained rapid acceptance amongst rice farmers boosted by the
subsidies. Farmers can in general not distinguish between pests and natural enemies, and they do
not know thresholds. They prefer chemicals which decimate pests immediately when either the
insect itself or its damage is evident. Rapid-action insecticides, such as those listed above are the
obvious choice to ensure dramatic, albeit short-term, control.
2.3 BPH CONTROL BY PLANT RESISTANCE

As early as 1967 varieties resistant to BPH were found by mass screening at IRRI (Pathak et al.,
1969) (Table 1). Two genes for resistance were named: the dominant Bph 1 gene and the recessive
bph2 gene. Later two other genes for resistance were identified: the dominant Bph3 gene and the
recessive bph 4. The Bph l and bph 2 genes are closely linked (Lakshminarayana and Khush 1977) as
are the Bph3 and bph4 genes (Sidhu and Khush 1979, Ikeda and Kaneda 1981). Closely linked genes
cannot be easily incorporated into the same plants.
TABLE 2
Insecticides and applications as recommended in the Philippine Masagana 99 program
Before transplanting (BT):

25 days BT: Soaking of seeds in carbofuran
21 and 14 days BT: Broadcast: carbofuran
diazinon
BHC + carbaryl
MIPC
21, 14, and 7 days BT: Spray: azinphosethyl
parathion
chlorpyrifos
thioquinox
diazinon
endosulfan
malathion
fenitrothion
mercaptophos
monocrotophos
phophamidon
toxaphene +
parathion
Afer transplanting (AT):
30 days AT
for stem borer: Broadcast: carbofuran
diazinon
BHC + carbaryl
MIPC
BHC
lindane
or spray: azinphosethyl
chlorpyrifos
thioquinox
diazinon
endosulfan
fenitrothion
malathion
mercaptophos
BHC
lindane
phosphamidon
toxaphene + parathion
For any leaf damage: Spray parathion
50 days AT: Broadcast any insecticide as mentioned under 30 days AT, for stem
borer control
IR20: as above, but only at 10 deadhearts/m2
75 days AT: Broadcast any insecticide as mentioned for stem borer under 30 days
AT
For hoppers and rice bugs Spray parathion
The genes for resistance were transferred to breeding lines in countries with BPH problems.
Varieties with genes for BPH resistance became the principal tool for BPH control (Pathak and
Saxena 1980). In the Philippines the various genes were present in many rice varieties registered by
the Philippine Seed Board (Table 1). IR26 was an early BPH resistant variety with the Bph 1 gene
– it saved Philippine rice production from massive BPH damage in 1972-73. The BPH outbreaks
were so serious that rice cultivation was prohibited in some areas until IR26 was distributed to
farmers.
The resistant varieties in combination with the chemical insecticides superbly suppressed BPH
for several years. During this time new resistant varieties were developed for subsequent distribu-
tion.
2.4 PUBLICATIONS ON BPH
The prospects of controlling BPH by plant resistance and chemicals seemed bright. To illustrate the
level of activity in resistance and chemical research we examined 1,291 publications dealing with
BPH published from 1967 to 1985. The publications were classified into the broad subject
categories of (1) plant resistance and chemical BPH control, and (2) biological and cultural BPH
control and (3) others (e.g., on taxonomy, migration, physiology). Most BPH research has focused
on plant resistance and chemical control (Fig. 2).
Fig. 2. Number of publications on BPH research (1967-1985) arranged to topic.

3. PROBLEMS
3.1 BIOLOGICAL CONTROL

Natural biological control, or natural control, is a common phenomenon - natural enemies regulate
pest populations at lower densities than otherwise occur. Populations in all natural systems, being
tropical, temperate, terrestrial or marine, are invariably influenced by some form of natural biologi-
cal control. Introduction of new pests in areas where its natural enemies do not occur, or removal
of natural enemies with insecticides mostly results in pest outbreaks (DeBach 1964, van den Bosch
et al., 1982). BPH is normally kept under biological control by diverse communities of natural
enemies - a fact to which little attention is given (Greathead 1979). Lists of BPH biological control
publications and natural enemies exist (Yasumatsu and Torii 1968, Greathead 1979, Chiu 1979), but
few report on their ecology and effectiveness (see chapter 7).
The natural enemies of rice pests include predators (Fig. 1B) and parasites (Yasumatsu and
Torii 1968, Chiu 1979, Nickel 1964, Kenmore 1980, Kenmore et al., 1984, Ooi 1986), and insect
pathogens (Rombach 1987, Rombach et al., 1987). The specific roles of most of the natural enemies
in rice are not well understood. Their combined action remarkably suppresses RPH and other pests.
Although Walker (1962) lists more than 1,400 insects attacking rice, only few ever cause economic
damage - and about one hundred species can attain a pest status (Kisimoto 1984). Even most of
these species generally occur in very low numbers and only occasionally escape control.
3.2 KEY PESTS OF RICE

Taking the numbers of insects feeding on the crop into account, there are only few key insect pests
of rice. Key pests occur on a regular basis, and cause substantial damage, even when not directly
provoked by man’s activities. Of course, all pests of cultivated crops can simply be regarded as a
result of man’s activities — he planted the crop as a monoculture in the first place. However,
virtually all rice pests, including BPH are pests because of crucial changes in agricultural practices.
These practices include: mismanagement of chemical pesticides, cultural measures such as double
cropping, wide spread planting of one or few genetically similar rice varieties, planting of long-dura-
tion rice varieties, rattoon practices (growth of a second and third crop from stubbles of the first
crop), and excessive use of fertilizers. Important key pests are stem borers (Pyralidae and Noc-
tuidae), the leaffolders Cnaphalocrocis medinalis (Guenee), Marasmia spp., and, infrequently,
armyworms such as Spodoptera spp. and Mythimna separata (Walker) and semiloopers and hairy
caterpillars (Noctuidae).
3.3 RESURGENCE
Destruction of natural enemy populations (e.g. by insecticide applications) can lead to secondary
pest outbreaks where former relatively rare insects become primary pest species (DeBach 1964,
Huffaker and Messenger 1976). In general BPH is such a secondary pest. Large BPH populations
build up in the field after insecticide applications (Kenmore 1980, Kenmore et al., 1984). Broad
spectrum insecticides kill natural enemies, in addition to BPH. BPH has a faster rate of increase
compared to most natural enemies and in sprayed fields BPH populations explode. Therefore,
insecticides are blamed for evoking BPH resurgence on rice in tropical southeast Asia (Heinrichs
et al., 1982, Heinrichs and Mochida 1984, Reissig et al., 1982, Kenmore 1980, Nishida 1975, Ooi
1980).
Predators are the major biological control agents of BPH. They include spiders (Lycosidae,
Tetrangnathidae, and Micryphantidae), crickets (Gryllidae), and small hemipterans such as Cyrtor-
hinus lividipennis. Information on their population dynamics is given by Kenmore et al. (1984).
Parasites of BPH are abundant; Chiu (1979) lists 19 species of parasites of BPH eggs, and 16
attacking nymphs and adults. The parasites undoubtedly have an influence on BPH numbers, but
whether they significantly suppress populations is not known. However, due to their size and active
habit parasites are, in general, very sensitive to pesticides.
3.4 BPH PEST STATUS

BPH rarely occurs as an economic pest in the tropics and is not a key pest. BPH was already a pest
in Korea and Japan before chemical insecticides were used. However, the ecology of BPH in these
temperate areas is different from tropical areas in that the seasonal massive immigration of BPH in
the temperate zones upsets the natural balance in favor of the pest. Similarly, BPH can be an
economic pest on tropical rice after immigration from resurgence areas; however, this resurgence
is caused by insecticides.
3.5 AGRONOMICAL ASPECTS

In tropical rice, double or triple cropping can enhance BPH population growth (Mochida and
Suryana 1979), and allows multiple generations of other pests, such as the leaffolder, to develop.
The green revolution with its irrigation projects and other cultural measures promoted planting of
several crops per year and insect pests, including BPH, increased. Synchronized planting and crop
rotation were suggested as a measure to control BPH (Oka 1979). This practice results in a fallow
period over large areas — which will lead to unstable pest and natural enemy populations. It can be
argued that pest species with a high biotic potential (e.g., BPH) benefit more from instability than
do predators and parasites — and field experiments at IRRI and in Mindanao (Philippines) do
suggest this effect. However, the minimum area for synchronous planting experiments should be
about 1,200 ha, an estimation based on BPH dispersal data (Loevinsohn 1984). Clearly, research on
this aspect of BPH population dynamics is needed to clarify the effectiveness of synchronous
planting in BPH control. However, although changing planting schemes might be technically useful,
it is socially unacceptable — planting dates are based on a myriad of factors, of which BPH risk is
only a minor one.
3.6 PRESENT PROBLEMS

Curing the 1970s and the early 1980s the BPH posed tremendous problems, and was referred to as
“a threat to Asian rice production” (IRRI 1979). However, recently BPH populations decreased in
some areas, including the Philippines, for reasons not well understood, but a decrease in insecticide
use after the Masagana 99 ‘insecticide boom’ is probably the major reason. Insecticide applications
rose more than 500% from 1966 to 1979 in Central Luzon, Philippines (Loevinsohn 1987). This was
also the period when BPH became a serious problem. A similar situation was reported for Malaysia
(Ooi 1986). Despite the present slowdown insecticide use on tropical rice is still massive: In 1982
US $13.3 billion worth of pesticides were sold worldwide, of which about 30% (US $4 billion) was
sold in the developing world (GIFAP 1983). This represents about US $1.3 billion in insecticides for
Asia and we estimate that the southeast Asian rice farmers, excluding the Chinese, spend at least
US $250-500 million/year on insecticides. China now produces several insecticides, including
monocrotophos and chlorinated hydrocarbons and Chinese insecticide usage is steeply increasing.
3.7 CASE STUDIES

3.7.1 Guadalcanal, Solomon Islands
A decade of heavy insecticide use against the cutworm, armyworm, and BPH (resurgence popula-
tions) decimated populations of natural enemies in the isolated rice growing area on Guadalcanal.
A dozen insecticide applications were made per crop (equivalent to about 1 spray per week). Broad
spectrum insecticides such as malathion, parathion, BHC, and toxaphene were used. The popula-
tions of natural enemies were destroyed to such an extent that interruption of chemical treatments
resulted in almost total yield loss to hopperburn, even when BPH resistant rice varieties were
planted. The problems increased to such a degree that rice cultivation on Guadalcanal was
terminated in 1987. The Solomon Islands are now importing rice bought on the world market.
3.7.2 Java and Sumatra, Indonesia

Insecticides were subsidized in Indonesia, and their use on rice became extreme. They were often
applied when not needed. Consequently, BPH became a major pest over vast areas in Java and on
Sumatra. In 1985-86 the point was reached that no resistant rice varieties were available, pesticide
use was extreme, and still hopperburn severe. Also, BPH vectored virus diseases, ragged stunt and
grassy stunt, became serious. This situation dominated large rice growing areas on Java and
Sumatra, an estimated 100,000 to 200,000 ha, probably Asia’s largest area ever afflicted by this
insect. Increased insecticide use only worsened the situation. In 1986 President Soeharto issued a
decree which officially adopted integrated pest management and natural enemy conservation as the
state policy for insect control on rice; it banned the use of 59 insecticides for use on this crop. Only
a few moderate to highly selective insecticides can now be used.
3.8 RICE RESISTANCE TO BPH

During 1972-1973 IR 26 was widely planted all over the Philippines. This variety has the first
identified gene (Bphl) for resistance. However, within two years BPH was able to damage IR26 in
Mindanao, an area where the variety originally was resistant (Feuer 1976). Soon BPH outbreaks on
other varieties with the Bph1 gene were reported from many other areas of the Philippines.
The introduction of IR36, IR42 and other varieties with the bph2 gene brought immediate and
successful control of the BPH outbreaks. IR42 lasted six years before it also became susceptible in
1982 in the Philippines (Peralta et al., 1983). Similar cycles of BPH outbreaks on varieties with the
Bph1 and bph2 genes have occurred in other countries in southeast Asia (IRRI 1979). Recently
varieties with the Bph 3 and bph4 genes were released. These have not yet been damaged by BPH in
the Philippines. However, moderate BPH populations can survive on plants with the Bph3 gene
(IR60, in Mindanao, Philippines) and high populations thrive on plants with the bph4 gene (IR66,
Negros, Philippines). So, over the past 15 years each new gene for BPH resistance broke down
within several generations and needed replacement by a new gene. In the greenhouse Pathak and
Heinrichs (1982) selected a virulent population that overcame resistance conferred by the bph 2
gene within 7 generations; similar adaptation rates were found by other researchers (e.g., Lee and
Choi 1981).
In the Philippine rice fields the widespread planting of varieties with the Bph 1 gene led to high
BPH mortality in 1973. However, BPH quickly adapted to the gene and IR26 was destroyed in 1976.
The BPH genes that allowed it to multiply on rice varieties with the Bph1 gene have high probability
of fixation (Hedrick 1983) in BPH populations; IR26 and related varieties were grown extensively
and immigration from other genotypes was negligible. IR36 and IR42 (bph2) were planted in areas
where IR26 was destroyed. This again led to high mortality, but BPH also adapted to the bph2 gene.
Recent results indicate that adaptation has not been lost after almost 10 years absence of Bph1
varieties (K.D. Gallagher, unpublished 1987). Therefore, adaptation to the bph2 gene occurs
without loss of adaptation to the Bphl gene. BPH able to survive and multiply on varieties with the
Bph3 and bph4 genes were recently found. From past experience, it is likely that BPH will thrive on
these varieties without losing past adaptations. If new resistant genetic materials will be incor-
porated in future varieties the “boom and bust’’ cycle (Robinson 1975) is expected to continue.
Planting rice varieties with vertical (unstable) resistance to BPH simply sets the selection and
adaptation procedure in motion. Thus rice breeding programs should emphasize the development
of rice varieties with stable resistance to BPH. Plant resistance is one of the various tactics for BPH
control that should be integrated with other tactics in a strategy primarily based on biological
control by natural enemies.
4. PROSPECTS
4.1 CONSERVATION OF NATURAL ENEMIES
On tropical rice the conservation of natural populations of predators and parasites is the prime
strategy for stable BPH control. The judicious use of chemical pesticides is the key to natural enemy
conservation. However, key pests will continue to infest fields and cause crop damage. Therefore,
efforts to control key pests without harming natural enemies are more relevant to BPH management
than controlling BPH populations after natural enemies are destroyed and BPH has resurged.
Therefore, selective control agents for lepidopteran key pests must be developed.
We are all aware of the consequences of pesticide resistance in insects, environmental Pollution
on a vast scale and, last but not the least, general cost increases of chemical pesticides. Therefore,
the importance of biological control, within an IPM framework, cannot be overstated.
4.2 PLANT RESISTANCE

Plant resistance for BPH control has a place in the total rice IPM package. New genes for BPH
resistance are found by screening and genetic analysis (Martinez and Khush 1974, Khush 1979). The
genes are mainly present in Assam rice types from north eastern India which is one of the areas of
Oryzae origin (Khush et al., 1986). New resistant varieties will continually be released in the future.
However, reliance of plant resistance as a sole tool in BPH management is unstable. Gene
rotation, and use of multilines have been suggested as strategies for gene release (Saxena and
Barrion 1985). In gene rotation a scheme is devised in which varieties with the different genes for
resistance are planted sequentially. However, BPH populations do not lose adaptation, and the final
population will be adapted to all genes. Recent experiments in Indonesia confirm these implications
(K. Sogawa, pers. comm., 1987).
New resistant rice varieties should be used sparingly. Selective pesticides can be used to contain
small scale outbreaks. The use of broad spectrum pesticides should definitely be discouraged; these
compounds hasten BPH adaptation to the resistant plants.
4.3 PESTICIDES AND BPH ADAPTATION

Natural enemies may slow the rate of adaptation of BPH populations on resistant varieties. The
reason for this phenomenon is as follows: Adapted insects live longer (Gallagher 1988) and
therefore lay more eggs compared to less adapted insects. The fitness differential increases each
day the insect lives and can produce more offspring. This is true in the absence of mortality factors.
However, natural enemies shorten the lives of all insects. They have thus a relatively more profound
impact on longer living insects, (i.e., the chance of being eaten in 5 days is 5 times as high as the
chance for 1 day). Thus, natural enemies will delay BPH adaptation to resistant rice varieties. Broad
spectrum insecticides remove natural enemies as a mortality factor. Large BPH populations will
occur in the absence of natural enemies and large populations mean increased probabilities of
adaptation. Thus, broad spectrum insecticides hasten BPH adaptation to plant resistance.
4.4 INSECTICIDE APPLICATION AND TIMING

Proper choice of insecticide and timing of the insecticide application can maximize the impact on
BPH and minimize effects on natural enemies (Aquino and Heinrichs 1985). Also, granular
applications of systemic insecticides have a wide spectrum for phytophagous insects and carry less
risk for BPH resurgence. Granules affect predators less than sprays; this partly explains the
popularity of carbofuran for use on rice. However, most granular insecticides emit insecticidal
vapors, and can kill natural enemies dwelling on the water or the lower parts of the plant (Koyama
1971). Indeed, BPH resurgence after carbofuran applications of granules was occasionally observed
at IRRI.
Heinrichs (1979) provides a table which compares the contact and systemic activity of various
rice insecticides. In general, carbamates are more toxic to BPH than organophosphate and
chlorinated hydrocarbon insecticides (Fukuda and Nagata 1969). Unfortunately, BPH has a general
low susceptibility for insecticides compared to other insects such as the green leafhopper (GLH
Nephottetix virescens (Choi and Lee 1976). At present over 35 different insecticides are recom-
mended for use against BPH all over Asia, based on effectiveness and commercial availability.
Organophosphates (diazinon) and several carbamates (BPMC, carbofuran and carbaryl) are most
widely used.
4.5 BPH SELECTlVE INSECTICIDES

Few moderate to highly selective insecticides for BPH control are available. Several carbamate
products (e.g., BPMC) can be used to suppress populations without major natural enemy destruc-
tion. However, buprofezin is highly selective and is the suggested compound for BPH control.
Routine use of this product is probably not economical. However, BPH is a rare problem in well
managed tropical rice fields, and buprofezin can be used as an emergency tool for its control on a
small scale.
4.6 JAPAN AND KOREA

In temperate Korea and Japan BPH does not overwinter and immigrates yearly (Kisimoto 1979).
Therefore, the development of populations of pests and natural enemies differs substantially from
tropical rice cultivation where populations of host and prey are continually present. In temperate
areas epidemics of BPH are initiated by long-distance migration from mainland China and the
immigrating BPH populations can overwhelm populations of natural enemies. Here serious crop
loss can be prevented by applications of insecticides in late July and early August (Kisimoto 1971,
1976). However, in both countries spray decisions are based on early season immigrant numbers
from light traps and not on actual field populations at time of spray. We think that applications with
selective pesticides and spray decisions based on actual field population numbers will reduce
applications. Furthermore, fewer applications will conserve natural enemies and thus even further
reduce the need for chemicals. Large scale field tests to compare these strategies with conventional
practices are needed.
4.7 INSECTICIDE-PLANT RESISTANCE PACKAGE

The combination of high levels of rice resistance for BPH and chemicals is unfortunate because the
deleterious effects of the insecticides are not noticeable as long as the plants remain resistant. But,
high levels of insect resistance in rice tends to break down (Gallagher 1988) due to selection of
virulent populations.
In practice, farmers are, unaware of the effect that insecticides have on natural enemies and the
importance of their conservation because plant resistance masks the deleterious effects of insec-
ticides on populations of natural enemies. Over time, BPH populations adapt to the plant resis-
tance, and their numbers explode in the absence of natural enemies. The importance of natural
enemies is not recognized by the farmer, because "the insecticides worked", while in reality his BPH
populations were kept low (despite insecticides) by plant resistance. This farmer conditioning leads
to a grossly distorted BPH management concept.
The few examples above illustrate that chemical pesticides alone are not the panacea for rice
pest problems. However, specific and effective chemical insecticides are available and have a role
in the BPH management strategy. For example, Dimilin, which contains diflubenzuron, is a molting
inhibitor for use against leaf feeding lepidoptera, and Applaud, which contains buprofezin, is a
contact action molting inhibitor for use against plant- and leafhoppers. In addition some insect
pathogens are available, and these products are highly specific for one pest species or a group.
Several insect pathogen products were tested on rice (a review of these tests is given by Rombach
et al., 1987). Products containing Bacillus thuringiensis Berliner are probably most promising for
selective lepidoptera control; B.t. products are already used to some degree in China (Hussey and
Tinsley 1981). These safe and specific chemical and microbial insecticides should be included in
IPM programs if any sprays are necessary.
Buprofezin is recommended for use and has been applied in BPH infested areas in Indonesia,
with favorable results on both BPH and populations of natural enemies; the spider community was
relatively unharmed while BPH populations were severely affected by the chemical.
5. CONCLUSION
We can conclude that (1) control of BPH with plant resistance and pesticides is not stable, and that
(2) biological control is stable in itself, but can be destroyed by mismanagement of pesticides. The
key to BPH control is selective control of other rice pests. This will conserve natural enemies, and
BPH problems will be minimized.
This conclusion has been known for many years, published, and discussed in various forums.
However, BPH outbreaks still occur on a large scale. Obviously, farmers are still making inap-
propriate decisions and misusing pesticides.
Sufficient scientific information is available for implementation of basic rice IPM systems in all
Asian rice areas; if necessary, fine tuning of these systems can be carried out on a local level.
However, local decision makers, policy makers, farmer advisors, and farmers need training in IPM
concepts and techniques. Training and extension are now the most important aspects of Asian rice
IPM.
IPM is fundamentally different from other technological packages such as distribution of rice
seeds or pesticides. In IPM the farmer makes a control decision based on levels of insects and
natural enemies in his field. Relatively simple sampling decision models were recently developed
for several rice pests (Shepard et al., 1988); the decisions are based on numbers of pests as well as
natural enemies. The farmer should be aware of the consequences of his actions, especially when
pesticides are involved. The farmer must be familiar with the local environment, pest insects, and
natural enemies. Farmer training must be recognized and incorporated into national programs.
One decade ago the Western world started to move from a dependence on large scale heavy
industry to societies based on information and services (Toeffler 1980). In rice production we have
an analogous situation; the era of Asian BPH control by technological packages of seeds and
pesticides has passed with successes and failures – the era of information on local decision making,
farmer training, and fine tuning of IPM programs has just started. The Asian farmer needs full
support to make this change.
ACKNOWLEDGEMENTS
We thank Drs. B. M. Shepard, P. E. Kenmore, A. P. Gutierrez, D. W. Roberts, E. A. Heinrichs, and
J. C. van Lenteren for sharingwith us their insights in rice entomology and IPM. Drs. D. W. Roberts
and A. P. Guiterrez are thanked for funding our respective projects. Maureen Rombach is thanked
for help with the preparation of the manuscript and the figures. This research was funded by grants
from the Office of the Science Advisor, US Agency for International Development (Washington)
and The Jessie Noyes Foundation (New York); both agencies are thanked for their support.
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IV. IMPLEMENTATION OF
RICE IPM SYSTEMS
12
Development and Implementation of

Rice IPM in Japan
KEIZI KIRITANI
National Institute of Agro-Environmental Sciences
Yatabe, Tsukuba, Ibaraki, 305, JAPAN
KUNIHIKO NABA
Hiroshima Prefectural Agricultural Experiment Station
Higashihiroshima, Hiroshima, 739-01, JAPAN
1. NEED FOR RICE IPM IN JAPAN

A CROPPING RESTRICTION ON RICE has been enforced in Japan since 1970 due to an oversupply of
domestic rice. During the last three decades, a total of 3,170,000 ha of rice fields in 1967 was
decreased by 30% to 2,250,000 ha in 1983. Concurrently, the aim of the national rice production
program has been changed from increasing the rice yield to the labor-saving production of better
quality rice. Because some 80% of Japanese rice producers manage less than 2 ha of rice fields they
depend on incomes from non-agricultural sources and spend a minimum amount of time in rice
farming. Therefore, labor-saving rice production practices have the highest priority in Japan.
Since World War II, the gain of rice yield per ha has been closely linked to an increase in the
amount of pesticide produced (Fig. 1). This suggests that the increase in rice yield was achieved
partly by the increasing use of pesticides. The recent demand for high grain-quality rice has
encouraged the preventative application of insecticides to control heteropterous bugs. Additional-
ly, the price of domestic rice is relatively higher, i.e., 3 and 6 times as high as in the EC and USA,
respectively; the relative cost of pesticide application is very low. Rice IPM in Japan has to be
714 RICE IPM IN JAPAN
Fig. 1. Annual changes in rice yield, and the amount of pesticides and insecticides produced in
Japan. Insecticides for rice total about one-half of the total amount of insecticides produced.
Fig. 2 Percentage of rice area affectcd by major insect pests in Japan. RSB: striped stem borer, GRL: green rice
leafhopper, WBPH whitebacked planthopper, RLB: rice leaf beetle, RWW: rice water weevil.
KIRITANI AND NABA 715
Fig. 3. Percentage of rice area affected by insect-borne disease in Japan. RSV: Stripe, RDV: Dwarf, RYD: Yellow
dwarf.
developed by compromising the following demands: reduction of production cost, including the
minimum use of pesticides and fertilizer, and the use of labor-saving techniques.
2. SHIFT OF TARGET PESTS SINCE AROUND 1970

Since the early 1970s, significant savings of labor have been attained by the mechanization of both
transplanting and harvesting. Mechanization has caused a significant change in the status of insect
pests and insect-vectored diseases compared to that of the 1960s (Figs. 2 and 3).
2.1 PEST STATUS BEFORE 1970

The yellow stem borer, Scirpophaga incertulas which is monophagous on rice, disappeared from
southern Japan after the introduction of organochlorine insecticides. Some univoltine insects such
as the black rice bug, Scotinophara lurida, and the rice stink bug, Lagynotomus elongatus, also
decreased remarkably before 1960.
The striped stem borer, Chilo suppressalis, was the most serious pest throughout Japan until
1960 and at its peak level infested 42% of the total area of rice fields. Accordingly, this species
developed resistance to BHC and parathion because of heavy insecticide applications to control it.
An ever-decreasing trend in the infested area, however, has been observed since the late 1960s and
this insect occurred in less than 10% of the rice fields in the 1980s. This decrease was attributed to
changes in such agronomical practices as the early planting of rice, use of high tiller-number type
varieties and mechanization of both transplanting and harvesting rather than to the intensive use of
insecticides (Kiritani 1979).
By contrast, populations of the green leafhopper, Nephotettix cincticeps, the small brown
planthopper, Laodelphax striatellus, and the heteropterous stink bugs, which are polyphagous and
m ultivoltine, have remained constant or increased since 1970. N. cincticeps populations began
increasing on early-season rice in southern Japan in the 1950s when the insecticides were intro-
duced in rice fields for the control of C. suppressalis. N. cincticeps, having developed resistance to
organophosphates by the early 1960s and then to carbamates by the early 1970s (Hama 1975,
Asakawa and Kazano 1976), affected 40-50% of the rice fields annually during the last two decades
(Fig. 2).
2.2 PEST STATUS AFTER 1970

2.2.1 Change in Pest Status due to Machine Planting
The machine planting of early maturing rice varieties has resulted in shifting the transplanting date
forward and in increasing plant density. It induced changes in pest status of some species which
reproduce at the vegetative stage of rice growth. Although 100,000-200,000 ha of rice fields had been
annually infested by the rice leaf beetle, Oulema oryzae, in eastern Japan during the 1960s, the
infested area increased to some 600,000 ha in the middle 1970s after the introduction of machi ne
planting. Moreover, the rice plant weevil, Echinocnemus squameus, and the small rice miner,
Hydrellia griseola, have increased in abundance over the country, particularly in northern Japan,
since the late 1970s. The rice skipper, Parnara guttata, which was abundant in the late planted rice,
has decreased recently.
Since the early 1970s, the machine planting of rice was followed by the nursery tray insecticide
treatment, where granules are applied onto seedlings just before transplanting. The nursery tray
treatment is very effective in controlling early season insect pests (e.g., O. oryzae) and insect-borne
virus diseases. Consequently, the area infested by these pests began to decrease or level off. The
incidence of O. oryzae in some regions in the Tohoku district decreased from one-sixth to one-eighth
of its maximum.
The nursery tray treatment has several advantages compared to that of conventional broadcast-
ing. It requires only one half of the conventional insecticide rate and less labor, and has a longer
residual effect. However, because it is prophylactic, it is not economical if the incidence of target
pests is less than the control threshold.
2.2.2 Change in Occurrence of Insect-Borne Virus Diseases

In the 1960s, epidemics of rice stripe disease vectored by L. striatellus occurred in association with
the introduction of early planting middle-season rice. But an ever-decreasing cultivation of wheat
and barley, the host plants for the first generation of the vector, resulted in reduction in the
incidence of stripe since the late 1960s. However, the increased cultivation of wheat and barley since
the late 1970s promoted the recent prevalence of stripe in some districts. Consequently, newly
released stripe resistant varieties have rapidly replaced conventional varieties over a large hec-
tarage in the Kanto district. Because these varieties are resistant only to stripe, but are susceptible
to L. striatellus, an increase of the vector population on these varieties is becoming a problem
particularly for growers of the conventional varieties (Takayama 1984).
~~
In southern Japan two epidemic periods of the rice dwarf disease vectored by N. cincticeps have
occurred during the last 40 years (Kiritani 1983). The first epidemic, 1953-1966, coincided with an
extension of early rice cultivation, while the second, 1967-1974, started when the area planted to
wheat and barley was less than half of the previous maximum. The second epidemic was charac-
ferized by the spread of dwarf disease to middle-season rice areas. However, from 1973 when the
infested area attained a maximum of 330,000 ha, the incidence of dwarf continued to decline as a
result of the encouragement of winter and spring ploughing of fallow paddy fields and winter
cropping. The wide adoption of the nursery tray treatment also contributed to a decrease in the
incidence of dwarf to less than one-third of maximum incidence.
Rice yellow dwarf disease, of which the causal agent is a mycoplasma, is transmitted by N.
cincticeps. It occurred in over 200,000 ha in the early 1960s and 30,000 ha in the 1970s and then
decreased to 5,000 ha in the 1980s. This decrease was due to the destruction of the transmission
cycle by early harvesting, ploughing down infected ratoon plants, and the nursery tray treatment.
2.23 Increase in Occurrence of Long-Range Migratory Insects

The whitebacked planthopper, Sogatella furcifera, the brown planthopper, Nilaparvata lugens, and
the rice leaffolder, Cnaphalocrocis medinalis, are long-range migratory insect pests that are
believed to immigrate from mainland China. Before the 1960s, these oversea migrating pests were
chemically controlled simultaneously with the first generation of C. suppressalis. The omission of C.
suppressalis control due to the current reduced infestation appears to have encouraged outbreaks
of these migrating pests. Though C. medinalis was a minor pest until the early 1960s, its high
incidence has continued since 1967 when the first outbreaks were observed in the Kyushu and
Hokuriku districts.
The first sign of resistance to organophosphorous insecticides in S. furcifera was recognized in
1985 (Hosoda unpublished). N. lugens had developed resistance to organophosphates by the early
1970s and then to carbamates by the late 1970s (Nagata et al., 1979, Hosoda 1983). These patterns
of resistance are considered to reflect the insecticide applications in the place of origin of these
migrating insects.
2.2.4 Build-up of Insect Pests Affecting Grain Quality

Production of better grain-quality rice has been emphasized since 1970. If there are more than two
spotted grains per 1,000 grains, the lot of rice is ranked at a lower grade in proportion to the
percentage of spotted grains. The cause of spotted grains is mainly heteropterous stink bugs, but
sometimes the rice plant weevil, Echinocnemus squameus, the rice thrips, Baliothrips biformis, and
the rice white-tip nematode, Aphelenchoides besseyi are responsible for the spotted grains. These
pests have induced increasing frequency of pesticide application after rice heading. Especially, the
calendar-based control of the heteropterous stink bugs became common for early-maturing
varieties grown in the mountainous areas. The prevalence of stink bugs is believed to have resulted
from the increase of fallow paddy fields as a result of the rice cropping restriction.
The control threshold for stink bugs is so low that rice growers control them prophylactically.
For example, in the case of the southern green stink bug, Nezara viridula, the control threshold
corresponding to 0.1% of spotted grains is two adults per 50 net strokes at the milky stage of rice
plant growth (Nakasuji 1973). The control of stink bugs also involves the difficult task of protecting
rice from their continuous attacks from heading until harvest.
2.2.5 Invasion of the Rice Water Weevil

The rice water weevil, Lissorhoptrus oryzophilus, a species indigenous to America, was first dis-
covered in Aichi prefecture in central Japan in 1976 (Tsuzuki and Isogawa 1976). L. oryzophilus is
believed to be the first exotic species established in the history of rice cultivation in Japan, All of the
rice insect pests, except such overseas migrant species as N. lugens, are endemic in Japan. After ten
years of its first occurrence, the area invaded by 1985 amounted to 770,000 ha of rice fields covering
Honshu, Shikoku, Kyushu and the Nansei islands. In 1986, this weevil was discovered in Hokkaido,
the last uninfested prefecture remaining in Japan. No effective natural enemies of this weevil have
been recorded in Japan.
Recommendations to control L. oryzophilus are delay of transplanting to evade peak incidence
of immigrants and management of water with intermittent and shallow irrigation to prevent oviposi-
tion and feeding by adults. These cultural methods, however, have not commonly been practiced
because of problems associated with water availability. Instead, chemical control by the nursery tray
treatment and/or the application of granules onto the surface of irrigation water have widely been
practiced. Thus, the invasion of L. oryzophilus resulted not only in an increase of control costs but
also in the nation-wide utilization of the nursery tray treatment.
3. COMPONENTS FOR THE RICE IPM PROGRAM

The IPM program should be carried out as shown in Fig. 4. The current status of available
components for IPM are described below.
3.1 MONITORING AND FORECASTING

The nation-wide forecasting network for rice pests is composed of approximately 600 staff members
who work at the Prefectural Agricultural Experiment Stations and the District Plant Protection
Service Stations. The locality-based information concerning the occurrence of rice pests and
control recommendations are delivered to rice growers through wire-relays, telephone service and
television, by way of local governments, the Agricultural Cooperatives and the Agricultural Mutual
Aid Associations. Since the late 1970s, the pest surveillance data in several prefectures have been
processed with computer simulation models for use in forecasting.
During 1960-75, several ecology-oriented research studies were conducted on natural popula-
tions of such rice pests as N. viridula (Kiritani and Hokyo 1970), N. lugens (Kisimoto 1965, Kuno
1968), S. furcifera, N. cincticeps and L. striatellus (Kuno 1968), N. cincticeps (Hokyo 1972) and N.
cincticeps and the rice dwarf virus disease (Kiritani et al., 1970; Nakasuji and Kiritani 1972; Nakasuji
Fig. 4. General flow-chart of implementing IPM programs for rice in Japan

(adapted by modifying Kiritani 1976).
et al., 1975,1985). Kiritani and his co-workers applied the systems approach to the IPM system of
rice dwarf disease by incorporating the interactions among rice plants, N. cincticeps, spiders and
virus, These studies contributed to more precise prediction of pest occurrence by elucidating the
regulatory mechanism of population density.
A systems model of rice dwarf virus epidemiology was first proposed by Nakasuji and Kiritani
(1972) and developed by Nakasuji et al. (1975) and Kiritani and Nakasuji (1977). However, some of
the relationships involved in these models were empirical. Thus, they are regarded as tactic models
that cannot be applied to different localities in their original forms. Although it is important to
develop such tactic models that are applicable to other localities, the necessity of time-and-labor
consuming studies precludes further development. Development of strategic models analyzing the
general behavior of systems by mathematical equations is also needed. Such a study has recently
been published by Nakasuji et al. (1985).
3.2 ECONOMIC INJURY LEVEL (EIL) AND CONTROL THRESHOLD (CT)

3.2.1 Establishment of the EIL
Twenty-two EILs of ten insect pests of rice were calculated assuming a gain threshold of 5%
(Andow and Kiritani 1983). Except for the EILs of O. oryzae (Emura and Kojima 1978, 1980;
Kojima and Emura 1979) and C. suppressalis (Koyama 1977), that were developed from life-table
data, these static EILs were calculated from loss assessment experiments.
A dynamic EIL incorporating models of plant growth, insect population dynamics and weather
conditions has not yet been developed for rice insect pests. Kiritani (1973) has suggested a tabular
key for decision-making in controlling planthoppers aiming at the dynamic EIL. The key arranges
CTs as a function of rice cropping seasons, planthopper species, wing type of adults and their
density. Because rice yields are predicted by the product of four components (see Table 1) arranged
along the time axis of plant growth, the quantitative analysis of the relationship between these
components and amount of damage due to insect pests will be another fruitful approach toward the
dynamic EIL.
TABLE 1
Rice insect pests affecting yield components
Insect Pests Panicles/m2 Grains/Panicle Ripened Grains(%) 1000-Grain Weight

SSB, 1st generation •
SSB, 2nd generation 0 • 0
RDV vectored by GRL •
Sap sucking by GRL • 0
RSV vectored by SBPH • 0 • 0
RWW •
RLB •
WBPH • 0
BPH • •
Note: Heavy damage: • , Light damage: 0
SSB: Striped stem borer, GRL:green rice leafhopper, RDV Dwarf, SBPH: small brown planthopper, RSV Stripe,
RWW: rice water weevil, RLB: rice leaf beetle, WBPH whitebacked planthopper, BPH: brown planthopper.
3.2.2 Use of the CT
Each prefecture annually publishes a pest control guide that is utilized by technical officials at the
Agricultural Extension Service Office and the Agricultural Cooperative Associations. Table 2
shows the number of prefectures specifying the CTs in their guides for 1985. Most of the CTs have
been established since the early 1970s. Almost all of the prefectures in western Japan, however,
published the CTs for S. furcifera and N. lugens in the 1960s.
Although an increasing number of prefectures are recommending the CTs, CTs for N. lugens
and L. oryzophilus, for example, appeared in the guides of only 20 and 12 prefectures, respectively,
corresponding roughly to one-half and one-quarter of the prefectures concerned. The reasons that
prefectures do not recommend CTs are as follows: (a) difficulty in predicting pest damage due to
lack of research on loss assessment; (b) many CTs so far developed are of the static type and only
applicable to a specific growth stage of rice plants; (c) deep-rooted thought of perfect control or
“sterilization” of the crop without permitting even slight damage prevents rice growers from
accepting the concept of CT (their overdependence on insecticides has been supported by the
relatively low control cost as compared to the monetary value of rice); (d) prophylactic controls
TABLE 2
Use of the control threshold (CT) for major rice insect pests in Japan
District No. of prefec- No. of prefectures specifying CT in the pest control guide
tures SSB WBPH BPH GRL SBPH RWW RLB Stink Bugs
Hokkaido 1 0 0 – – 0 – 0 0
Tohoku 6 5 2 0 1 0 2 3 0
Hokuriku 4 2 1 0 1 0 2 1 0
Kanto, Tokai 13 0 5 6 1 0 3 1 1
Kinki, Chugoku 11 2 4 8 2 2 5 2 0
Shikoku, Kyushu 12 1 5 6 1 1 0 0 0
TOTAL 47 10 17 20 6 3 12 7 1
Note: No occurrence (–)
SSB: striped stem borer, WBPH: whitebacked planthopper, BPH: brown planthopper, GRL: green rice leafhopper, SBPH: small
brown planthopper, RWW: rice water weevil; RLB: rice leaf beetle.
such as the nursery tray treatment and aerial applications of insecticide carried out according to the
prefixed flight schedules of helicopters, negate the utilization of CTs.
3.3 SELECTIVE INSECTICIDES

Use of selective insecticides that are only effective against target but not against non-target species
is very important in the implementation of rice IPM. Buprofezin, registered in Japan in 1984,
showed selective toxicity to several species of Homoptera by inhibiting the biosynthesis of chitin.
Because this insecticide is reported as being safe to spiders, predacious mites and wasps (Shibuya
1984), the risk of N. lugens resurgence may be kept to a minimum.
3.4 RESISTANT VARIETIES

Resistant varieties form another key component in rice IPM. The widespread planting of varieties
resistant to the rice stem maggot, Chlorops oryzae since around 1960 has greatly reduced damage
caused by this fly. Replacement of susceptible varieties with resistant ones also contributed to the
elimination of waika, a tungro-like disease, from Kyushu in the early 1970s (Shinkai 1977). As
mentioned earlier (Sec. 2.2.2), an increasing cultivation of varieties resistant to rice stripe disease
has been practiced in northern Kanto since 1982 and the area planted to these varieties reached
66,000 ha in 1985. Because these varieties are resistant only to the disease, but not to the vector, the
omission of chemical control has allowed an increasing vector density on these resistant varieties.
This in turn may result in a high incidence of the disease among susceptible varieties grown near the
disease resistant varieties. Thus, breeding of varieties resistant to both stripe disease and the vector
is needed.
4. IMPLEMENTATION OF RICE IPM PROGRAMS: CASE STUDY

There have been significant advances in the development of individual components to be utilized
for rice IPM programs (e.g., studies on pest population dynamics and the development of control
measures). On the other hand, the implementation of IPM programs by farmers is only partial
because of the various reasons mentioned earlier. Nonetheless, IPM oriented practices or cam-
paigns have been in progress somewhere in Japan at varying degrees as discussed below.
4.1 REDUCTION IN FREQUENCY AND RATES OF INSECTICIDE APPLICATION AND

UTILIZATION OF INSECTICIDES LESS TOXIC TO NATURAL ENEMIES
The widespread use of BHC for controlling rice insect pests including C. suppressalis during the
1950-60s induced the build-up of N. cincticeps as a result of disruption of natural enemies (e.g.,
spiders). Also it resulted in the development of resistance to BHC in C. suppressalis and BHC
residues caused contamination of the environment including food and human milk. Kiritani and his
co-workers at the Kochi Prefectural Institute of Agricultural and Forest Science experimentally
demonstrated that even BHC granules lethally affect spiders through food-chain toxicity (i.e. from
irrigation water through rice plants and leafhoppers to wolf spiders) (Kiritani and Kawahara 1973).
As a result, the use of BHC for rice pest control was restricted in Kochi prefecture in 1969, two years
before the nation-wide ban on the use of BHC.
The first IPM pilot project started in Kochi and was aimed at enlightening the importance of
IPM to extension officers and scouts at agricultural cooperatives (Kiritani et al. 1971, 1972). The
objectives of this project were to demonstrate the (1) feasibility of the replacement of BHC with
organophosphates in the control of C. suppressalis; (2) effectiveness of selective insecticides; and
(3) feasibility of reducing rates and frequency of insecticide application. This project, using
demonstration fields located in different regions, was carried out by participation of persons from
various agricultural sectors including farmers. The three-year project demonstrated the following
results: (1) Organophosphates (i.e., MEP and diazinon) were more effective in controlling C.
suppressalis and gave rise to a greater yield than the conventional control using BHC; (2) In the
selective insecticide plot, where BPMC was used for control of N. cincticeps and chlordimeform,
MEP and cartap for C. suppressalis, the ratio of prey (mainly N. cincticeps) to spiders changed in
favor of the predators after insecticidal treatment. The rice yield of the IPM plot was nearly equal
to the plot with the conventional control schedule in spite of the fact that the frequency of
applications was reduced by 1-3 times and the rate was lowered to one-half; (3) Satisfactory control
of rice dwarf disease was attained by the nursery tray treatment with ethylthiometon at a rate of
one-half that of the conventional treatment (Kiritani 1974). This method was not only labor saving
but also as effective as that of 2-3 broadcast applications of insecticide in the rice field.
4.2 DECISION-MAKING USING CTS FOR SIMULTANEOUSLY OCCURRING PESTS

Damage due to C. suppressalis has been declining in Akita prefecture as in other parts of Japan
since around 1970. Nevertheless, there was no sign of reduction in the aerial spray against C.
suppressalis. Koyama (1973, 1975, 1977, 1978) at the Akita Prefectural Agricultural Experiment
Station determined the CTs for the first generation of C. suppressalis and O. oryzae which occur
simultaneously. If the occurrence of these two pests is less than the respective CTs, aerial control is
deleted.
The process of implementing the program is as follows: (1) A high correlation between the
percentages of injured leafsheaths and dead hearts occurred in the first generation. This is because
larval densities during the corresponding period were relatively stable. Therefore, the CT for
C. suppressalis was determined as 12% injured leaf sheaths at the time of insecticidal application.
(2) A total of 245 fields located in the prefecture were regularly surveyed for the occurrence of
C. suppressalis from 1971 by staff members of the District Plant Protection Service Stations. The
survey showed that the annual incidences of C. suppressalis and O. oryzae were always below the CTs
established for both species (Kobayashi and Koyama 1977). Eventually by 1977, aerial spray against
the first generation of C. suppressalis was suspended (Fig. 5). (3) Chemical control including aerial
sprays against the second generation has continued due to the absence of a CT for this generation.
Fig. 5. Changes in the area of occurrence of Chilo suppressalis and area treated with
insecticides in Akita prefecture (Koyama 1985).
4.3 PARTICIPATION OF FARMS IN PEST SURVEILLANCE FOR THE DECISION-

MAKING OF LOCALITY-BASED CONTROL
In view of the decreasing occurrence of C. suppressalis in Niigata prefecture since the early 1970s,
monitoring of C. suppressalis by teams of technical staff at the Agricultural Cooperative Associa-
tions and the Agricultural Mutual Aid Associations and by voluntary farmers has been initiated. On
the basis of these data, decision-making for the community-based control of C. suppressalis was
made after group discussions led by the staff of the District Plant Protection Service Station. Further
details of this “bottom-up” approach are explained below.
The follow-up surveillance for damage due to C. suppressalis was initiated on 1,500 ha in 1968
after insecticide application had been discontinued. This campaign was supported by rice growers
and the area subjected to surveillance increased greatly after 1975. As a result, Kojima and Emura
(1983) at the Niigata Agricultural Experiment Station standardized the monitoring procedure to
determine the practical CT for the first generation of C. suppressalis . Accordingly, pest surveillance
(see Table 3) in 1984 was extended to 52% of the municipalities in the prefecture covering 89,000
ha (i.e., 61% of the total rice fields). The surveillance showed that in 54,000 ha of rice fields. C.
suppressalis control was not necessary. The final decision for omission of control, however, was
made by agreement of all concerned farmers. It is worthy to mention that many active farmers
participated voluntarily in monitoring C. suppressalis in nine municipalities in 1984.
TABLE 3
Decision chart for control of the first generation of Chilo suppressalis used by
Ojiya City Plant Protection Association in Niigata prefecture (Emura 1982.)
Monitoring time: Grade Injured hills (%) Guidelines for control decision
Null 0 Control is unnecessary
Late June Light 1-10 Continued scouting for rice fields with a higher % of injured hills
(First generation of
C. suppresalis ) 11 - 29 Collective area-wide control
Medium 30 - 59 Treat heavily injured fields followed by collective area-wide control
Severe 60 - 89 Treat all the early planted fields followed by collective area-wide control
Very severe 90 or more Conduct collective area-wide control twice
Note: Sample size is 25 hills per rice field.

In the area where control for the first generation of C. suppressalis was recommended, sudden
outbreaks of the second generation occurred over 15,000 ha in 1984. The outbreaks confirmed the
importance of surveillance. It was also revealed that the outbreaks were partly due to the develop-
ment of resistance in C. suppressalis to some organophosphates. In the advanced municipalities, O.
oryzae and rice blast disease, in addition to C. suppressalis, have been monitored in reference to the
practical CTs for these pests.
4.4 COMMUNITY-BASED CONTROL OF THE RICE WATER WEEVIL USING THE CT

Earlier, the complete extermination of L. oryzophilus was attempted when its distribution was still
limited to a few localities. Presently, it has extended its distribution throughout Japan. Meanwhile,
a CT was established for this species by Tsuzuki et al. (1983). In 1983 when L. oryzophilus was first
discovered in Hiroshima prefecture, the CT was used for the decision-making of insecticidal control
by the local plant protection teams composed of technical officers of various agricultural bodies and
active farmers. The teams as a whole worked about 850 man-days per year to monitor the popula-
tion density of immigrating adults and the percentage of injured hills in about 8,000 sample rice
fields chosen at a rate of 2-3 fields per 10 ha.
Decisions for chemical control in each locality were based on the incidence of various rice
pests, including L. oryzophilus, with reference to the transplanting date. Decisions were made
through discussion participated in by all members of the monitoring teams and technical officers of
the District Plant Protection Service Station and the Agricultural Improvement Extension Service
Office. Consequently, where the nursery tray treatment against O. oryzae was already being prac-
ticed additional control against L. oryzophilus in the newly invaded area was omitted. Similarly, rice
fields planted after mid-June were not treated with insecticides, because little damage was expected
in the late planted rice. In spite of an increasing infested area and density of L. oryzophilus, the area
of untreated rice fields amounted to 23%, 60% and 21% of the total infested area in 1983,1984 and
1985, respectively.
4.5 COMPREHENSIVE PLANT PROTECTION APPROACH BY

ACTIVE FARMER GROUPS
To minimize pesticide use and to avoid human poisoning, the treat-when-necessary control of rice
pests has been practiced in Fukuoka prefecture by groups of 200 rice growers under the leadership
of Une (1984) at the Fukuoka Agricultural Improvement Extension Service Office since 1980. The
objectives are as follows: (1) Transfer of technical information on rice IPM to farmers; (2) Im-
plementing pest control with the minimum use of pesticides; (3) Monitoring leaf- and planthoppers
and natural enemies by means of a sight counting board (Fig. 6); (4) Promoting sales of the rice
produced.
This campaign assisted by the Agricultural Cooperative Association had made a drastic change
in the pesticide-application-calendar. The 1985 calendar recommended 1 and 3-4 applications in
the nursery tray and production field, respectively, while as many as 7 and 5 applications in the
Fig. 6. Use of the sight counting board. Rice plants are beaten and the number of insects that fall
on the board are estimated (Une 1984).
nursery tray and field, respectively, were recommended in 1980. The rice produced in this way has
been on the market since 1983 and amounted to 360 tons annually in 1984 and 1985.
5. FUTURE STRATEGY AND PROSPECTS
5.1 PROPOSAL OF THE STATISTICAL EIL TO MINIMIZE

INSECTICIDE APPLICATION
In the Philippines, the gain threshold ( =cost of control/market value of rough rice) for C. medinalis
is calculated as $29.30 (U.S. dollars) per ha/$0.15 per kg = 195 kg/ha (Bautista et al., 1984). This
value is equivalent to 8.5% of the average rice yield in the Philippines (ie., 2,300 kg/ha in 1982).
Mochida (1986) also calculated it as 6.6% for black bug control in the Philippines. Because the price
of rice and the rice yield per unit area in Japan are 5 and 2.5 times as high as those in the Philippines,
respectively, the estimated gain threshold is as low as 1% of the rice yield. This surprisingly low level
of the gain threshold is due to the government supported rice price.
Therefore, even if the amount and number of insecticide applications increase further in the
near future, their effects on production cost in Japan would be little. The diverse and complicated
insect pest fauna from the seed bed to harvest and the demand for high quality rice grains would
stimulate the more intensive use of insecticides. However, the price of rice has recently approached
its support limit, so labor-saving and low-cost technologies are required in rice production. Rice
growers are also becoming more prudent in use of pesticides for various reasons already men-
tioned.These factors will encourage the development of EILs and CTs for various pests and their
wider use in the future.
Because pest control recommendations are utilized over a wide area and include several
thousand rice growers, they are made in favor of chemical treatment. When an overall density of
pests in one locality is just equal to the CT or EIL, the pest density in half of the fields would be less
than CT or EIL. Consequently, Kidokoro and Kiritani (1982) have proposed the use of a statistical
EIL for small size farm conditions. This gives a statistical projection for the probability that a given
field has a pest density greater than the EIL (Fig. 7). With a within-field-EIL of 30% hill infestation
for C. suppressalis, the proportion of fields with greater than 30% infestation can be calculated. We
can make recommendations to individual growers concerning the insecticide control with the
probability of risk that his field has a pest density greater than the EIL.
Fig. 7. Frequency curves of Chilo suppressalis injury to paddy fields, for four within paddy mean injury levels (m).
The frequency distribution follows a poisson distribution (Kidokoro and Kiritani, 1982). The area under
the curves that is to the right of the CT gives the proportion of paddies with infestations causing economic
damage.
5.2 COMBINATION OF CONTROL MEASURES FOR RICE IPM PROGRAMS

In developing and implementing the IPM system for rice in Japan, some socio-economic as well as
technological problems rather specific to Japan should be taken into consideration. In other words,
the IPM system should be tailored to fit Japanese conditions.
Rice is the staple food for Japanese, so the government policy is self-supporting of domestic
rice. The total area planted to rice is 2,246,000 ha, which is equivalent to 55% of the total arable land
in Japan. The average yield of rice is 4,590 kg/ha. The number of rice growers is 3.7 million, so 85%
of the farmers are growing less than 1 ha of rice (MAFF 1984). The production cost of rice in Japan
is estimated to be 5-6 times higher than that in the USA and the high price of rice in Japan reflects
this situation.
The small scale and intensive system of rice cultivation in Japan requires further reduction of
production costs through the saving of both labor and resources (e.g., pesticides), without reduction
in yield. The recent demand for the high-quality rice, however, likely works in favor of overuse of
insecticides, particularly after heading. On the other hand, the nursery tray treatment contributed
not only to the reduction of insecticide applications in terms of amount and frequency, but also to
the saving of labor. The prophylactic nature of this technique remains to be solved.
Consequently, development of EILs for important rice pests is urgently needed. The cor-
responding CTs can be established by population dynamics studies of the pests. The introduction
of the statistical EIL being associated with the field inspection by growers themselves will contribute
to the rational use of pesticides.
Because major rice insect pests in Japan are endemic, the efficacy of introduced natural
enemies will be limited. The ecological balance between insect pests and native natural enemies
should be maintained in favor of the latter through the manipulation of environmental conditions.
In this connection, utilization of more selective insecticides that are safer to natural enemies as well
as aquatic organisms and wildlife is important.
Many rice varieties of the japonica type are highly susceptible to rice leaf- and planthoppers
and insect-borne virus diseases. Utilization of pest resistant varieties is most promising for reducing
the production cost of rice as well as reducing the use of pesticides. Integrated use of resistant
varieties with pesticides and natural enemies will be the final IPM program in Japan. Additionally,
international cooperation on forecasting and control of the long-ranged migratory insect pests will
become increasingly important among the neighboring countries irrespective of their political
regimes. The development of IPM for rice in Japan should be approached from domestic and
international viewpoints.
ACKNOWLEDGEMENTS
Thanks are due to the following persons for their valuable comments in preparing the manuscript:
K. Emura, J. Koyama, Y. Une, K. Nakazawa, T. Kidokoro, H. Kawamoto, A. Fujiie, E. Yano, A.
Tatara.
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Kobayashi, J. and J. Koyama. 1977. Studies on the sampling method for the survey of rice pests. Ann. Rep. Plant. Prot. North.
Japan. 28 : 30-37. (In Japanese).
Kojima, A and K. Emura. 1979. Threshold density of the rice leaf beetle, Oulema oryzae Kuwayama, for insecticidal control.
II. Estimation of the threshold density. Jpn. J. AppL Ent. Zool. 23 : 1-10. (In Japanese, English summary).
Kojima, A., and K. Emura. 1983. Decreased incidence of rice stem borer, Chilo suppressalis Walker, in Niigata Prefecture
and control system corresponding to the reduced levels of this insect pest. J. Niigata Agric. Expt. Stn. 33 : 73-80 (In
Japanese, English summary).
Koyama, J. 1973. Studies on the diminution of insecticide application to rice stem borer, Chilo suppressalis Walker. I. The
relation between the damage caused by rice stem borer and the yield of rice. Jpn. J. Appl. Ent. Zool. 17 : 143-153. (In
Japanese, English summary).
Koyama, J. 1975. Studies on the diminution of insecticide application to the rice stem borer, Chilo suppressalis Walker. II.
The economicinjury level of the rice stem borer and its predictive estimation. Jp J. Ent. Zool. 19 : 63-69. (In Japanese,
English summary).
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Pyralidae). Appl. Ent. Zool. 12 : 213-224.
Koyama, J. 1978. Control threshold for the rice leaf beetle, Oulema oryzae Kuwayama (Coleoptera: Chrysomelidae). Appl.
Ent. Zool. 13 : 203-208.
Koyama, J. 1985. Pest control and ecology: conditions for the establishment of integrated pest management in Japan.
Shokubutu-Boeki 39 : 455-460. (In Japanese).
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Ministry of Agriculture, Forestry, and Fisheries (MAFF). 1984. Rice cropping in Japan. Division of Crop Production,
Agricultural Production Bureau. Tokyo: Chikyu-sha. 464 p. (In Japanese).
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1986. IRRI & TARC. (In Japanese, English summary).
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236 p. (In Japanese).
13
Development of an Integrated Pest Management

System for Rice in Latin America
G. WEBER
Centro Internacional de Agricultura Tropical
Apartado Aereo 6713
Cali, Colombia.
O. PARADA
Federacion Nacional de Arroceros
Apartado Aereo 52772
Bogota, Colombia.
1. INTRODUCTION
RICE IS ONE OF THE MAJOR food crops in Latin America with an averageper capita consumption of
28 kg white rice per year. The highest consumption is in Guyana (112 kg/yr) and the lowest is in
Guatemala (4 kg/yr) where maize is still the staple food crop. Total production in 1983/85 (FAO
1986) was about 16.5 million mt of unhulled (paddy) rice with average yield of 2.4 mt/ha for favored
upland rice, 4.2 mt/ha for irrigated rice, and 1.0 mt/ha for unfavored upland rice (CIAT 1985).
Average production costs were 161 US$/mt of paddy in 1984/1985, varying between 209 US$/mt in
Colombia and 90 US$/mt in Chile (Pulver 1985).
Total production costs for irrigated rice, expressed in equivalents of kg paddy/ha to allow for
more easy comparisons between countries, has in tropical countries like Colombia and Ecuador
reached 5,160 kg/ha and 4,000 kg/ha respectively. This has caused low profitability of rice cropping
because average yields in these countries are around 5,230 kg/ha and 4,140 kg/ha, respectively.
Plant protection costs in Colombia and Ecuador contribute 24.9% and 18.4% respectively to the
total cost of production. Pest control expenses are thus a major cause for increasing costs and
734 INTEGRATED PEST MANAGEMENT SYSTEM IN LATIN AMERICA
decreasing profitability of rice cropping in these countries. (FEDEARROZ-ICA-CIAT 1986,

PNA-INIAP-CIAT 1987).
The average number of insecticide applications is 5.2 per crop in Colombia, 3.5 in Costa Rica
and 3.7 in the Dominican Republic. These applications are not only a matter of economic concern,
taking into account that 60-80% are preventive calendar-scheduled sprayings (Weber 1987) but
they are also a cause of considerable concern because of their ecological and toxicological effects.
Insecticide resistance, for example has been shown to occur in at least three rice insect species in
the Americas; Spodoptera frugiperda, Tagosodes orizicolus (Diego Trujillo 1988 unpublished) and
Lissorhoptrus oryzophilus (Bowling 1972). Resurgence of the rice planthopper T. orizicolus can
easily be induced by certain organophosphate or pyrethroid insecticides (ICA 1983 unpublished,
Weber 1986). Insecticide residues in grain oil and straw have been found in Brazil and Colombia
(ITA 1982, Ramos et al., 1986). Thus present pest control practices are not acceptable from an
economical, ecological or toxicological point of view.
Although information on pest biology, pest damage and tolerance of rice to insects has been
available in the Americas for the last two decades, only recently has an integrated pest management
concept been developed and implemented in close cooperation with several institutions in the
tropical zone. The following describes the development and implementation of rice IPM in Latin
America.
2. DEVELOPMENT OF THE IPM-CONCEPT

A short characterization of some basic IPM components is given. This is followed by a discussion of
the integration of these components for the management of some major and minor phytophagous
insects.
2.1 CHARACTERIZATION OF IPM COMPONENTS

2.1.1 General Plant and Crop Characteristics
Sixty two per cent of the favored upland and 72% of irrigated rice in Latin America is planted to
modern semidwarf varieties, whereas traditional varieties dominate under unfavored conditions
mainly in the cerrados of Brazil (CIAT 1985). Most of the modern varieties have three general
characteristics that increase their tolerance to insect attack:
• High initial vigor allows plants to quickly pass through the early seedling stage and thus
escape deleterious damage by leaf-mining (Hydrellia) and leaf-feeding (Spodoptera) insects;
additionally this assures good recovery from previous attack.
• High tillering increases crop plasticity and provides tolerance to attacks by certain early
seedling pests (Euetheola, Phyllophaga, Spodoptera) stem boring pests (Diatraea) and stem
sucking pests (Tibraca, Euschistus) .
• Medium duration varieties of 120-130 days assure a vegetative growing period of 60-65 days,
which again increases the plants capacity to recover from early damage.
WEBER AND PARADA 735
Almost all of the rice area in Latin America is direct seeded in dry or puddled soils. Only a
few countries like Ecuador, Peru and Dominican Republic have some transplanted rice
areas. Direct seeded rice crops generally have two critical periods:
The early seedling stage, when pests such as leaf miners (Hydrellia), leaf-feeders
(Spodoptera), plant cutting (Euetheola) and sucking insects (Tagosodes, Hortensia,
Draeculacephala) immigrate into the crop before beneficial arthropods have established
an ecological equilibrium.
The reproductive stage from booting to flowering, where the plant concentrates its
resources increasingly on panicle development and shows less capacity to tolerate or
recover from certain damage. However, well established biological control reduces the
risk of pest attack by leaf feeding (Mocis, Thioptera), leaf rolling (Syngamia, Marasmia),
leaf sucking (Tagosodes), grain sucking (Oebalus, Nezara) and stem boring (Diatraea,
Rupela) insects.
2.1.2 General Characteristics of Phytophagous Insects

Several key parameters are used for a general characterization of phytophagous insect biology,
ecology and epidemiology. They provide guidance in evaluating the insect’s damage potential and
in analyzing key aspects of its management (Table 1).
Ecological Strategy: All phytophagous insects are classified according to their general ecologi-
cal strategy on the r-K continuum. Plant- and leafhoppers and spider mites are rather typical
r-species with high capacity for population increase, wide density fluctuations and wide genetic
adaptability. The rice water weevil is a typical K-species with a rather low fecundity, high longevity
and an extreme adaptation to its special ecological niche. Whereas the management of r-type
species requires a reduction of short-term population fluctuations, breeding varieties that tolerate
the normal fluctuations and strengthening natural, density dependent antagonists, the K-type
species is managed by reducing populations below the economic threshold by altering environmen-
tal conditions to make them less favorable for their survival.
Damage in Plants: Most insects affect the plant directly through their feeding and additionally
cause indirect damage by transmitting plant pathogens and/or increasing the risk of higher weed
infestations. The attack by most foliage damaging pests during the seedling stage causes little direct
damage because the plant usually recovers from even a 100% loss of leaf area. However, the
reduced capacity of the crop to compete against weeds is an important indirect effect in upland and
irrigated rice under direct seeding conditions.
Field Colonization: Some pests are already in the field at planting time. Early season upland
soil pests including Euetheola, Phyllophaga or Neocutilla should be monitored and controlled
before planting. Most phytophagous insects in rice, however, immigrate into the field after planting
and their short-term importance depends on the size of the immigrating population. Stink bugs, for
example, cause immediate crop damage upon their arrival by sucking grains. Their reproduction
and population build-up occurs in graminaceous weeds rather than in rice and the major biological
control organisms attack the egg and nymphal stages. Thus the size of the immigrating population
TABLE 1
General characterization of some phytophagous insects in rice fields in Latin America
Insect species (common Ecological Crop damage Field colonization Biological control Damage Pest status
name) strategy potential
Direct Indirect Potential Stability
Tagosodes orizicolus r Phloem suck- Virus trans. Immigration + High Medium High Major
(rice planthopper) ing multiplication
Hydrellia spp. r-(K) Leaf mining Reduction of Immigration + Low Medium Medium Occasional
(whorl maggot) plant vigor oviposition (irrigated)
Spodoptera frugiperda r Leaf feeding Reduction of Immigration + High Low Medium Occasional
(fall armyworm) plant vigor oviposition
Oebalus spp. (r)-K Grain sucking Grain discol. Immigration Low Low Medium Occasional
(stink bug)
Euetheola spp. K Seedling pest Reduction of Present in soil or Low – Medium Occasional
(black beetle) crop density immigration (regional)
Phyllophaga spp. K Seedling pest Reduction of Present in soil Low – Medium Occasional
(white grubs) crop density (regional)
Syngamia spp. r-K Leaf rolling – Imigration + High High Medium Occasional
(leaf roller) oviposition
Elasmopalpus lignosellus r-K Seedling cut- Reduction of Immigration + Low – Medium Potential
(corn stalkborer ting crop density oviposition
Diatraea spp. r-K Dead hearts – Immigration + High Medium High Potential
(sugarcane borer) oviposition (regional)
Tibraca limbativentris r-K Dead hearts – Immigration Medium – High Potential
(stink bug) (regional)
Hortensia similis r Xylem suck- – Immigration + High High Low Potential
(leafhopper) ing oviposition
Panoquina spp. r-K Leaf rolling – Immigration + High High Low Potential
(leaf roller) oviposition
Lissorhoptrus spp. K Root cutting Fungal infec- Immigration + Low – High Potential
(water weevil) tions oviposition
Rupela albinella r-K Stem boring Lodging Immigration + High High Low Potential
(stem borer) oviposition
basically defines the damage in the rice crop and in managing this pest it is necessary to concentrate
on reducing the build-up in natural hosts and minimizing immigration into the rice fields. Most
lepidopterous pests (e.g. Spodoptera, Mocis, Thioptera, Syngamia and Diatraea) immigrate as adults
and only the late larval stages may affect the crop by their feeding activity. Biological and cultural
control mechanisms have time to reduce the population density during the egg and early larval
stages. Thus management of these pests requires the strengthening of mortality causing factors
during these early development stages.
Insects with short generation cycles (e.g. Tagosodes, Hortensia and Hydrellia ) may complete
several generations within the same crop. Their field colonization is by immigration and subsequent
multiplication and their management requires a strengthening of all factors that regulate pest
densities in a density-dependent manner and preventing a population build-up with special atten-
tion to the risk of inducing resurgence.
Biological Control: A characterization of the effectiveness of biological control for every pest
in the rice field requires the evaluation of two basic aspects; the potential of existing biocontrol
agents to control and regulate the pest population and their stability, that is their capacity to reduce
density fluctuations in the medium term. Most phytophagous insects in rice fields in Latin America
have a high potential of biological control with a medium to high stability. However, some soil
inhabiting pests (e.g. Euetheola, Phyllophaga and Lissorhoptrus) have a rather low potential of
biological control under most conditions. The increase of this potential has to be a long term
research goal. High potential of biocontrol by itself does not automatically assure high stability as
in the case of Spodoptera frugiperda, where the low stability of biological control, closely linked to
the strategy of the insects to ‘escape’ by migrating, is the major factor contributing to pest densities
that cause plant damage.
The above mentioned characteristics provide a general description of the damage potential of
the major rice insects. In Tagosodes, Diatraea, Tibraca and Lissorhoptrus the damage potential is
especially high because of the capacity for extreme population fluctuations and/or their severe
damage to rice plants. Only Tagosodes can presently be considered as a widespread major pest.
Diatraea, Tibraca and Lissorhoptrus have regional importance as major pests but are potential pests
all over Latin America (Table 1).
2.1.3 Varietal Characteristics
Breeding for insect resistance is only part of the overall strategy for developing commercially
acceptable rice varieties. Breeding efforts require the prior identification and availability of
resistant parents, an adequate level of resistance and efficient screening methods. Genetic resis-
tance to rice insects in Latin America has been identified for at least Tagosodes orizicolus, Hydrellia
griseola, Diatraea saccharalis, Lissorhoptrus oryzophilus, Syngamia sp., Spodoptera frugiperda, Elas-
mopalpus lignosellus, Chaetocnema sp., and spider mites. Analyzing the efforts required for breed-
ing resistant varieties and comparing them with the availability of alternative control measures, has
justified germplasm screening for T. orizicolus, H. griseola, D. saccharalis, and L. oyzophilus. In the
case of leaf rollers, S. frugiperda, Chaetocnema and spider mites the selection of highly susceptible
varieties is avoided on the basis of field observations.
For every insect a certain type of resistance is desired (Table 2). In the case of the T. orizicolus,
germplasm with tolerance to mechanical damage is selected, and all lines that possess some medium
to high degree of antibiosis against the insect are discarded. This concept of selecting only tolerant
material is based on greenhouse experiments that show a high probability of selecting new biotypes
in a relatively short time by growing varieties with antibiosis. Seventy six per cent of the presently
grown varieties in Latin America and more than 90% of the varieties in the tropical zone are
tolerant to the planthopper, and no biotype outbreaks are known although some of these varieties
have been grown on a large scale for more than 10 years. Differences in general aggressiveness
seem to exist. Present varieties tolerate the natural pest fluctuations and allow the beneficial
arthropods to regulate population density below economic thresholds. Field results from 20
commercial fields all over Colombia indicate that no insecticide applications were required for T.
orizicolus in the tolerant varieties. However, maximum insect densities passed the economic
threshold for susceptible varieties in 35% of the fields. Thus, because of varietal tolerance direct
damage by T. orizicolus was insignificant and biological control was allowed to regulate population
densities below the thresholds (Fig. 1).
TABLE 2
Type of resistance desired in an intergated pest management approach for rice in Latin America
and present status of resistance in the 54 major commercial varieties.
Insect species Type of resistance Modern commercial varieties

resistant (%)
Tagosodes orizicolus Tolerance to mechanical damage 76
Hydrellia spp. Intermediate resistance 46
Diatraea spp. Antibiosis and tolerance 87
Lissorhoptrus spp. ? -
Syngamia sp. Prevent high susceptibility 921
1
Data for only 13 of the commercial varieties.
Hydrellia leaf mining attacks are restricted to irrigated rice with major attacks during the rainy
season. These attacks can destroy direct seeded fields if they occur during the 1-3 leaf stage. About
46% of the present varieties have intermediate resistance against the leaf miner and general
germplasm screening started in 1986 in order to increase the number of resistant varieties. The type
of resistance widely available in the germplasm bank is a combination of intermediate resistance to
larval attack and good recovery of damaged plants.
Diatraea is presently not considered as a pest in most areas of Latin America because 87% of
the widely grown varieties are resistant. However, some varieties and parents from unfavored
upland rice such as IAC 165, IRAT 124 and TOX 1011-4-1 are highly susceptible to this stem borer.
Therefore breeding lines having a Diatraea susceptible parent have to be screened in order to
prevent the release of Diatraea susceptible varieties and Diatraea thus becoming a new insect
problem in irrigated or favored upland rice. Antibiosis against larval feeding in the leaf sheath
before penetrating the stem and tolerance to attack by high tillering are the major mechanisms of
resistance.
Fig. 1. Maximum number of Tagosodes orizicolus encountered in 20 commercial fields all over Colombia in
relation to action thresholds (A.T.) for susceptible and tolerant varieties in the seedling stage (A.T.I.) and
advanced growing stage (A.T.II) (Source: FEDEARROZ 1987).
The rice water weevils, Lissorhoptrus, Onychilus, Oryzophagus, Helodytes and other genera are
increasing in importance regionally. Germplasm evaluation for resistance is still in the early
research stages.
2.1.4 Agronomic Practices

Many agronomic practices that have favorable or adverse effects on certain phytophagous insects
are available (Table 3). In some insects the use of adequate crop management can reduce their
economic importance to insignificant levels. The integration of pest management into an overall
concept of crop management must be carried out on a regional level.
Most phytophagous insects on rice in Latin America are not monophagous on this host plant
and may reach higher populations on other host plants. If the sequence of rotated crops is not
planned adequately, crop rotation may build up potential pests like Diatraea, Blissus or Nezara to
economically damaging populations in maize or soybeans. These pests then migrate into rice.
2.1.5 Chemical Control

The use of insecticides that are most compatible with other control tactics is one of the recom-
mended short-term control measures for rice pest management. Data from the European OILB-
research group on the side-effects of pesticides on beneficial arthropods is incorporated into the
IPM-field guide, although more adequate data on selectivity of pesticides against the major benefi-
cial arthropods in rice fields in Latin America would be desirable (Hassan 1984). The use of
resurgence-inducing insecticides is discouraged.
TABLE 3
Crop management practices and their effects on insect pests in rice.
Effect on insect species

Crop management practice Increasing Decreasing
Land preparation at beginning of dry season – Euetheola
Fine grained seedbed Phyllophaga
High seeding density Neocutilla
High water level Elasmopalpus Spodoptera
Agrotis
Spodoptera
Hortensia
Hydrellia Spodoptera
Lissorhoptrus Euetheola
Ochetina Neocutilla
Elasmopalpus
Phyllophaga
Chaetocnema
Poor land levelling Hydrellia
Lissorhoptrus
Ochetina
Deficient homogeneity in seeding density Hydrellia
High nitrogen fertilization Syngamia
Marasmia
Hortensia
Draeculacephala
Deficient control of weeds –
Echinochloa –
Setaria Oebalus –
Brachiaria Mocis –
Aeneolamia –
Zulia –
Deois –
Andropogon Sipha
Mocis
Crop rotation Diatraea
Rice-rice Tagosodes Blissus
Hydrellia Nezara
Rupela Tagosodes
Rice-maize Spodoptera Hydrellia
Diatraea Rupela
Blissus Nezara
Elasmopalpus Tagosodes
Rice-soybean Nezara Hydrellia
Acrosternum Rupela
Diabrotica Diatraea
Blissus
Phyllophaga
TABLE 4
Integration of IPM components to manage some insect pests of rice in Latin America.
Insect species Adequate variety Crop management Biological control Chemical control
Tagosodes Tolerant to insect, – Protect beneficials Do not cause resurgence
resistant to hoja blanca ++ +
++
Hydrellia Intermediate resistance Homogenous seeding den- Protect beneficials Economic threshold
++ sity, adequate levelling +
++
Spodoptera High initial vigor Excellent seedbed prepara- Protect stability of Water management
tion, adequate seeding den- biological control ++
sity, water management +
Euetheola and Phyllophaga High initial vigor, high tillering Land preparation at begin- – Economic threshold
ning of larval infestations; ++
crop rotation with legumes
++
Oebalus – Efficient weed control – Economic threshold
+ ++
Syngamia Prevent high susceptibility Adequate fertilization with Protect beneficials Economic threshold
N ++
++
Diatraea Resistant Adequate crop rotation Protect beneficials
++ ++
+ + Essential component, + major component, – complementing component.
2.2 INTEGRATION OF COMPONENTS
The integration of the control tactics into an overall pest management concept is being done on a
regional basis. For the tropical zone, examples of integrated management programs for some pests
are shown in Table 4.
The risk of economic damage from T. orizicolus and hoja blanca virus was high until the 1970s,
when traditional or improved varieties susceptible to the insect and to the virus predominated.
With the introduction of T. orizicolus tolerant varieties in the 1970s, the risk of mechanical damage
from hopperburn decreased. The additional incorporation of an intermediate level of resistance
against the virus in the 1980s has reduced the risk of virus outbreaks. The development and
introduction of integrated crop management including insect monitoring and economic thresholds
since 1985 has reduced the probability of insect outbreaks to extremely low levels mainly by
reducing the number of insecticide applications and by preventing the use of resurgence-causing
insecticides. Also, the introduction of virus-resistant varieties together with a predictive system of
vector-percentages on a regional level in 1989-1990 reduced the risk of virus outbreak to low
probabilities (Table 5).
TABLE 5
Consecutive steps to an intergrated management of Tagosodes, hoja blanca, and Hydrellia
Tagosodes
Varietal resistance Crop management Probability of damage
Time scale Tagosodes Hoja blanca Tagosodes Hoja blanca
Before 1968 Susceptible Susceptible Conventional High High
1970-80 Tolerant Susceptible Conventional Medium High
1980-85 Tolerant Intermediate Conventional Medium Medium
1985-90 Tolerant Intermediate Adequate Low Medium
After 1990 Tolerant Resistant Adequate + Low Low
predictive system
Hydrellia
Varietal resistance Crop management Probability of damage
Susceptible Conventional High
Susceptible Adequate Medium
Intermediate Adequate Medium-Low
Intermediate Adequate + early egg Low
monitoring
Routine evaluation of germplasm for Hydrellia resistance started in 1986, although many earlier
released varieties have moderate resistance similar to IR 40. Combining adequate crop manage-
ment practices like homogeneous seeding densities and adequate field levelling with moderately
resistant varieties reduces the risk of Hydrellia damage to medium-low probabilities. In order to
prevent destructive attacks of very early immigrating Hydrellia in the 1-2 Ieaf stage, field monitoring
including an evaluation of the percentage of plants with eggs is required (Table 6).
Similar recommendations are incorporated into the IPM manual on the basis of present
knowledge to provide control in the short term and to manage populations in the medium to long
term. For most secondary pests field monitoring and use of economic thresholds has shown that
these insects are of very low economic importance.
3. IMPLEMENTATION OF IPM IN COLOMBIA
3.1 PROCESS OF IMPLEMENTATION

The Colombian National Federation of Rice Growers (FEDEARROZ), the National Research
Institute (ICA) and CIAT in 1986 initiated a cooperative effort to design and implement an
integrated pest management system for rice farmers in Colombia. After an analysis of current pest
control practices, pests and recommendable alternatives (FEDEARROZ-ICA-CIAT 1986), an
IPM field guide was designed and tested by local field technicians in confirmation trials in commer-
cial fields throughout Colombia. Commercial fields of 5-10 ha size were divided in two parts, one
managed conventionally according to the farmer’s traditional practices and the other part accord-
ing to the IPM field guide.
Based on analysis of the results of the confirmation trials the original IPM-field guide was
modified to suit more closely the needs and problems in the different regions of Colombia. The
addition of risk thresholds allowed a reduction in the number of required field visits from 13 to 7,
with visits focussing on the critical crop stages. The field guide consists of three parts:
Field Evaluation: There is an evaluation sheet for seven different crop stages indicating the
required samples. For most insects indirect sampling methods are required. This involves evaluat-
ing the number of leaves damaged by defoliating, mining, rasping or rolling insects and tillers
damaged by stem cutting or boring insects. The farmer is not expected to identify the insect species
but rather has to recognize the different types of damage. Only planthoppers, leafhoppers and
beneficial spiders have to be collected with the insect net and counted and stink bugs are evaluated
during flowering by counting the number of insects per m2 .
Decision: Critical thresholds are indicated for all pests using a sequential threshold for optimal
sample size, a risk threshold for intensity of sampling and an action threshold for the need of control
(Table 6).
Action: If pest attack passes the action threshold, the farmer has to identify the adequate action
to control the insect on the short term and the adequate management to prevent similar attacks in
the future. The action part, therefore, includes tables indicating pest characterization, control and
management methods on the short and medium term, a characterization of pesticides including
their effect on beneficials and a characterization of varieties.
TABLE 6
Integrated pest management of rice in Colombia. Decisions according to critical thresholds.
Sample Evaluation Thresholds according to crop stage 0 to 3
A. SOIL INSECTS
Sample: 12 × 1 m No. of white grubs in 3 m 2 Risk threshold: More than 30 larvae at beginning of dry season
lineal 0.20 × 0.25 cm = 3 m 2
No. of black beetles in 3 m2 Action threshold More than 5 adults before planting
No. mole crickets in 3 m2 Action threshold: More than 3 mole crickets before planting;
Emergence-seedling 0–1 Seedling-tillering 1–2 Tillering-elongation 2–3
B. INSECTS IN FOLIAGE ST RT AT ST RT AT ST RT AT
Sample: 5 times 10 tillers No. damaged tillers in 5 × 10 tillers 2 2–5 6 2 2–5 6 4 5–7
Sample: 5 times 10 tillers No. leaves with eggs of Hydrellia 50

with 3 youngest leaves
No. leaves damaged by Hydrellia 30 60 60
No. leaves damaged by leaf feeders 15 15–30 30 30 30–60 60 60
No. leaves damaged by leaf rollers 25 25

% leaf area affected by rasping 20 20–30 30 20 20–30 30 30
C. SWEEP NET
Sample: 3 times 10 double No. approx. of planthoppers 200 400 400

sweeps
No. approx. of leafhoppers 250 500 500
(contd.)
TABLE 6 (contd.)
Sample Evaluation Thresholds according to crop stage 3 to 7
Elongation-booting 3—4 Booting-Pan.emerg 4—5 Pan.emere-Flowerine 5—7
B. INSECTS IN FOLIAGE ST RT AT ST RT AT ST RT AT
Sample: 5 times 10 tillers No. damaged tillers in 5 × 10 tillers 4 5-7 8 2 – 3
Sample: 5 times 10 tillers No. leaves damaged by leaf feeders 40

with 3 youngest leaves
No. leaves damaged by leaf rollers 25
Sample: 10 times 10 tillers No. leaves damaged by leaf feeder 4 4-12 12 4 12

with flag leaf
No. leaves damaged by leaf roller 4 4-12 12 4 12
C. SWEEP NET
Sample: 3 times 10 double No. approx. of planthoppers 400 400
sweeps
No. approx. of leafhoppers 500 500
D. PANICLE PESTS
Sample: 10 times 1 m 2 No. of stink bugs per 10 m 2 40 40
ST = Sequential threshold – If attack passes over this level, take two more samples.
RT = Risk threshold – If attack passes over this level, take another sample in 3-5 days.
AT = Action threshold – If attack passes over this level, take control action.
3.2 FIELD RESULTS FROM COLOMBIA

Managing 20 commercial fields in Colombia during 1986–87 according to the IPM field guide
required a control action in 10 cases (FEDEARROZ 1987). Hydrellia leaf mining was the most
common problem, because the predominating variety, Oryzica 1, is highly susceptible. Dead hearts
and leaf rasping mainly caused by stem sucking pentatomids and flea beetles respectively, passed
action thresholds in 10% of the upland fields. Spodoptera and Mock caused leaf feeding damage in
seedlings and flag leaves, respectively, surpassing thresholds in 5% of the fields (Table 7).
TABLE 7
Percentage of 20 commercial fields located throughout Colombia,
where insect attacks passed economic thresholds
Insect/attack Crop stage Fields passing threshold (%)

White grubs in soil Before planting 0
Black beetles in soil Before planting 0
Mole crickets Before planting 5
Cut seedlings Seedling stage 0
Eggs of Hydrellia Seedling stage 0
Leaf mining Seedling stage 15
Leaf rasping Seedling stage 10
Dead hearts Tillering-booting 10
Leaf feeding Tillering-booting 5
Leaf rolling Tillering-booting 0
Flag leaf feeding Booting-flowering 5
Flag leaf rolling Booting-flowering 0
White panicles Flowering 0
Stink bugs Flowering 0
Source: FEDEARROZ 1987
An average of 0.5 applications per crop was required when using IPM against the countrywide
average of 5.2 applications of insecticides in conventionally managed fields in 1985. Applications
and insect control costs were reduced by more than 90%, permitting savings in production costs of
80 to 120 US $/ha.
Field IPM demonstrations are part of a general approach to improved crop management in
Colombia, including the demonstration of adequate weed control, disease management and seeding
practices. Total savings from integrated crop management are between 170 and 225 US $/ha or an
equivalent of 0.8–0.9 tons of paddy per ha (Table 8). These possible savings and the increase in
profitability attract the farmers’ attention and guarantee a high level of success in transferring the
integrated crop management concept as a general alternative to present practices.
4. CONCLUSIONS
Rice is a relatively young crop in the Americas and only a few insects can be mentioned as major or
potentially destructive pests. This is in clear contrast to the current practice of 2–6 preventive
insecticide applications in rice all over the tropical zone, Central America and the Caribbean.
TABLE 8
Present production costs in kg of paddy rice/ha for planting, weed, insect and disease control compared to costs when
following an integrated crop management system (ICMS) in Colombia.
Practice Central Zone Coastal Zone Llanos

Present ICMS Savings Present ICMS Savings Present ICMS Savings
Planting 513 308 205 513 308 205 513 308 205
Weed control 474 263 211 395 263 132 395 263 132
Insect control 407 115 292 256 29 227 289 73 216
Disease control 322 89 233 356 97 259 576 245 331
Total 1716 775 941 1,520 697 823 1,773 889 884
Source: FEDEARROZ-ICA-CIAT, 1986.
An adequate characterization of the plant and crop development and the ecology of the
phytophagous insects as well as the possible control methods is the first step in developing an IPM
concept. In order to assure stability and reliability of pest management no single control method
should be used. Therefore, antibiosis resistance is avoided in the highly adaptive rice planthopper,
T. orizicolus, preferring instead the combination of varietal tolerance to the mechanical damage of
the insect, varietal resistance against the virus and adequate protection of beneficials to prevent pest
resurgence.
Field results from Colombia show large potential savings in combining of IPM with other
adequate crop management practices. Therefore, the incorporation of IPM into an overall crop
management approach that demonstrates to farmers a considerable increase in crop productivity
and profitability is essential in order to attract farmers’ attention.
Similar efforts in other countries such as Ecuador also show considerable savings by using
adequate crop management practices. The urgent task of demonstrating the use of IPM in the
farmers’ fields lies ahead for rice IPM proponents in Latin American countries.
REFERENCES
Bowling, C.C. 1972. Status of rice water weevil resistance to aldrin in Texas. J. Econ. Entomol. 65: 1490
Centro Internacional de Agricultura Tropical (CIAT). 1985. Situacion actual de la prcduccionde arroz en America Latina
y el Caribe. Informe de la Sexta Conferencia Internacional del Arroz para America Latina y el Caribe, CIAT, p.
137-168.
Food and Agriculture Organization (FAO). 1986. Production Yearbook. Rome.
FEDEARROZ. 1987. In Resultados deal Manejo Integrado de Plagas. Investigation Arroz 1986-1987, Fedearroz, Bogota,
Colombia pp. 283-319.
FEDARROZ-ICA-CIAT. 1986. Resultados del Plan Nacional para majorar el manejo del cultivo del Arroz en Colombia.
Documento de trabajo. 152.
Hassan, S.A., 1984. Testing the side effects of pesticides on beneficial arthropods. In Judicious and Efficient Use of
Insecticides on Rice. International Rice Research Institute, Los Baños, Philippines,
ITA. 1982. Contaminacao de altimentos processados. Instituto de Tecnologia de Alimentos. Campinas, Brazil.
PNA-INIAP-CIAT. 1987. Plan Nacional para majorar la situacion del arroz en Ecuador. Documento de trabajo, 247 p.
Pulver, E. 1985. Costos de produccion en America Latina y el Caribe; una guia para identificar problemas de produccion.
In lnfome de la Sexta Conferencia Intemacionol de Arroz para America latina y el Caribe. Centro Internacional de
Agricultura Tropical, Cali, Colombia, pp. 105-116.
Ramos, C.E., R. Rodriguez, B. Mejia and J. Torres. 1986. Intoxicacion asintornatica por pesticidas organofosforados en
bovinos alimentados con socos de arroz (Oryza sativa L.). Revista Association Colombiana de Medicos Veterinarios y
Zootecnistas (Bogota, Colombia) (37): 12-16.
Weber, G . 1986. Manejo integrado de plagas. Un ahorro y una inversion. Arroz en las Americas 7(2): 1-5.
Weber, G. 1987. Approach Io intergrated pest management in rice in Latin America. International Conference on Plant
Protection in the Tropics, Manila, Philippines, 1987.
Subject Index
AC 1423 532 Antibiosis 533-534, 539-540, 689, 138, 748

Acephate 521-522, 668 Antixenosis 533-534, 536, 689
Acoustic behavior 414, 424 Ants 191, 367, 554, 560, 567, 592
Action threshold 689, 738-739, 743-748 Aphids 81, 83, 365, 367, 462-463
Aerial sprays 723-724 Applaud (see Buprofezin)
Afghanistan 391, 400, 408, 427, 430, 432, 442 Apple grain aphid 463
Africa 3, 8, 12, 38, 68, 76, 85, 99, 101, 108, 178-179, 182, Aquatic 386
240, 242, 246, 270, 272, 280, 364, 376-377, 388-389, Aquatic pests 168, 557-559, 571
398, 409, 411, 413, 415, 460, 462-463, 488, 492-493, Argentina 366, 397, 428, 437
496, 504, 529
Arkansas 391
African pink borer 389, 405, 496, 527
Armyworm 408, 445, 495, 527, 550-555, 560-564,
African rice borer 389
567-569, 571, 634, 640, 643, 669-670, 689, 700, 702
African rice gall midge 272, 375
Artificial diet 641, 645
African rice hispa 459
ASD 7 520, 522, 530, 684
African striped stem borer 527
ASD11 662
African white stem borer 527
Asia 8, 10, 12, 242-243, 363-364, 388-389, 394, 413, 415,
Agroecosystem 498, 607-608, 614
420, 425, 427, 432, 454, 457, 463, 488-489, 495-496,
Akita prefecture 723-724 498-499, 504, 529-531, 561, 600, 613-614, 636-637,
Aldehydes 536 658, 681, 694, 696-697, 701-702, 705-706
Alkanes 536 Asian rice gall midge 372, 532
All India Coordinated Rice Improvement Project Asiatic rice borer 389
(AICRIP) 532 Assam 704
Allelochemics 536, 561, 690 Assam Rice Collection (ARC) 532
Allomones 536 Asynchronous planting 496-499, 501, 504-510, 564-566,
Allopatric populations 537-538 607
Almond moth 640 Augmentation of natural enemies 586-587
American rice stalk borer 391-392 Australia 3, 76, 368, 388-389, 403, 409, 413, 420, 422,
American rice stem borer 389 424, 427, 432, 435, 442, 449-450, 463, 488, 490, 570
American sugarcane borer 389 Azinphos ethyl 671, 697-698
Amino acids 369, 534-535, 561, 644 Azolla 456, 559
Andhra Pradesh 532, 538
Angola 405, 419-420, 459 Bacteria 427, 433, 463, 524, 614, 630
Angoumois grain moth 527 Bacterial leaf blight 541-542, 558
Antarctica 3, 488 Baculoviruses 642
750 SUBJECT INDEX
Bangladesh 368-370, 372, 379, 390, 394, 400-401, 403, Bund plastering 550
407, 414, 420, 422, 427, 430, 432, 435, 437, 439-443, Bund shaving 550, 554, 571
449-450, 454, 457, 463, 525, 532, 537, 590, 600, 658
Buprofezin 659-660, 705-706, 722
Barley 366, 421, 445, 453, 463, 562, 716-717
Burkina Faso 375-377, 396, 405
Barnyard grass 469
Burma 3, 368-369, 372, 389-390, 392, 394, 398, 400,
Barriers 550, 554
407-408, 411, 414, 420, 422, 427, 430, 432, 437, 439,
Basmati 444 442, 449-450, 457, 525
Beating 554
Burning 550-551
Benefit-risk ratio 674
Burrower bugs 97
Benin 376-377, 396
Burundi 405, 409
Berseem fodder 562
BG276-5 532
CAB International Institute of Entomology 600
BG400-1 532
BHC 658-659, 665, 669, 698, 702, 715, 722-723 Cabbage 447, 449, 634
Bhutan 372, 390, 394, 400, 407-408, 414, 420, 422, 427, California 372, 386, 445, 453-454
430, 432, 437, 439, 442, 449-450, 457, 638 Cameroon 375-377 ,396, 405, 459
Bihar 538 Canada 444, 463
Biological control 10, 517-521, 524, 543, 560, 585-608, Cannibalism 365
613-646, 673, 681, 688-690, 699-701, 703, 736-737, Carbamates 658, 667-668, 696-697, 704-705, 716
741
Carbaryl 660, 666, 668, 698, 705
Biotechnology 540-542, 614
Carbofuran 521-522, 659-661, 665, 668-669, 671-672,
Biotype 375, 423, 425, 522, 529-530, 537-540, 543, 572,
682, 684, 697-698, 704-705
661, 673, 682-684, 689-690, 703-705, 738
Carbonyl compounds 536
Bird cherry oat aphid 463
Carbosulfan 659, 684
Bird damage 542
Carboxylesterase activity 668
Black beetles 380-381, 736, 744, 748
Black bug 369-370, 527, 551-552, 555, 558, 560, 562, 564, Caribbean 530, 748
568, 587, 618-619, 623, 626, 629, 632-633, 715, 727 Caribbean Islands 428
Black-streaked dwarf virus 421, 530 Cartap 723
Blast disease 541-542, 726 Caseworm 9, 63, 437-439, 495-496, 504, 506, 527,
Bloodworm 527 541-542, 552-553, 555, 558-559, 566, 568, 570-571,
Blue beetle 460 669, 689
Blue leafhopper 526 Castor 449
Bonfires 555 Caterpillars 555, 558, 564, 566, 568, 571, 614, 616
Boom and bust cycle 703 CB 1 531
Botanical insecticides 660 CB 2 531
BPMC 660, 665, 668, 684, 689, 697, 705 Central America 366, 389, 397, 428, 444, 447, 530, 748
Brachypterous 410, 421, 423, 427-429 Central Luzon 702
Brazil 5, 383, 389, 405, 428-429, 437, 468, 618, 621, 637, Central Rice Research Institute (CRRI) 532
734
Centro Internacional de Agricultura Tropical (CIAT)
Brownbeetles 136 530, 743
Brown planthopper 9-10, 364, 422-425, 427, 493, 495,
Cereal thrips 470
497-501, 503-506, 518-520, 522, 526, 529-530,
535-538, 540-542, 549, 555, 559, 561-562, 566, 572, Cereals 447, 493-494
585, 596-603, 606, 616, 620-621, 623-628, 630-632, Chad 410
637, 645-646, 658-667, 671-673, 681, 685-686, 688, Chafer beetles 380-381
693-707, 717 Chemical control 10, 517-518, 524, 560, 657-673,
Brown semilooper 643 681-684, 688, 690, 696, 699, 706, 718, 722-723, 726,
Brunei 407, 432, 435 739, 741
Bufencarb 660 Chilo iridescent virus (CIV) 643, 645
SUBJECT INDEX 751
China 5–6, 8, 215, 251, 370, 372, 374, 390–391, 394, 398, Critical thresholds 743–745
400, 402, 407–409, 411, 414, 420–422, 425, 427–428, Crop
430–432, 434, 437–444, 446, 449–450, 452, 457, 461, area 550, 562
463, 488, 524–525, 530, 538, 551–552, 556, 563, background 550, 559
566–567, 586–587, 621, 625, 635, 637, 641, 657, 661,
diversification 556
668–669, 673, 681, 684, 688–689, 695, 702, 705–706,
intensification 9, 498–500, 502–504, 507–509, 562,
717
572
Chinaberry 660
rotation 550, 561, 572, 690, 701, 739–741
Chlordimeform 659, 671, 723
sanitation 687
Chlowpha 659, 670–671, 697–698
Cross resistance 529, 658, 666, 668
Ciba-Geigy 697
Crushing insects 550–551, 554
CICA4 530
Cuba 366, 388, 429, 468, 633
CICA6 530
Cultivars
Classical biological control 586–588
commercial 524
Climate 488, 524, 662
obsolete 524
Cloning 542
special purpose 524
Clover 562, 634
Cultural control methods 10, 383, 459, 487, 490–491,
CO 15 531
496, 510, 517–518, 520, 524, 543, 549–551, 556–572,
CO 18 531–534 673–674, 681, 689–690, 699–700, 715, 718–719, 737,
CO 21 531–534 739–742
CO 42 662 Cutworm 72, 551, 634, 646, 702
Coconut beetle 616 Cyanofenphos 665
Coconut moth 588 Cypermethrin 666, 671
Coleoptera (key to species) 135–136, 138–182 Cyprus 409
Colombia 404, 428, 733–734, 743, 748, 748 Cytoplasmic polyhedrosis virus (CPV) 643
Colombian National Federation of Rice Growers
(FEDEARROZ) 743 Dahomey (see also Benin) 376–377
Colorado potato beetle 625 Damselflies 671
Common armyworm 444 Dark-headed stem borer 9–10, 389, 392, 494
Common cutworm 449, 643 DDT 658
Comoro Islands 391 Deadhearts, stem Sorer 370, 377, 390, 393, 398, 552–554,
Congo 405, 419–420 556, 559, 641, 669, 723, 736, 748
Consewation of natural enemies 703, 705, 741 Deepwater rice 373, 400, 403, 439, 453, 493, 567, 590
Continuous cropping 562–563 Deltamethrin 661, 666, 671, 683, 689
Control tactics Density-dependent 503, 505
integration of 606–607, 641, 673–674, 681–690, Dermaptera (key to species) 128, 130
695, 103, 705, 739–742 Diazinon 658, 661, 665–666, 668, 671–672, 698, 705, 723
Control threshold 719–728 Differential cultivars 538
Corn 445, 449, 625, 641 Diflubenzuron 706
Corn semilooper 643 Dimilin (see Diflubenzuron)
Corn stalkborer 736 Diptera (key to species) 14, 17, 19, 20–38
Costa Rica 405, 734 Direct sowing 557, 590, 594, 735, 738
Cotton 447, 557, 586, 641, 695 Diseases 363, 412–414, 419, 421, 423, 427, 463, 495, 511,
Cotton bollworm 642 524, 532, 557, 571, 614, 682, 715–716, 722, 748–747
CPMC 668 DNJ 45 532
CR55 532 Dominican Republic 468–470, 734
CR 56 532 Donor parents 543
CR 94-13 533 Double cropping 9, 392, 425, 489–491, 494, 497–498,
Crickets 38, 365, 599, 701 502–505, 507–510, 563–564, 572, 607, 700–701
752 SUBJECT INDEX
Dragging 554 Ethoprophos 670

Dragonflies 130 Ethylan 521-522, 660, 662, 689
Draining fields 461,557,560,571 Ethylparathion 672
Drought 369, 376, 380, 407, 431, 452, 542, 550, 565 Ethylthiometon 723
Dryland rice 556-557, 559-560, 567,571 Europe 8, 68, 243, 364, 371, 421, 430, 444, 463
Dry season 374, 489-490, 492, \494, 502-503, 507, 562 European Community (EC) 713
Durable resistance 539 European corn borer 625, 635
Dust formulations 663, 671 Exclusion cage
DV12 532 for predation studies 604
Dwarf virus 413-44, 420 Exotic parasitoids 587, 607
Early-maturing rice cultivars 563-564, 572, 684, 686, 716 Fall armyworm 447-448, 570, 736
East Asia 205, 221, 242, 245,251, 299, 307, 446, 493, 531 Fallow period 495-498, 504, 506, 562, 564, 567-568, 572,
Ecological guild 493 701, 717
Ecological strategy 735-736 Fatty acids 536
Economic injury level 522, 719-720, 727-729 Feeding deterrent 533
Economic threshold level 602, 614, 658, 661-662, 667, Feeding preference 597
669, 671, 697, 718, 728, 741-743, 748 Feeding rates 561, 598, 603
Ecotypes 396 Fenitmthion 665-666 668
Ecuador 405, 733, 748 Fenthion 665-666
Egypt 366, 372, 419, 452, 531 Fenvalerate 665-666, 668
Elcar 642 Fertilizer management 549, 561, 687
Electromagnetic energy 549 Fertilizers 8-9, 403, 433, 435, 444, 549-550, 560-561, 569,
Electronic monitoring of insect feeding (EMIF) 533 571, 694-695, 697,700, 715, 740-741
Electrostatic sprayers 663 Field crickets 364
Elevation 524 Field geometry 550, 556, 571
Endosulfan 671, 689, 698 Field resistance 536, 540
Entomogenous bacteria 638-642 Fiji 366, 422, 435-436, 450, 463, 586-588
Entomogenous fungi 616-638 Fish 671
Entomogenous nematodes 645-646 Flea beetles 179, 748
Entomogenous viruses 642-645 Flooding 365, 550, 553-554, 557, 560, 567, 571-572
Entomopathogens Flooding stubble 572
biotechnology use. 614 Foliar sprays 662-664, 670, 673, 688, 698, 723
description of 614, 616-621, 624-633, 635-640, Forecasting
642-645 network 718
literature of 614, 632-633, 639 techniques 601
manipulation of 614 France 452, 635, 641
mycological terminology of 614-415 French Guiana 405
practical application of 614, 617-618, 621, 623-643, Fukuoka prefecture 726
645-646 Fungi 399, 427, 433, 463, 524, 614, 616-638, 690, 736
production of 621-623
Entomophthoralean fungi 671-672 Gain threshold 719, 727
Entomophthorales 617, 636-638 Gall formation 373-375, 570
Environmental modification 488 Gall midge 364, 373-376, 489, 495, 518, 527, 532-533,
Environmental resistance 497 538-539, 541-542, 553-561, 563-566, 568, 670-471,
Epizootics 614, 616-617, 621, 632, 634, 636, 638 682, 686
Esters 536 Gambia 405,437
Eswarakora 532-533 GEB24 532
Ethiopia 389, 419-420, 459 Gene deployment strategies 539,541
SUBJECT INDEX 153
Gene pyramiding 539 Guinea-Bissau 375

Gene rotation 540, 704 Guyana 383, 405, 447–448
Genetic diversity 537, 539, 543
Genetic engineering 542 Hainan Islands 402
Genetic maps 542 Haiti 468
Genetic transformations 542, 690 Handpicking of insects 550–551, 570
Geographical deployment 540
Harvest method 550, 569
Georgia 392
Hawaii 394, 450–451, 586–587
Germany 452
HCH (see BHC)
Germplasm
Hemiptera (key to species) 78, 80–129, 132
bank 539
collection 524, 526 Hesperiid 209, 250
evaluation 526–529, 737–739, 742 Hispa 457–459, 527, 552, 554, 558, 560, 568, 571
preservation 524 Hoechst 697
storage 525 Hoja blanca virus 429, 530, 741–742
Ghana 248, 375–377, 405, 410, 419, 437, 638 Hokkaido district 718, 721
Giallume 463 Hokuriku district 721
Glutathione -S transferase 668 Honduras 405
Gold–fringed rice borer 389–390 Honeydew 422, 425, 427, 429, 435, 443, 534
Gondwanaland 3–4 Hong Kong 390, 400, 407, 414
Granular insecticides 600, 659, 664, 670, 672, 688, 698, Honshu 718
704, 718
Hopperburn 364, 370, 423–424, 427, 627, 702, 742
Granulosis viruses 643–644
Horizontal resistance 539, 689
Grape colaspis 382–383, 561–562, 568
Host plant resistance 517–543,681-690
Grapes 382
Gm 366–367, 375, 377, 379, 383–384, 386, 390–391, genes for 526–530, 537–539, 696–697, 699, 702–703
401, 408, 413, 417–418, 421, 427–428, 431, 437, 441, mechanism of 526, 528, 533–537, 738
444–447, 449, 451–452, 461–463, 466, 468, 489, 493, use in IPM 519–524, 533–537, 539, 543, 699, 703–705,
496, 562, 567–568 729
Grass webworm 63, 643 Use of 487, 517–543, 737, 742
Grasshopper 31, 38, 46, 245, 408–412, 551–552, 554–556, variable 537–538, 703
560, 564, 567–568, 591, 638 Host plants
Grassy stunt 412, 423, 541, 660, 688, 702
alternate 367–368, 370–371, 378, 382–383, 386,
Grazing livestock 550, 570 390–392, 394, 396, 398, 401, 406, 409,
Greater wax moth 645 411–412, 415, 417, 420–422, 428, 432, 437,
Green hairy caterpillar 447, 626, 634–635, 640, 700 439–440, 446–447, 450, 460, 464, 470,
Green leafhopper 413–415, 495, 500, 504, 506, 508, 518, 493–494, 496, 569, 607
521–523, 526, 529, 533, 535, 537–538, 540–541, 557, primary 367
559, 561–562, 565–566, 585, 596–597, 599, 617, secondary 367
620–621, 626–627, 629–630, 645, 660, 663–664,
HR42 532
667–668, 682, 685, 688, 705, 714–715, 720–721
Green muscardine fungus 618 HR63 532
Green revolution 10, 494, 498–499, 537, 585, 682, 694, Humidity 375, 377, 402, 409, 414, 417, 425, 428, 434,
696, 701 455–456, 459, 466, 488–491, 559, 645
Green semilooper 446, 496, 506, 624, 634, 700 Hymenoptera (key to species) 182,184–279
Greenhorned caterpillar 632, 641–642 Hyphomycetes 617–636
Guadalcanal 702
Guam 435 IAC 165 738
Guatemala 405 IET 6286 532
Guinea 375 Inaction levels 607
754 SUBJECT INDEX
India 5-6, 215, 221, 364, 366, 368-370, 372, 374-375, International Institute of Tropical Agriculture (IITA)
379-381, 383, 385, 388-394, 400-404, 407-412, 414, 525
416-417, 420-422, 424-425, 427-428, 430, 432-433, International Rice Research Institute (IRRI) 8, 11,
435, 437-444, 446-447, 449-450, 452, 454, 457, 425-426, 454, 494, 499-501, 504, 525-526, 529-533,
459-460, 462, 466, 470, 525, 531-532, 537-538, 540, 537-538, 541, 593, 627, 632, 658, 663, 665, 667, 683,
551, 556, 566, 587-588, 625, 646, 659, 667, 669-670, 695, 701
681-682, 687 Intercropping 571
Indian meal moth 640, 690 Internodal borer 389
Indica 5 Introduction of natural enemies 586-589, 607
Indonesia 5, 10, 221, 366, 369-370, 372, 378-379, 390, Inundative releases 586-588, 607
394, 400, 403-404, 407, 411, 414, 420, 422, 425-427, IPM-field guide 743, 748
430, 432, 437, 440-442, 446, 449-450, 454, 457, 467,
IR8 520, 530, 695
520, 525, 532, 538, 540, 565, 620, 638, 681, 687,
695-697, 702, 706 IR20 519, 531, 698
Insecticides 10, 401, 432, 455, 487, 517, 519-520, 522-523, IR22 523, 529, 664
566, 568, 572, 585, 590, 600, 602, 613, 641, 657-674, IR24 530
682-684, 688-690, 694, 696-697, 706-707, 713-716, IR26 425, 522, 530, 538-539, 673, 682, 696, 699, 702-703
722, 728, 748 IR28 523
advantages of 658 IR36 455, 509, 519-520, 522-523, 530-532, 534, 536-537,
amount used 714, 728 539, 682, 696, 702
application 522, 602, 722 IR38 533
application equipment 518, 658-659 IR40 531, 533, 742
application methods 660, 662-667, 670, 682, 698, IR42 510, 520, 533, 696
704 IR46 7, 673
cost of 657-659, 702, 713, 748 1R48 530
disadvantages of 657-659 IR50 531
effect on grain yields 658 IR52 530
effect on natural enemies 603, 605-407, 613, 657, IR54 531
659-661, 663, 666, 669, 671-672, 674, 682 IR56 529
effective 659-660 IR60 530, 703
factors affecting efficacy of 660-661, 668-670 IR62 530, 606
hazards to humans 672-673, 726 IR66 703
integration of 673-674, 688-690, 699, 706, 729, IR532 E 576 531
739 IR1820-52-2 531-532
mixtures of 659, 663, 668, 697-698 IR1917 606
number of applications 726-728, 742, 748 Iran 394
rates 426, 522, 661, 666-667, 673, 682, 722-723 IRAT 525, 738
residues 659, 671-673, 722, 734 Iridescent viruses (1Vs) 644
resistance to 524, 657-658, 661, 665-669, 674, Irrigated rice 367, 371, 373-374, 376-377, 379, 391,
683, 734 405-406, 423, 425, 437, 440, 450, 456, 492, 494, 558,
selective use of 613-614, 660, 664, 669, 672, 689, 563, 565, 571, 697, 733-734, 738
695, 702, 704-705, 719, 722-723, 729, 739 Irrigation 10, 426, 489, 494-495, 498-499, 501, 504, 507,
subsidy 696 510, 563, 571
timing of application 659, 661-662, 666-671, 682, Isofenphos 659, 670
697, 704, 717, 723 Isoprocarb 659-660, 665, 668, 697-698
tolerance limits 672 Isoptera (key to species) 182, 187, 280
Instituto Colombiano Agropecuario (ICA) 530, 743 Isoxathion 666
Integrated pest management 10, 519, 529, 585-586, 600, Israel 419
602, 606, 608, 674, 681-682, 684-687, 689-690, 694, Italy 366, 389, 394, 438, 452, 463
703-704, 706-707, 713-729, 733-748 Ivoty Coast 375-377, 396, 405, 419, 459
SUBJECT INDEX 755
Japan 5, 8-9, 11, 201, 215, 223, 364, 366-367, 370, 378-379, Leaf roller 63, 642, 736-737, 744-745
386-388, 394, 396, 400-403, 407, 409, 411, 414, 418, Leaf worms 552
420, 422, 425, 427-428, 430, 432-435, 439-442, 446, Leaf yellowing 414
449-450, 452-454, 456, 461, 463-464, 466-467, 490, Legume 447, 686, 690, 741
497, 499, 525, 529-531, 551-552, 555-556, 559, 562,
Lepidoptera (key to species) 57, 59, 61-78
565-566, 569-570, 572, 586-591, 632-633, 641,
657-658, 660, 665-666, 669, 681, 687-688, 696, 701, Lespedeza 562
705, 713-729 Lesser cornstalk borer 389, 527
Japonica 729 Lesser grain borer 527
Java 203, 221, 490-491, 557, 564, 566 Leuang 152 532
Javanica 5, 702 Liberia 375-376, 525
Jaya 682 Light trap 365, 371, 374, 403, 428, 492, 494, 499-500,
502, 505, 550, 552, 554-556, 571, 661, 670, 705
Jute 449
Lindane 672, 698
Juvenile hormone 659
Locust 554, 570
Long-duration rice varieties 563
K-type species 735-736
Longhorned grasshopper 602
Kampuchea 369-370, 372, 392, 394, 400, 407, 420, 422,
427, 430, 432, 437, 439-442, 449-450, 457, 525 Louisiana 5, 388, 391
Kanto district 716, 721-722 Lowland rice 376, 495
Karnataka 532 Lowland irrigated rice 492
Kencana 673
Kenya 391, 419-420, 459, 493 Macropterous 410, 421, 423-424, 427-429
Kerala 437, 659 Madagascar 381, 405, 411, 413, 415, 419-420,432, 437,
450, 459, 494, 499
Kerosene 551, 554-555, 571, 657
Madhya Pradesh 443
Key pests 683, 700-701
Maharashtra 461
Kinki district 721
Maize 6, 391, 408, 421, 427, 444, 463, 493-494, 562.
Kochi prefecture 396, 403, 722-723
739-740
Korea 5, 8, 378, 394, 407, 411, 420-422, 425, 427, 430,
Maize borer 389
432, 434-435, 439, 441-442, 446, 449-450, 452, 461,
Maize weevil 527
463, 525, 530, 626-628, 633, 666, 701, 705
Malagasy 3, 459, 525
Kyushu district 401, 418, 718, 721
Malathion 658, 665, 669, 698, 702
Malawi 391, 419, 437
Lady beetles 592, 594
Malaysia 9, 215, 221, 366, 369-370, 379, 390, 392, 394,
Land preparation 549, 568, 740
400-403, 407, 409, 411, 414, 418-420, 422, 427, 430,
Law 372, 392, 394, 400, 403, 407, 420, 422, 427, 430, 432,
432, 435, 437, 439, 441-442, 446, 449-450, 452, 457,
437, 439, 441-442, 449-450, 454, 457, 525 465, 467, 470, 494, 502, 525, 537, 586-589, 600-603,
Large rice grasshoppers 408 632, 658-659, 661, 681, 687, 702
Latin America 3, 11-12, 90, 397, 488, 492, 504, 733-748 Male-sterile 531, 541-542
Leaf beetles 461, 552, 555, 558, 560, 566, 568, 714, 716, Mali 375-377, 396, 459
720-721 Manchuria 439
Leaffolder 63, 184, 209, 408, 432-435, 505, 527, 539, Mangrove swamp 493
541-542, 553, 555, 557-558, 560-564, 568-571, 586,
Marcescent process 621-622, 625, 636
590, 593-595, 608, 613, 634, 636-641, 644, 669-670,
700-701, 717 Masagana 99 Program 696-698, 702
Mauritius 405, 437
Leafhoppers 83, 85, 87, 280, 412, 416-417, 420, 524, 526,
532, 534, 541, 552, 555-558, 560, 562-564, 566, 568, Mauritius pink borer 389
570, 590, 595, 608, 613, 616, 618, 621, 625-628, 632, Mealybug 364, 368-369, 443, 553-554, 568
636-637, 642, 658, 671, 688, 697, 706, 722, 724, 729, Mechanical control methods 10, 549-556, 560, 570-571
735-736, 743-745 Medium maturing cultivars 563, 734
Leaf miner 452-453, 564, 568, 738 MEP 723
756 SUBJECT INDEX
Mercaptophos 698 Nansei Islands 718

Metalkamate 661–662, 669 Natural control 700
Methyl parathion 660–661, 667–668, 671, 688 Natural enemies 12, 424–425, 487, 489, 498, 503, 508,
Mexico 389, 391, 405, 428, 447 510, 520–5219 523, 551, 558, 560, 563–564, 566,
Microbial insecticides 614 571–572, 585–608, 613–646, 661–662, 687, 696,
Microencapsulation 663 700–701, 703–706, 718–719, 729, 738–739, 743, 748
Midges 559 Natural hybrids 524
Migration 365, 367, 385, 425, 428, 441, 444, 499, 519, Neem cake 660
701, 705, 717–718, 729, 737 Nematodes 524, 633, 645–646
Millets 408, 421, 427, 444, 449, 463, 466, 470 Neotropical corn borer 389
Minnesota 469 Nepal 368–369, 390–391, 394, 400, 407–408, 414, 422,
Minor genes 536 427, 430, 432, 437, 439, 442, 449–450, 457, 488,
524–525
MIPC (see. Isoprocarab)
Netting 570–571
Mind bug 521, 594, 601, 603, 671, 684–685, 688
Neuroptera (key to species) 135
Mites 566, 722
Mitsubishi 697 New Guinea 366, 372
Mixed cropping 550, 556 New York 469
New Zealand 442, 463
MLO disease 534
Moderately resistant cultivars 522, 531, 533–534, 539, Nicaragua 405
662, 673, 682–684, 687, 741–742 Niger 375, 396
Modem varieties 9–10, 499, 504–505, 538, 542, 558, 571, Nigeria 375–377, 396, 400, 410, 419–420, 437, 462
585, 593, 687, 695, 697, 734, 738 Niigata prefecture 723–725
Molasses 630 Nitrogen 369, 375, 403, 414, 426, 428, 433, 435, 443, 459,
Mole cricket 364–365, 553–555, 560, 562, 567, 570, 744, 560–561, 565–566, 687, 740–741
748 Noctuid borers 388
Molecular biology techniques 541, 543 Non-preference (see Antixenosis)
Molting inhibitor 706 North Africa 452, 463
Mongolia 427 North America 8, 236, 463, 718
Monocrotophos 662, 665–666, 671, 697–698, 702 Nuclear polyhedrosis viruses (NPVs) 620, 642–643
Monoculture 493, 700 Nursery tray treatment 716–718, 722–723, 726–727
Monogenic resistant 683 Nyasaland (see also Malawi) 381
Monsoon rains 374–375, 384, 390, 402, 428, 430, 451,
466, 552, 563, 565, 567 Oats 421, 445, 453, 463
Moonlight 403, 555 Ob 677 533
Morocco 366, 419 Oceania 8
Mosquito 374, 645 Odonata (key to species) 130–135
Mozambique 437 Oil palm 586
MTMC 663, 671 Oils 536, 555
Muda scheme 600 Onion leaf 373, 375
Mudgo 535, 696 Orange-leaf 412, 414, 420
Mulberry trees 641 Organochlorines 672, 697, 702, 704, 715
Multigenic resistance 689 Organophosphates 401, 668, 696–697, 704–705, 716, 723,
Multilines 540, 704 726, 734
Multiple resistance 532–533, 658, 668, 682 Oriental locusts 495
Murunaga 139 531 Orissa 532, 538
Mycological terms used 615 Orthoptera (key to species) 38–54
Oryzanone 536, 561
N 22 521 Oryzica 1 748
Naled 666 Ovicidal action 659, 668
SUBJECT INDEX 757
Oxalic acid 535 Physical control methods 10, 549-556, 570-571

Phytoalexin 533
Paddy root weevil 385 Phytotoxicity 571, 659
Pakistan 369-370, 372, 379, 391, 394, 400-401, 403, Pigeonpea 557
407-409, 414, 420, 422, 427, 432, 439-442, 449-450, Pink borer 389, 407-408
457, 470, 525, 687 Pink stem borer 5, 389, 407, 640-643
Pale-headed striped borer 389 Plant
Panama 405 age 432, 534-535, 540, 549-550, 552, 558, 563,
Panicle thrips 470 571-572, 590, 592-593, 604, 662, 744-745
Pankhari 205 537 anatomy 536
Pannetti 531 chemistry 536, 607
Papua New Guinea 368, 370, 379, 394, 407, 414, 422, damage 367, 369, 374, 412-413, 421-423, 425,
427, 430, 432, 437, 442, 449-450, 457, 466 427, 429-430, 433, 437-438, 440, 442, 453,
Parasites 402, 508, 517, 520, 551, 559, 571-572, 613-614, 455, 457, 459-462, 465, 470, 550, 558-559,
673, 684, 690, 700-701 571, 673, 693, 719, 725, 735-736, 743, 745
Parasitization 373, 402, 645, 661, 669-670 density 375, 736, 740
Parasitoids 585-597, 599-600, 602-603, 606-608 growth stages 6, 364, 395, 403, 734, 744-748
Parathion 658, 667-669, 698, 702, 715 maturity 520, 550, 557, 572
Parthenogenetic 367, 369, 386 morphology 536, 607
Pathogen physiology 536
insect 508, 517, 559, 568, 585-586, 613-646, 684, senescence 364
700, 706 Planthoppers 83, 90, 280, 412, 428, 524, 526, 539, 541,
plant 517, 571, 735 552, 555, 558, 560-564, 566, 568-570, 590, 595-596,
Pecky rice 463-464, 467, 469 608, 613, 616, 618, 621, 625-628, 632, 636-637, 642,
Pentadecanal 536 658-659, 671, 673, 697, 706, 720, 726, 729, 735-736,
Permethrin 665-666, 668 743-745
Peru 405, 586 Planting
Pesticides 9, 561, 728-729, 743 dales 491-492, 504-505, 507
Phagostimulants 534 density 57
Phenotypes 491 method 456-457, 459, 550, 557, 716
Phenthoate 663 time 507, 550, 557, 563-565, 607, 684, 686-688,
Pheromone 433 701, 715-716, 718-719
Plowing 496, 549, 553-554, 567, 572, 717
Pheromone traps 556
Philippine Seed Board 699 Pollinators 671
Polyculture 493
Philippines 8, 221, 366, 369-371, 379, 381, 390, 392, 394,
400-401, 403, 407, 412, 414, 420-422, 424-427, 430, Polygenic resistance 531, 536, 539
432, 436-442, 446-447, 449-451, 454-455, 463-464, Polymorphism 441
466, 491-492, 494, 499-500, 503-505, 508, 525, 532, Potash 403
537-538, 562, 587-590, 593, 596-597, 618, 620-621, Potassium 435, 561
625-626, 628, 632-635, 638 643-645, 658, 665, 668, Potato 690
673, 681-682, 695-696, 699, 701-703, 727 Potato ladybug 625
Phloem feeding 364, 425, 533-534, 538 Predators 378, 508, 517, 520, 567, 570, 572, 585-586,
Phorate 659, 681 589-608, 613-614, 629, 669, 684, 690, 700-701
Phosalone 671 Prey : predator ratio 597-598, 600-601, 603-604
Phosphamidon 659, 672, 698 Primitive cultivars 524
Phosphate 375, 667 Probing 536
Phosphorous 435, 561 Propaphos 666, 668
Photoperiod 441 Propoxur 665, 671
Photoperiod insensitive cultivars 563 Protoplast techniques 542
Photoperiod sensitive cultivars 563-565 Pruning 550, 558, 571
758 SUBJECT INDEX
Ptb 10 532 Rice dwarf disease 412, 565, 715, 717-718, 720, 723,
Ptb 18 532-533 Rice earcutting caterpillar 442
Ptb 21 532-533 Rice garden 507, 562
Puerto Rico 368, 468 Rice grasshopper 410
Purple stem borer 389 Rice leafminer 537
Pyralid borers 388 Rice moth 643
Pyrethroid insecticides 666-668, 734 Rice plant weevil 384, 542, 716-717
Pyrethrum 555 Rice skipper 28, 440-441, 635-636, 641, 643, 716
Pyridafenthion 665-666, 669 Rice stalk borer 389
Pyrimioxythion 671 Rice-stripe virus 421, 530, 715-716, 720, 722
Rice striped horer 389, 394-395, 526, 531
Quinalphos 659, 671 Rice weevil 527
Rice yellow dwarf disease 715, 717
Ragged stunt 423, 542, 688, 702 Rice-free peritd 563
Ragi stem borer 389 Risk threshold 743-745
Rainfed lowland rice 371, 376, 406, 412, 414, 437, 443, R-K continuum 735-736
464, 492-493, 504 Rockefeller Ibundation 541-543
Rainfed rice 373, 377, 504, 508, 558, 571 Rodents 524, 554, 571
Raipur 538 Roguing 550, 553-554, 571
Ratna 531 Rollers, for insect control 555, 571
Rattoon 416, 461, 550, 570, 700, 717 Root aphids 80, 364-365, 367, 553, 558, 560, 564, 568
RD4 532 Root bugs 567-568
RD9 532 Root dipping 670
Reoviruses 642 Root feeders 164, 559
Repellent, insect 557 Root soaking 667
Reproductive stimulation 426 Root weevils 383, 553-555, 558, 560-561, 564, 566, 568
Resistance to insecticides 658 Root-zone application 664-665, 667, 671, 688
Resistant varieties 10, 425, 428, 435, 518-519, 526-527, R-type species 735-736
572, 585, 606-607, 662, 666, 673, 681-690, 696-497, Rusty plum aphid 462
699, 702, 704, 719, 722, 737-738, 742, 748 Rwanda 405, 459, 498-499,
Resurgence, insecticide induced 10, 522, 555, 585, 606, Ryuku Islands 400, 407, 461
658, 661-662, 666-667, 671, 674, 682-683, 694-695,
700-702, 722, 734, 739, 741-742 Sabah 215, 586
Rhodesia 420 Saline soils 372, 542
Rice Sapan Kwai 537
area 5-6, 713 Sarawak 404, 450-451
classification 5 Scarab beet1t-s 380, 552
consumption of 3, 733 Screening mcthods 526
cost of prcduction 733, 747 Seed bugs 555-556, 561, 564, 566, 568, 570
gmwth stages 6, 734-735 Seal midgcs 364, 371-372
importance of 3, 695, 733 Sad treatnwnt 698
origin 3 Seedling flies 364, 527
production 696, 713, 733 Seedling maggot 26, 380, 560-561, 564, 568
species 4 Seedling root coating 667
yields 696, 714, 733 Semiaquatic insects 560
Rice blast 399 Semidwarf varieties 734
Rice breeding 539 Senegal 375-377, 396, 405, 419, 459
Rice bug 245, 527, 551-555, 619, 671-672, 689, 697 Sequential release of varieties 539
Rice delphacid 428-429, 526, 529-530 Sequential sampling 602
SUBJECT INDEX 759
Sequential threshold 743, 745 Spiders 12, 14-15, 520-521, 592, 600-603, 616-617, 620,
Sexual dimorphism 401 626, 629, 638, 661, 665, 669, 671, 684-685, 694, 701,
Shakti 532, 682 706, 719, 722, 743
Sheath blight 542 Spiders (key to species) 283, 285, 287-340
Shikoku district 718, 721 Spittlebugs 618, 621
Siam 29 532 Spotted stem borer 390
Siberia 421, 461 Spray volumes 662
Sierra Leone 366-367, 375-377 Sri Lanka 369-370, 372, 374, 379, 390-391, 394, 400-402,
407-408, 412, 414, 420, 422, 427, 430, 432, 436-437,
Sight counting board 726-727
442, 449-450, 457, 470, 525, 532, 538, 658, 666, 670
Sikkim 391, 400
Stable resistance 703
Silica 435, 459, 537, 552, 561
Staggered planting 507, 520, 563, 565
Silicic acid 535
Stalk-eyed fly 376-377, 496, 527, 529
Silkworm 633, 641, 643
Stem borers 29, 63, 370, 380, 388-390, 398-401, 403, 407,
Silver shoot 373, 532, 554
459, 490, 493, 495496, 498-499, 510, 518, 532,
Singapore 379, 407 536-537, 541-542, 549-553, 555-556, 558-572,
Single cropping 498, 503-504, 510, 568 585-588, 590, 592-595, 600, 603, 607-608, 613-614,
Sinica 5 632, 635-636, 640-642, 645-646, 658, 668-669, 673,
Sinna Sivappu 684 697-698, 700, 738
ß sitosterol 535 Stem maggot 364, 378-379, 527, 537, 555, 722
Small moth borer 389 Sterols 535
Small purple-lined borer 389 Sticky board 601
Small rice grasshopper 411 Stink bugs 463-464, 552, 613, 616, 618, 632, 715, 718,
Small rice miner 716 721, 735-736, 743, 745-748
Smaller brown planthopper 421-422, 526, 530, 562, 566, Storage insects 529, 541-543, 557
659-660, 665-666, 715, 720-721 Straw destruction 550, 569-570, 572
Soil fertility 369 Strepsiptera (key to species) 14, 16-18, 280
Soil injection 671 Stresses, plant 372, 375, 403, 414, 426, 428, 433, 435, 444,
Soil pests 567, 735, 737, 744 455, 459, 558, 572
Soil types 384, 524 Stripe disease 422, 660
Solomon Islands 407, 422, 424, 435, 618, 630-631, 702 Striped stem borer 389, 490, 536, 541-542, 552, 559, 567,
Sooty mold 423, 429 569-570, 572, 591, 635, 640-641, 668-669, 687-688,
Sorghum 391, 463, 470, 493 714-715, 720-721
Sorghum stem borer 389 Stubble burning 550, 569, 684, 687
Sorjan system 557 Stubble destruction 568-570, 572, 687
South Africa 420, 459 Sucking inhibitors 535
South America 5, 8, 12, 240, 389, 397, 430, 443, 447, 463, Sucking stimulants 535
468, 529-530 Sudan 375, 391, 397, 405, 410, 419-420, 459, 695
South American white borer 389, 405, 527 Sugar 534-535, 553, 561
South Asia 10, 221, 240, 243, 248, 280, 307, 374, 403, 409, Sugarcane 381, 390, 407-408, 421, 461, 463, 553
412, 422, 493, 529, 531, 538, 586, 588-589, 606-607, Sugarcane borer 389, 527, 587, 736
608 Sugarcane internodal borer 389
South Carolina 5 Sugarcane leathoppers 621
Southeast Asia 3-4, 10, 205, 221, 243, 248, 299, 307, 374, Sumatra 702
389, 403, 409, 412, 422, 436, 488, 493, 529, 531, 538, Sunlight 550, 641, 643
586, 588-589, 606-607, 668, 695, 701, 703 Superstitious insect control practices 550
Southern green stink bugs 467-468, 718 Surekha 532
Soybeans 6, 562, 634, 641-642, 739-740 Surinam 366, 405, 428
Spain 389, 394, 463 Surveillance system 601-602, 681-682, 706, 718, 723-727,
Spider mites 735 743
760 SUBJECT INDEX
Survey, of natural enemies 599-601, 608 Transitory yellowing 413-414

Swaziland 376-377, 459 Trap crop 549-550, 566, 569, 572, 674, 687-688
Sweet flag 557 Triazophos 660
Sweet potato 449, 553 Trinidad 389, 405, 587
Symbiotes 422 Triple cropping 503, 563, 701
Synchronous planting 487, 509, 520, 550, 561, 565-568, Triveni 520, 540, 662, 673
572, 607, 686-687, 690, 697, 701 Tropical Development Research Institute (TDRI) 536
Systemic insecticides 668, 670, 704 Tungro virus 412-414, 419-420, 522-523, 529, 534-535,
540, 542, 559, 565, 570-571, 667
Taiwan 215, 221, 223, 366-367, 390, 394, 400-403, 407, Tunisia 419
409, 411-412, 417, 420-422, 426-427, 430, 432, 437, Typhoons 565
440-442, 446, 449-450, 452, 454, 457, 461, 463, 529,
563, 569, 657-658, 661, 665-666, 668 U.S.A. 3, 5, 366, 368, 371-372, 382, 386, 388-389, 391,
Tamil Nadu 667, 669-670 397, 428, 444, 447-449, 452, 463, 468-469, 525, 529,
Tanjung Karang irrigation scheme 600 570, 589, 728
Tanzania 391, 405, 419-420, 450, 459, 499 U.S.S.R. 366, 439, 525, 625
Temperature 9, 375, 382, 387-388, 395-396, 402, 405, Uganda 391, 419-420, 450, 459, 499
407, 410, 414, 416-417, 419, 422-425, 428, 431-432, ULV sprayer 663
434, 441, 453, 456-457, 462, 488-489, 491, 494, 553, Upland rice 364, 367, 371, 373, 379-381, 391, 400,
560-561, 564-565, 567, 621, 637, 643 404-406, 423, 427, 430, 441, 443, 450, 458, 461, 464,
Termites 567 492-493, 503-504, 625, 738
Terpenoids 536 Uruguay 437
Tetrachlorvinphos 666 Utri Rajapan 540, 673
Texas 391, 396 UV light 641
Thailand 203, 215, 366, 369-370, 372-375, 379, 390-392,
394, 400-401, 403, 407-408, 414, 420, 422, 427, 430, Vegetable crop 690
432, 439-442, 449-450, 454, 457, 463, 465, 489, 525, Velvetbeans 634
532, 537-538, 589, 638, 670, 695 Venezuela 389, 405, 437
Thrips 55, 430-432, 470, 527, 532, 554, 558, 560-562, 564,
Vertical resistance 703
571, 673, 717
Vietnam 369-370, 372, 378-379, 390, 392, 394, 400,
Tiger beetle 141 407-408, 417, 420, 422, 427, 430, 432-433, 437,
Tillage 383, 550, 567-568 439-442, 446, 449-450, 457, 525, 532, 537, 540, 638
Timothy 453 Violet stem borer 389
TKM 6 531-532, 536 Viruses 413-414, 419, 421-423, 427, 463, 508, 522, 524,
TN 1 520-522, 662, 684 530, 534, 555, 563, 566, 568, 570, 572, 614, 667, 688,
Tobacco 447, 449, 607 702, 716, 719, 729, 736, 748
Togo 375, 459 Volatile chemicals 535, 561
Tohoku district 721
Tokai district 721 Waika virus 414, 419, 722
Tolerance, plant 533-534, 533-540, 558, 565, 673, 690, Water bugs 592, 599
735, 738, 741-742, 748 Water management 550, 559, 741
Tomato 690 Water stress 369, 371, 571
TOX 1011-4-1 738 Water weevils 386, 499, 527, 529, 556, 559-560, 568, 633,
Toxaphene 698, 702 646, 688, 714, 718, 720-721, 735-736, 739
Traditional controls 569 Water-foxtail 418
Traditional cropping systems 657, 743 waxes 368-369, 536
Traditional cultivars 526, 538, 558, 563, 571, 593, 734, WC1252 532
742 WC1253 532
Transgenic plants 542 WC1257 532
Transitory rice virus 414 WC1263 531-532
SUBJECT INDEX 761
Weeding 549-550, 568-569, 747 Wild rice 3-5, 375, 394, 400, 415-416, 489, 506, 524, 531,
Weeds 8, 366, 371, 375, 401, 416, 418, 422-423, 428-429, 538, 540-542, 569
447, 449, 453, 457-458, 463-465, 496, 524, 554, Wildlife 377
566-571, 603, 607, 687, 735, 740, 748-747 Wilted stunt virus 423
West Africa 396, 439, 444, 463 Wolf spider 591, 598
West Bengal 403
West Indian Islands 397, 447 Xylem feeding 533, 535, 538
Wetland rice 557, 560, 568
Wet season 374, 503
Yeasts 630-631
Whale oil 555, 657
Yellow disease 463
Wheat 6, 366, 421-422, 445, 453, 463, 470, 543, 562, 625,
641, 716-717 Yellow dwarf 413-414, 418, 562, 570
White borer 389, 398 Ycllow mottlevirus 460
White grubs 364, 380-382, 493, 552-553, 560-561, Yellow rice borer 389, 569
563-564, 567, 570, 625, 736, 744, 748 Yellow stern borer 9-10, 389, 392-393, 401-404, 492,
White leafhopper 412, 526, 616 495-496, 500, 502-504, 506, 508, 520, 526, 531, 536,
White leaf disease 429 541-542, 552, 557, 559, 561, 563, 567, 569, 590-591,
594, 602, 668, 682, 715
White muscardine fungus 625-626
White rice borer 389 Yellow-orange leafvirus 413
White stem borer 9, 389, 403-404, 489, 491, 496, 687 Yield losses 7-10, 369-371, 373-374, 377, 379, 383, 385,
387-388, 390, 394, 401, 404, 406, 411-412, 414, 418,
White-tip nematode 717
421, 427, 433, 443, 445, 450, 454-455, 457, 461, 463,
Whitebacked planthopper 426-428, 521-522, 526, 530,
465, 467, 469, 503, 517, 552, 593
541-542, 557, 596-597, 599, 626, 645, 659-660,
665-666, 682, 714, 717, 720-721
Whiteheads, stem borer 390, 393, 397, 403-404, 406, 408, Zaire 437, 459
553-554, 556, 646 Zambia 366, 419-420
Whorl maggot 9, 454-456, 527, 541, 549, 557-559, Zigzag leafhopper 420-421, 526, 529
561-562, 564, 570, 667, 682, 736 Zimbabwe (see Rhodesia)
Wide hybridization 540-541 Zinc 561
Species Index
Achaea janata 72, 74, 227 Alopecurus aequalis 417-418, 430

Achromobacter nemotophilus 646 Alopecurus fulvus 378
Aciagrion occidentale 133, 135, 137 Amauromorpha accepta metathoracica 215-216
Acigona chrysographelia 68-69 Amauromorpha accepta schoenobii 215-216
Acrida exaltata 409 Amphiareus 111-112
Acrida turicata 409 Ampittia maro 59, 62
Acrida willemsii 43, 47 Ampycoides pallidicosta 398
Acrostemum 740 Anagasta (Ephestia) kuhniella 640
Acrotylus humbertiana 409 Anagrus 257
Adoretus 26 Anagrus armatus 246, 269
Adoretus luridus 166-167, 169 Anagrus flaveolus 263, 267
Adoretus ranunculus 165-167 Anagrus frequens 266, 269
Aelopus affinis 409 Anagrus incamatus 263, 267
Aeneolamia 740 Anagrus optabilis 263, 265
Aeolodenna brachmana 176-177 Anagrus panicicolae 263, 261
Aeolopus tamulus 409 Anagrus perforator 263, 267
Aeolothtrips fasciatus 55, 58 Anagrus sp. A 263, 266, 269
Aeschynteles maculatus 464 Anagrus sp. B 266, 269
Aethus indicus 99-100 Anagrus spp. 596
Agamennis 645 Anaphothrips obscurus 57, 60
Agamennis unka 645 Anaxipha longipennis 39, 42, 599
Agelena labyrinthica 305, 308 Ancylolomia chrysographella 389
Ageratum conyzoides 470 Andrallus spinidens 106, 109
Agriocnemis femina femina 133-134 Andropogon 740
Agriocnemis pigmaea 133-134 Andropogon gayanus 410
Agromyza oryzae 34-35, 279, 537 Andropogon tectorum 398
Agropyron sp 450, 452 Anerastia pallidicosta 398
Agropyron kamojo 430 Anisops kuroiwae 94, 96
Agrostis tenuis 452 Anisops sp. 94, 96
Agrothereutes lanceolatus 220, 222 Anoecia comi 366
Agrotis ipsilon 72, 74 Anoecia fulviabdominalis 366-367
Ailopus thalassinus tamulus 45, 47 Anoecia sp. 366
Alisma plantago-aquatica 384, 452 Anomala 26
Allium cepa 452 Anomala dimidiata 381
764 SPECIES INDEX
Anomala humeralis 166, 169 Bacillus thuringiensis 542, 614, 638, 640-642, 673, 690,
Anomala sulcatula 166, 169 706
Anoplogenius microgonus 156-157 Bactra venosata 220-221
Antestia anchora 106, 109 Bagnallia biformis 430
Anthicus sp. 638 Baliothrips biformis 430, 717
Apalochrus rufofasciatus 181, 185 Beauveria 618, 632
Aphanogmus fijiensis 233-234 Beauveria bassiana 618-619, 622-628, 632-633, 635-636,
Aphanogmus sp. 233-234 672
Aphelenchoides besseyi 717 Beauma brongniarti 625, 636
Aphis gossypii 463 Beauveria densa 625
Aphis craccivora 463 Beauveria spp. 625
Apluda mutica 368 Beauveria tenella 625
Arachis hypogaea 410 Beckmania crucaeformis 407
Araneus 339-340 Bellis perennis 452
Araneus diadematus 339-340 Berosus sp. 168, 171
Araneus inustus 339-340 Bessa remota 588
Arbela nitidula 120-121 Bianor hotingchiehi 285, 287, 289
Archicolliuris bimaculata 148, 150-151 Blissus 739-740
Arctosa janetscheki 313, 316 Blumia wightiana 470
Argiope aemula 331, 334 Bothriochloa pertusa 470
Argiope bruennichii 331, 334 Brachiaria 740
Argiope catenulata 331, 334 Brachiaria mutica 397, 432, 437, 442, 450
Argyria sticticraspis 390 Brachiaria ramosa 431
Argyrodes bonadea 304, 306 Brachiaria reptans 368
Argyrodes fissifrons 304, 306 Brachiaria setigera 442
Argyrodes miniaceus 301, 303-304 Brachinus sp. 145, 149
Argyrodes saganus 301, 303 Brachymeria albotibialis 275, 277-278
Argyrodes sp. 304, 306 Brachymeria excarinata 275, 278
Argyrophylax 29 Brachymeria jayaraji 275, 278
Argyrophylax fransseni 29-30 Brachymeria lasus 275, 278
Argyrophylax nigrotibialis 29-30 Brachymeria megaspila 275, 278
Argyrophylax phoeda 29-30 Brachysiphoniella montana 463
Arundinella hirta 430 Bracon chinensis 201, 204, 586-581, 591
Arundinella metzii 460 Bracon onukii 203-204
Aspergillus 618 Bradina admixialis 225
Asytasia gangetica 470 Brevennia rehi 80, 82, 368
Atherigona exigua 379, 527 Bromus inermis 452
Atherigona oryzae 26-27, 379-380, 527 Brumoides lineatus 172, 174
Atractomorpha crenulata 434, 409 Brumoides suturalis 172, 174
Atractomorpha psittacina 43-44 Busseola fusca 211
Atypena (=Callitrichia) formosana 311, 319, 321
Aularches miliaris 41-42 Calcarisporium ovalisporun 630-632
Avena fatua 417 Callitriche verna 384
Avena sativa 442, 452, 468 Callitrichia spp. 638
Axonopus compressus 450 Camaricus formosus 291, 294
Azadirachta indica 660 Camponotus sp. 283, 286
Camponotus spp. 191, 194
Bacillus cereus 641 Campsicnemus sp. 24-25
Bacillus subtilis 641 Campsomeris annulata 233-234
SPECIES INDEX 765
Campsomeris collaris quadrifasciata 229, 231-232 Chilo plejadellus 68-69, 71, 389, 391
Campsomeris nr. aurulenta 231, 233-234 Chilo polychrysus 9-10, 70-71, 207, 238, 389-390, 392,
Campsomeris sp. A 229, 232 587-588, 591
Campsomeris sp. B 231-232 Chilo sacchariphagus 217, 220, 238, 240
Campsomeris sp. C 231-232 Chilo simplex 394, 586, 588
Campsomeris sp. D 231, 234 Chilo spp. 29, 248
Carcelia exisa 28, 30 Chilo suppressalis 70-71, 201, 203, 205, 211, 215,
Cardochiles philippinensis 213, 216, 594 217-218, 220-221, 223, 225, 227, 238, 388-390, 394,
Cardiochiles sp. 213, 216 396, 494, 526, 531, 565, 586-588, 591, 635-636,
Carex olivacea 440 640-641, 645-646, 668-669, 687-689, 715-717, 719,
722-726, 728
Caryanda diminuta 46, 49
Chilo zacconius 70-71, 389, 396-397, 527
Casinaria ajanta 223-224
Chilo zonellus 211, 390
Casinaria pedunculata pedunculata 223, 226
Chilotraea auricilia 390
Casnoides interstitialis 602
Catantops pinquis 50, 52 Chilotraea polychrysa 392, 494
Celosia argentia 470 Chiracanthium japonicum 295, 298
Centeterus alternecoloratus 218-219 Chiracanthium lascivum 297, 300
Centistes medythiae 205, 208 Chiracanthium nr. unicum 297-298, 300
Ceracia spp. 31-32 Chiracanthium sp. A 295, 298
Ceraphron sp. A 233-234 Chiracanthium sp. B 295, 297-298
Ceraphron sp. B 233-234 Chiracanthium sp. C 297, 300
Ceromyia silacea 31-32 Chironomus crassiforceps 20-21
Chaetocnema 119, 137, 740 Chironomus dorsalis 20-21
Chaetocnema abyssinica 181, 183 Chironomus javanus 21-22
Chaetocnema basalis 181, 183 Chironomus kiiensis 20-21, 372, 507
Chaetocnema concinnipennis 181, 183 Chironomus sp. 372
Chaetocnema kenyensis 179, 181, 183 Chironomus tepperi 20-21
Chaetocnema pallidipes 181, 183 Chirothrips manicatus 55, 58
Chaetocnema pulla 181, 183 Chlaenius bioculatus 158, 161
Chaetocnema sp. 737 Chlaenius circumdatus 156, 159
Chaetogeoica polychaeta 366 Chlaenius inops 156, 159
Charops bicolor 227-228, 230 Chlaenius quadricolor 160, 163
Charops brachypterum 227-228 Chlaenius sp. A 158, 161
Charops ganges 227-228 Chlaenius sp. B 158, 161
Chelisoches morio 130, 132 Chlaenius sp. C 158, 161
Chelonus munakatae 205-206 Chlaenius sp. D 160, 163
Chelonus sp. 205-206, 640 Chlaenius virgulifer 156, 158-159
Chenopodium album 382 Chlaenius xanthospilus 156, 159
Chilo 390, 494, 551, 562, 642, 645 Chloethrips oryzae 430
Chilo agamemnon 68-69, 389 Chloriona furcifera 426
Chilo auricilius 70-71, 217, 389, 390 Chloris barbato 368, 470
Chilo diffusilineus 70-71, 250, 397 Chloris virgata 459
Chilo hyrax 70-71 Chlorops oryzae 33-34, 201, 378, 456, 537, 722
Chilo ignefuscalis 211 Chondracris rosea 48, 52
Chilo infuscatellus 217, 221 Chrotogonus sp. 409
Chilo loftini 389 Chrysodeixis chalcites 643
Chilo oryzae 394 Chrysopa basalis 135, 137
Chilo partellus 70-71, 389-390, 493, 632 Chrysopogon aciculatus 406, 410
Chilo phaeosoma 389 Cicadella spectra 412
766 SPECIES INDEX
Cicadulina bipunctata 87-89 Conocephalus longipennis 41-42, 44, 248, 591, 602
Cicindela chinensis 145, 147 Conocephalus maculatus 41, 44, 669
Cicindela elisae 141, 144 Copidosomopsis nacoleiae 266, 269, 593-594
Cicindela ferriei 141, 143, 147 Coptotermes formosanus 182, 187, 280, 284
Cicindela japana 143, 147 Cordius sp. 645
Cicindela japonica 143, 147 Cornus spp. 367
Cicindela kaleea 138, 142 Cotesia (= Apanteles) angustibasis 207, 210, 594
Cicindela lacrymosa 141, 143, 146 Cotesia (= Apanteles) opacus 207, 210
Cicindela nr. ferriei 141, 144 Cotesia (= Apanteles) prodeniae 207, 210
Cicindela nr. kaleea 138, 142 Cotesia agilis 209, 212
Cicindela nr. lacrymosa 143, 146 Cotesia baoris 209-210
Cicindela nr. sexpunctata 136, 138, 140 Cotesia cypris 209-210
Cicindela sexpunctata 136, 139 Cotesia flavipes 209, 212, 591
Cicindela sp. A 136, 139 Cotesia kariyai 211-212
Cicindela sp. B 136, 140 Cotesia plutellae 209, 212
Cicindela sp. C 138, 140 Cotesia ruficrus 211-212
Cicindela sp. D 138, 142 Cotesia sesamiae 211, 214
Cicindela specularis 143, 146 Crambus suppressalis 394
Cicindela striolata 141-142, 144 Creatonotus gangis 70, 74
Cicindela sumatrensis 141, 144 Cricotopus sylvestris 371
Cirphis unipuncta 444 Crocothemis servillia 131, 134
Cletur punctiger 126-127, 464 Cuphocera varia 26-27
Cletus trigonus 126-127 Cybister japonicus 160, 163
Clivina sp. 152-153 Cybister tripunctatus orientalis 162-163
Clubiona japonicola 297, 302 Cybister tripunctatus subsp. A 162-163
Clubiona kurilensis 299, 302 Cyclosa insulana 333-334
Clubiona lena 299, 302 Cylindromyia spp. 26-27
Clubiona sp. A 297, 300 Cylindrorthorax melanocephala 170-171
Clubiona sp. B 299, 302 Cylindrothorax sp. 170-171
Clubiona sp. C 299, 302 Cymbopogon caesius 368
Clubiona vigil 297, 300 Cymbopogon nardus 407
Cnaphalocrocis 184 Cymodema basicornis 125, 127
Cnaphalocrocis jolinalis 432 Cymoninus turaensis 125, 127
Cnaphalocrocis medinalis 63, 65, 205, 207, 209, 213, 215, Cynodon dactylon 368, 373, 379, 416, 420, 437, 441-442,
225, 227, 266, 432, 434, 436, 505, 527, 541, 634-635, 452, 454, 457
637, 639-641, 643-644, 669, 700, 717, 727 Cyperus difformis 377
Coccinella repanda 175, 177 Cyperus diffusa 446
Coccinella septempunctata 175, 177 Cyperus iria 430, 437, 468
Coelinidea oryzicola 201-202 Cyperus japonicus 407
Cofana 280 Cyperus rotundus 368, 384, 411-412, 420, 430, 437, 442,
Cofana spectra 17, 85, 88, 412, 526, 616, 621 452, 470
Coix lachryma-jobi 406 Cyrtacanthacris tatarica 48, 52
Colaspis flavida 382 Cyrtococcum patens 437
Colaspis occidentalis 383 Cyrtophora sp. 333-334
Colaspis prasina 383 Cyrtorhinus 638
Colemania sphenarioides 409 Cyrtorhinus fulvus 108-109
Coleosoma blandum 305, 308 Cyrtorhinus lividipennis 108-109, 521, 591, 594, 596-598,
Colocasia esculenta 370 600-604, 638, 660, 669, 671, 684-685, 688-689, 701
Conidiobolus coronatus 617, 637 Cyrtorhinus mundulus 108-109
SPECIES INDEX 767
Drapettis (Elaphropeza) sp. B22-25

Dactylis glomerata 452, 461 Drassodes sp. 295-296
Dactylispa 176 Drypta geniculata 152, 155
Doctylispa bayoni 178, 180 Drypta japonica 152, 154-155
Dactyloctenium aegyptium 368 Dyschiriognatha quadrimaculata 322, 325
Delphacodes furcifera 426 Dyschiriognatha tenera 322, 325
Delphacodes striatellus 421 Dysdercus superstitiosus 108-109
Delphax furcifera 426
Deltocephalus dorsalis 420 Echinochloa 740
Deois 740 Echinochloa colona 368, 373, 384, 394, 412, 415, 420,
Deraecoris vittatus 110-111 430, 432, 437, 442, 446, 452, 454, 457
Diabrotica 740 Echinochloa crus-galli 392, 394, 396, 407, 415-417, 428,
Diacamma sp. 191, 193, 196, 283, 286 430, 454, 457, 468, 470
Diacamma vagans 283, 286 Echinochloa frumentacea 407
Diadegma akoensis 223, 226, Echinochloa holubii 459
Diatraea 562, 735, 737-741 Echinocnemus 553
Diatraea auricilia 390 Echinocnemus bipunctatus 384
Diatraea polychrysa 392 Echinocnemus oryzae 165, 167, 383
Diatraea rufescens 240 Echinocnemus squamerus 165, 167, 384, 716-717
Diatraea saccharalis 68-69, 240, 388-889, 392, 397, 527, Echthorodephax fairchildi 184, 189, 596
737 Eclipta alba 660
Diatraea spp. 734, 736, 738 Ectomocoris atrox 118, 121
Diatraeophaga sp. 28, 30 Ectomocoris biguttulus 118, 121
Dichanthium annulatum 368 Ectrychotes crudelis 113-114
Dicladispa 176 Egadroma quinquepustulata 156, 159
Dicladispa armigera 176, 178, 180, 457-459, 527 Elasmopalpus 561-562, 740
Dicladispa gestroi 459 Elasmopalpus lignosellus 68-69, 389, 527, 736-137
Dicladispa striaticollis 178, 180 Elasmus albopictus 266, 271
Dicladispa viridicyanea 178, 180, 459 Elasmus claripennis 268, 271
Diectomis fastigiata 410 Elasmus corbetti 268, 271
Digitaria ciliaris 430 Elasmus philippinensis 268, 271
Digitaria sanguinalis 368, 382, 452, 468 Elenchus japonicus 14, 16
Digitaria zeyheri 459 Elenchus spp. 280, 284
Dimorphopterus cornutus novaeguineae 123, 125, 127 Elenchus yasumatsui 14, 16, 596
Diopsis apicalis 33-34, 378, 496 Eleusine coracana 368, 391, 407, 432, 441-442
Diopsis macrophthalma 33-34, 376-378, 527 Eleusine indica 367, 415-416, 454
Diopsis sp. 496 Empoasca kraemeri 637
Diopsis thoracica 248, 376 Empoascanara maculifrons 85, 87-89, 526
Diplachne fusca 459 Eniscospilus sokaguchii 221, 223-224
Diplacodes trivialis 131, 134 Enoplognatha japonica 304, 308
Diplocheila polita 154, 157 Enosima vectiferella 398
Diplonychus rusticus 95-96 Entomophaga grylli 638
Diploxys fallax 99, 102 Entomophthora aphidis 636
Diuraphis noocia 463 Entomophthora sphaerosperma 637
Dolomedes angustivirgatus 305, 308 Ephestia cautella 640
Dolomedes sulfureus 307, 310 Epilachna sp. 625
Dolycoris baccarum 464 Eragrostis aethiopica 459
Draeculacephala 735, 740 Eragrostis ferruginea 430
Drapetis (Drapetis) sp. 24-25 Eragrostis heteromera 456
768 SPECIES INDEX
Eragrostis interrupta 368 Fusarium oxysporun 632

Eragrostis nigra 470
Eretes sticticus 162-163 Galleria mellonella 645
Eriborus sinicus 227, 230, 586-587 Gambrus ruficoxatus 220, 222
Eriborus terebrans 227, 230 Gambrus wodai 220, 222
Eriborus vulgaris 227, 230 Gasteracantha mammosa 331-332
Erigone prominens 317, 321 Gasteracantha sp. 331-332
Erigonidium graminicola 319, 321 Gastrimargus marmoratus grandis 45, 47
Erynia delphacis 617, 636-637 Gastrimargus sp. 409
Erynia fumosa 637 Gea sp. 331-332
Erynia neoaphidis 636-637 Geocoris flaviceps 125-127
Erynia radicans 637 Geocoris jucundus 125, 127
Erynia radicans cicadelliphaga 637 Geocoris ochropterus 125, 127
Euborellia annulata 130, 132 Geocoris sp. 125, 127
Euborellia philippinensis 130, 132 Geoica lucifuga 80-82, 366
Eucoilidea sp. A 200, 202 Geoica sp. 366
Eucoilidea sp. B 200, 202 Geoica utricularia setariae 366
Eucolliuris sp. 148, 151 Geotomus pygmaeus 99- 100
Euconocephalus varius 39, 41-42 Gerris adelaides 95, 98
Euetheola 734-737, 740-741 Gesonula mundata 50, 52
Euetheola spp.. 736 Gibellula leiopus 620, 629, 638
Eumenes campaniformis 195, 197 Gibellula pulchra 620, 626, 638
Euplectrus chapadae 274, 278 Gibellula spp. 638
Euplectrus sp. 274, 276 Glyceria tonglensis 461
Eurytoma spp. 277, 281 Glycine max 382
Euschistus 734 Gnathonarium gibberum 319, 323
Euscyrtrus concinnus 39-40 Gonatocerus cicadellae 260, 262, 264
Euspudaeus sp. 591 Gonatocerus cincticipitis 260, 264
Exorista xanthaspis 33-34 Gonatocerus devitatacus 257, 270
Exoryza schoenobii 205, 207-208 Gonatocerus miurai 260, 264
Eysarcoris capitatus 101-102 Gonatocerus munarus 257, 260, 262
Eysarcoris guttiger 99, 102 Gonatocerus narayani 257, 261
Eysarcoris lewisi 464 Gonatocerus spp. 257, 596
Eysarcoris montivagus 99, 102 Gonorocerus urtardecanus 257, 261
Eysarcoris parvus 101-102, 464 Gonatopus lucidus 186, 190
Eysarcoris 26, 99, 236, 242, 245 Gonatopus yasumatsui 186, 189
Eysarcoris ventralis 101-102, 464 Goniozus nr. triangulifer 184, 189, 594
Gonista bicolor 43, 47
Festuca parvigluma 430, 452 Gonocephalum spp. 181-182, 185
Fimbristylis acuminata 450 Goryphus apicalis 221-222
Fimbristylis miliacea 368, 412, 454 Goryphus basilaris 220, 222
Fimbristylis tenera 368, 383 Goryphus lemae 221, 224
Forda sp. 366 Goryphus mesoxanthus mesoxanthus 221, 224
Formicomus sp. 182, 185 Gossypium hirsutun 409
Fragaria vesca 382 Gregopimpla kuwanae 218-219
Frankliniella tenuicornis 57, 60 Gryllotalpa africana 38, 40, 364
Fromicomus braminus 182, 185 Gryllotalpa orientalis 38, 40, 364
Fusarium 618, 632 Gryon nixoni 245, 247
Fusarium aleyrodis 632
SPECIES INDEX 769
Hahnio sp. 301, 303 Hirsutella spp. 621

Halictophagus bipunctatus 11–18, 280, 284 Hirsutella strigosa 629–630
Halictophagus spectrus 11–18, 280, 284 Hispa aenescens 451
Halictophagus thaiae 14, 18 Hispa armigera 451
Halydoia luteccomis 26–28 Hispa stygia 178, 180
Haplogonatopus apicalis 186, 190, 596 Hispellinus moestus 178, 180
Haplogonatopus atratus 188, 190 Holotrichia longipennis 381
Haplogonatopus oratorius 188, 190 Holotrichia seticollis 381
Haplogonatopus spp. 596 Hoiotrichia spp. 166, 169
Haplothrips aculeatus 55–56, 410 Hordeum vulgare 378, 401, 420, 442, 452, 468
Haplothrips ganglbaueri 55, 58, 470 Hotridipamera nietneri 123–124
Harmochirus brachiatus 283, 285, 281 Hortensia 135, 131, 140
Harmonia octomaculata 112, 174 Hortensia similis 136
Hasarius adansoni 288, 292 Hydaticus rhantoides 164, 167
Hebrus bergrothi 91, 100 Hydaticus vittatus 162–163
Heliothis (Helicoverpa) armigera 13–14, 218, 248 Hydrellia 200, 734–735, 131–138, 141–742, 144–748
Heliothis (Helicoverpa) punctigera 13–14 Hydrellia griseola 36–37, 219, 452, 454, 716,131
Heliothis zea 642 Hydrellia philippina 9, 36–31, 213, 270, 454–456, 527, 541,
Helodytes 739 604–605, 682
Hemarthria compressa 406–401 Hydrellia sasakii 36–37, 456
Hemiptarsenus sp. 214, 278 Hydrellria spp. 736, 738, 740
Herpetogramma licarsisalis 634, 643 Hydrellia tomiokai 36–31
Herperogramma (= Pachyzancla) stultalis 201 Hydrocharis morsus-ranae 452
Heterococcus rehi 368 Hydrolea zeylanica 410
Heteronychus lioderes 165, 161, 381 Hydrometra lineata 91, 100
Heteronychus mosambicus 381 Hydrometra orientalis 91, 100
Heteronychus plebeins 381 Hydrobomidius molitor 164, 161, 385
Heteropoda venatoria 288, 290, 292 Hydrophilus acuminatus 168–169
Heteropogon contortus 313 Hydrophilus nr. bilianeatus cashmirensis 168, 171
Heteroptermis respondens respondens 45, 41 Hygroplitis russatus 211, 214
Heteroptermis respondens 409 Hymenachne acutigluma 392
Heterorhabditis sp. 645 Hymenachne pseudointerrupta 370
Hibiscus esculentus 310 Hyparrhenia chrysargyrea 410
Hieroglyphus africanus 411 Hyparrhenia rufa 406
Hieroglyphus annulicornis 51, 53–54 Hypomeces nr. squamosus 165, 167
Hieroglyphus banian 51, 53, 408–409 Hypsosinga pygmaea 335, 338
Hieroglyphus daganensis 51, 53, 409–410 Hypsosinga sanguinea 335, 338, 340
Hieroglyphus furcifer 408 Hysteroneura setariae 81, 84, 462–463
Hieroglyphus nigrorepletus 51, 54, 408–410
Hieroglyphus oryzivorus 51, 53, 409 Imperata cylindrica 368, 430, 439, 441, 410
Hieroglyphus tonkinensis 51, 53 Inazuma dorsalis 420
Hikona formosana 422 Isachne dispar 432, 431
Hippasa greenalliae 301, 310 Isachne globosa 450
Hippasa holmerae 301, 310 Ischaemum aristatum 313
Hipppodamia variegata 175, 177 Ischatmum indicum 368, 415, 450, 470
Hirsutella 618 Ischaemum rugosum 367
Hirsutella citriformis 618–619, 622, 621–632 Ischaemum travancorense 460
Hirsutella gigantea 630 Ischnojoppa luteator 218–219
Hirsutella jonesii 629, 638 Ischnura senegalensis 133, 131
770 SPECIES INDEX
Iseilema laxum 368 Leucauge bengalensis 328, 332

Isotima chilonis 221, 224 Leucauge blanda 328, 332
Isotima dammermani 221, 224 Leucauge celebesiana 328, 330, 332
Isotima javensis 221, 224 Leucauge decorota 328, 332
Itoplectis narangae 217, 219 Leucopholis 26
Iva ciliata 382 Leucopholis irrorata 166, 169, 381
Leucopholis sp. 168-169
Laccophilus parvulus 160, 163 Levuana iridescens 588
Laccophilus sharpi 160, 163 Limnogonus fossarum 95, 98, 592, 599
Lagynotomus elongatus 464, 715 Limnogonus nitidus 97-98
Lamium album 452 Limnogonus parvulus 95, 98
Lampronota mandschurica 215-216 Lisarda sp. 116, 119
Laodelphax striatella 421 Lissorhoptrus 737, 739-740
Laodelphax striatellus 90-93, 184, 186, 188, 421, 526, 530, Lissorhoptrus brevirostris 388, 633
659, 715-716, 718 Lissorhoptrus oryzophilus 164, 167, 386, 499, 527,
Laphygma frugiperda 447 688-689, 718, 720, 126, 734, 737
Lurinia sp. 333, 336 Lissorhoptrus spp. 736
Larra carbonaria 193, 196-197 Livistona chinensis 441
Larra sanguinea 193, 195, 197 Lixophaga diatraeae 33-34
Leersia hexandra 373, 415, 418-420, 422-423, 432, 437, Locusta migratoria 495-496
446 Locusta migratoria manilensis 45, 47
Leersia japonica 417, 430, 456 Locusta migratoria migratorioides 246
Leersia oryzoides var. japonica 417, 456, 461 Lolium perenne 452
Leersia sp. 416 Loxoblemmus haani 38, 40
Lema oryzae 461 Lychnis dioica 452
Lema suvorovi 461 Lycosa chaperi 313, 316
Lemna minor 452 Lycosa pseudoannulata 520-521, 591, 596, 598, 605, 660,
Leptinotarsa decemlineata 625 665, 671, 684-685, 689
Leptispa pygmaea 176, 180, 460-461 Lycosa sp. 313, 316, 599, 694
Leptochloa chinensis 368, 446, 456, 466 Lycosa spp. 601-602
Leptochloa filiformis 397
Leptochloa panicoides 401 Macrocentrus philippinensis 213, 216, 594
Leptocorisa 126, 464, 466 Macrosiphum (Sitobion) avenae 81-82
Leptocorisa acuta 128, 132, 464, 466 Macroteleia crawfordi 246, 248-249
Leptocorisa biguttata 128-129 Macrotermes bellicosus 280, 284
Leptocorisa chinensis 128, 132, 464 Macrotermes gilvus 182, 187, 280, 284
Leptocorisa costalis 128-129 Mactrotermes natalensis 182, 187
Leptocorisa luzonica 128-129 Macrotermes sp. 182, 187
Leptocorisa oratorius 128-129, 245, 464, 527, 618-619, Maliarpha separatella 68-69, 242, 388, 398-400, 494, 496,
632, 672, 689 499, 527, 563
Leptocorisa palawanensis 126, 129 Marasmia 63, 184, 436, 735, 740
Leptocorisa pseudolepida 128-129 Marasmia exigua 63, 65, 435-436, 586
Leptocorisa solomonensis 126, 129 Marasmia patnalis 65-66, 436, 669
Leptocorisa spp. 464-465, 671, 697 Marasmia ruralis 63, 65, 436, 669
Leptocorisa tagalica 128-129 Marasmia sp. 634, 636-637
Lespedeza striata 382 Marasmia spp. 205, 207, 213, 225, 227, 266, 700
Leucania inferens 407 Marasmia venilalis 436
Leucania separata 442 Marpissa elongata 285, 289
Leucas aspera 470 Marpissa magister 285, 289
SPECIES INDEX 771
Medythia nigrobilineata 179-180, 205 Mocis 735, 737, 740, 748

Megamilus furcifera 426 Mocis frugalis 72, 74, 634, 643
Megaselia scalaris 24-25 Monochoria vaginalis 463
Melanaphis sacchari 81-82, 463 Monolepta bifasciata 179, 183
Melanitis 57, 236, 242 Monolepta signata 179, 183
Melanitis leda ismene 439, 632, 642 Monomorium pharaonis 282, 284
Melanitis leda 59, 62, 242 Montandoniola moraquesi 110-111
Melanitis phedima 59, 62 Monnoidea poecilia 245
Melia azadirach 660 Muhlenbergia mexicana 382, 452
Menida formosa 104, 107 Mycalesis 57
Menochilus sexmaculatus 175, 177 Mycalesis gotama 57, 62
Mermis sp. 645 Mycalesis nr. mineus 57, 59, 62
Mesovelia sp. 599 Mymar taprobanicum 260, 265
Mesovelia vittigera 120-121, 592
Mythimna 73, 236, 242, 551, 643
Metarhizium 618, 621, 672
Mythimna loreyi 78-79, 227
Metarhizium almun 616, 618, 620-621
Mythimna roseilinea 78-79
Metarhizium anisopliae 617-622, 625, 627, 632-633, 672
Mythimna separata 76, 79, 251, 442-444, 527, 620,
Metarhizium anisopliae var. anisopliae 618, 621 643-644, 689, 700
Metarhizium anisopliae var. majus 616, 618
Mythimna spp. 634
Metarhizium brunneum 621
Mythimna unipuncta 211, 242, 445
Metarhizium flavoviride var flavoviride 616
Mythimna venalba 76, 79
Metarhizium flavoviride var. minus 616, 618, 620-621,
Myzus persicae 463
623, 627
Metarhizium spp. 621, 632
Nabis capsiformis 122, 124
Metioche vittaticolis 39, 42, 591, 594, 603
Nabis sinofem 120, 121
Metopiusa rufus browni 218-219
Metopolophium dirhodum 463 Nabis stenoferus 122, 124
Mezira sp. 638 Naranga 570
Micraspis crocea 594-595, 598 Naranga aenacens 72, 74, 211, 217, 227, 486, 506,
Micraspis discolar 172-174 604-605, 624, 634
Micraspis hirashimai 173-174 Naranga diffusa 225, 446
Micraspis inops 113-174 Neanastatus cinctiventris 277, 281
Micraspis sp. 599 Neanastatus oryzae 217, 281
Micraspis spp. 592 Nemorilla maculosa 27-28
Micraspis vincta 173-174 Neoaplectana carpocapsae 645
Micraspis yasumatsui 173-174 Neocutilla 735, 740
Micronecta quadristrigata 92, 94, 96 Neorhizoecus epicopus 369
Microplitis pallidipes 213-214 Neoscona doenitzi 333, 336
Microplitis sp. 213-214 Neoscona nautica 335, 337
Microplitis tuberculifer 213-214 Neoscona scylla 335, 337
Microtormes sp. 280, 284 Neoscona theisi 333, 335-336
Microvelia atrolineata 660, 671, 689 Neozygites fumosa 637
Microvelia douglasi atrolineata 97, 100, 192, 596, 598, Nephotettix 280, 415, 418-419, 529, 555
603 Nephotettix afer 413, 415, 420
Microvelia douglasi douglasi 97, 100, 598 Nephotettix apicalis 415
Miscanthus sinensis 441 Nephotettix bipunctatus 416
Misumena sp. 293, 296 Nephotettix cincticeps 24, 85-87, 91, 186, 413-414,
Misumenops sp. 293, 296 417-419, 533-534, 667-668, 716-718, 722-723
Misumenops tricuspidatus 293, 296 Nephotettix impicticeps 416
772 SPECIES INDEX
Nephotettix malayanus 83, 86-87, 89, 413-415, 418-419, Odontomachus spp. 283, 286
506, 529 Odontoponera sp. 191, 194
Nephotettix modulatus 85-86, 90-91, 413, 415, 419-420 Odontoponera transversa 283, 286
Nephotettix nigropictus 17, 85-86, 90-91, 186, 413-415, Oebalus 236, 468, 735, 740-741
417-418, 420, 506, 529, 597, 667 Oebalus ornata 460-469
Nephotettix parvus 85-87, 91, 413-414, 419 Oebalus poecila 101-102, 468
Nephotettix sp. 660 Oebalus pugnax 101-102, 245, 468-469
Nephotettix spp. 22, 184, 188, 413-414, 585, 596-597, 599 Oebulus spp. 468, 736
Nephotettix virescens 10, 17, 85-87, 89, 414-417, 419, 496, Oedaleus abruptus 45, 47
506, 521, 523, 526, 529, 533-535, 537-538, 563, 597,
Oligosita 253
617-618, 625-626, 629-630, 645, 663-664, 667, 682,
Oligosita aesopi 255, 259
684-685, 688, 705
Oligosita brevicilia 255, 258
Neurothemis palliata 133-134
Oligosita consanguinea 253, 256
Neurothemis terminata terminata 133-134
Oligosita manii 255, 258
Neurothemis tullia tullia 133-134
Oligosita naias 255, 258
Nezara 242, 644, 735, 739-740
Oligosita nephotettica 255, 259
Nezara antennata 106-107, 236, 245, 464
Nezara viridula 106-107, 236, 243, 245, 464, 467-468 Oligosita shibuyae 255, 258
Nicotiana tabacum 442 Oligosita yasumatsui 255, 258, 596
Nilaparvata 434 Oncocephalus impudicus 116, 119
Nilaparvata lugens 10, 90-93, 364, 422-426, 428, 506, Oncocephalus pacificus 116-117, 119
519-520, 522, 526, 537, 563, 585, 596-597, 618, 620, Onychilus 739
627, 645-646, 659, 663-664, 666, 681-686, 688, Ooctonus sp. 257, 259
693-707, 717-718, 720 Ooencyrtus malayensis 277, 281
Nilaparvata oryzae 422 Ootetrastichus formosanus 270-271
Nilaparvata sordescens 422 Ophionea 592
Nilobezzia acanthopus 19-20 Ophionea indica 150, 153
Ninus insignis 125, 127 Ophionea interstitialis 150, 153
Niphadoses 389 Ophionea ishii hoashii 150, 153
Niphadoses gilviberbis 389 Ophionea ishii ishii 150, 153
Niphadoses palleucus 389 Ophionea nigrofasciata 150, 153, 599
Nomuraea 618 Ophionea sp. 595
Nomuraea rileyi 617, 424, 626, 634-635 Opius barrioni 213, 216
Nonagria inferens 407 Orgilus ashmeadi 205, 208
Notiphila 36, 200 Orius tantillus 110-111
Notiphila latigenis 31-38 Orseolia oryzae 11, 19, 221, 372-373, 527, 532, 537, 670,
Notiphila sekiyai 36-37 682, 686, 716, 723, 726
Notiphila similis 36-37 Orseolia oryzivora 17, 19, 272, 375-376
Nymphula depunctalis 9, 63-64, 437-438, 506, 527, 541, Orthetrum sabina 131-132
669, 689 Orthetrum testaceum 131-132
Nymphula nymphaeata 438 Oryctes 642
Nymphula stagnalis 437 Oryza 3, 524
Nymphula vittalis 439 Oryza alta 4
Nysius plebejus 464 Oryza australiensis 4, 541
Oryza barthii 4-5, 373, 396, 541
Ochetina 140 Oryza brachyantha 4, 541
Ochterus marginatus 92, 96 Oryza breviligulata 4
Ochthera brevitibialis 35, 37 Oryza eichingeri 4, 531, 541
Ochthera sauteri 35, 37 Oryza fatua 4
Odacantha sp. 150-151 Oryza glaberrima 3-5, 401, 525, 541
SPECIES INDEX 713
Oryza glumaepatula 4 Pachygnatha clercki 322,325

Oryza grandiglumis 4 Paecilomyces 632
Oryza granulata 4 Paecilomyces farinosus 619, 624, 628, 632, 635
Oryza latifolia 4, 401, 531 Paecilomyces fumosoroseus 632, 635
Oryza longiglumis 4 Paecilomyces lilacinus 623, 626, 629, 632-633
Oryza longistaminata 4-5, 541 Paecilomyces spp. 632
Oryza officinalis 4, 531, 541 Paecilomyces tenuipes 619, 635
Oryza meridionalis 4 Paederus fuscipes 175, 177, 592, 596, 599, 602
Oryza meyeriana 4 Paederus tamulus 175,177
Oryza minuta 4-5, 531, 541 Palexorista lucagus 31-32
Oryza nivara 4-5, 400, 541 Palexorista solensis 31-32
Oryza perennis 4 Pamerana nigritula 122, 124
Oryza punctata 4, 541 Panicum amplexicauIe 370
Oryza ridleyi 4, 541 Panicum antidotale 442
Oryza rufipogon 4-5, 400, 541 Panicum dichotomiflorum 468
Oryza sativa 3-5, 525, 531, 540-541, 543 Panicum maximum 407, 439
Oryza schlechteri 4-5 Panicum miliaceum 373
Oryza spontanea 4-5 Panicum ramosum 416
Oryza stapfii 5 Panicum repens 368, 379, 430, 437, 450, 452, 457
Oryzae 704 Panoquina spp. 736
Oryzophagus 739 Pantala flavescens 130, 132
Ostrinia fumacalis 225 Paracentrobia andoi 253, 256
Ostrinia nubilalis 217, 625, 635 Paracentrobia yasumatsui 253, 256
Otiorhynchus sulcatus 645 Paracletus cimiciformis 366
Oulema 178, 552 Paraeucosmetus pallicornis 123-124
Oulema astrosuturalis 179-180 Parahieroglyphus bilineatus 50, 52-53
Oulema melanopa 179-180, 221 Paralauterborniella subcincta 371
Oulema oryzae 179-180, 279, 461-462, 716, 718 Paraplea liturata 94, 96
Oxya 46 Paraplea sobrina 94, 96
Oxya chinensis 48-49, 409, 411 Paralanytarsus sp. 311
Oxya hyla intricata 46, 49, 638 Paratheresia clanpalpis 587
Oxya intricata 409 Pardosa 307
Oxya japonica japonica 46, 49 Pardosa annandalei 311, 314
Oxya japonica 246 Pardosa astrigera 309, 312, 669
Oxya minuta 46,49 Pardosa birmanica 311, 314
Oxya velox 46, 49, 246, 411 Pardosa isago 309, 312
Oxya yezoensis 48-49 Pardosa laura 307, 309-310
Oxyopes javanus 299, 302 Pardosa oakleyi 311, 313-314
Oxyopes lineatipes 301-302 Pardosa palustris 309-310
Oxyopes macilentus 299, 302 Pardosa pseudoannulata 309, 311-312
Oxyopes sertatus 301-302 Pardosa sumatrana 311, 314
Oxyopes sp. 671 Pardosa sutherlandi 311-312, 314
Oxyopes spp. 596 Pamara 28
Pamara ganga 61, 64
Pachybrachius inortatus 123-124 Parnara guttata 61, 64, 217, 268, 440-442, 641, 716
Pachybrachius nigriceps 123-124 Parnara mathias 635
Pachybrachius pacificus 123-124 Parnara naso bada 61, 64
Pachybrachius sobrinus 123-124 Parnara spp. 641
Pachydiplosis oryzae 372 Paspalum conjugatum 406, 437, 446, 450
774 SPECIES INDEX
Paspalum distichum 489 Pipunculus orientalis 22-24

Paspolum laeve 382 Pipunculus roralis 22-23
Paspalum paspalodes 457 Pipunculus spp. 596
Paspalum scrobiculatum 367-368, 373, 407, 415, 430, Pirata clercki 317, 320
441, 452, 454, 460, 470 Pirata meridionalis 315, 318
Paspalum thunbergii 407 Pirata piraticus 315, 318
Paspalum urvillei 397, 468 Pirata piratoides 315, 316, 320
Patanga succincta 48, 52 Pirata procurvus 317, 320
Pediobius atamiensis 268, 271 Pirata subpiraticus 315-316, 318, 665
Pediobius mitsukurii 268, 271 Pirates arcuatus 118, 121
Peirbaea orbata 31-32 Pirates atromaculatus 118-119
Pelopidas 28, 59 Pirates sparsus 118, 121
Pelopidas agna 61-62 Pisaura lama 305, 308
Pelopidas cinnara 223 Pistacia 366
Pelopidas mathias 61-62, 215, 223, 221, 440-442 Platygaster foersteri 235, 237
Penicillium 618 Platygaster oryzae 233, 237, 670
Pennisetum japonicum 430 Platygaster sp. 665
Pennisetum pedicellatum 432 Platyscelio abnormis 246, 249
Pennisetum purpureotyphoides 460 Platysentra (Cirphis) compta 646
Pennisetum purpureum 406, 442 Plebeiogryllus plebejus 39-40
Pennisetum subangustum 367, 406, 410 Plexippus setipes 288, 292
Pennisetum typhoides 409, 470 Plexippus paykulli 288-289
Pentagonica nr. erichson 154, 157 Plodia interpunctella 640
Pentagonica ruficollis 154, 157 Plutella xylostella 209
Phalaris arundinacea 418, 452 Poa annua 418
Phaneroptera furcifera 41-42 Poa compressa 382, 452
Phaneroptera gracilis 41-42 Poecilochroa sp. A 295-296
Pheidole sp. 282, 284 Poecilochroa sp. B 295-296, 298
Pheidologeton sp. 193, 196, 282, 284 Poecilotraphera taeniata 33-34
Pherosophus javanus 145, 148-149 Polididus armatissimus 113-114
Pheropsophus jessoensis 148, 153 Polygonum lapathifolium 382, 452
Pheropsophus occipitalis 148, 151 Polynema sp. 260, 265
Pheropsophus sp. A 145, 149 Polypogon fugar 407, 452
Pheropsophus sp. B 145, 149 Polyrachis spp. 191, 194, 282
Phidippus sp. 288, 292 Polytoxus fuscovittatus 112, 114
Philodromus spinitarsis 290, 292 Polytoxus selangorensis 112, 114
Phleum paniculatum 418, 452 Polytoxus sp. A 112, 114
Phragmatiphila 388-389 Polytoxus sp. B 113-114
Phragmites australis 394, 418, 430, 452, 461 Polytremis pellucida 61-62
Phragmites communis 440 Polytrias amaura 470
Phragmites karka 407 Popillia cupricollis 381
Phyllophaga 734-737, 740-741 Potamogeton natans 384
Physopleurella armata 112, 114 Potanthus confucius 59, 62
Piezodorus 242, 245 Potentilla bicolor 382
Piezodorus guildinii 236 Proboscidocoris sp. 110
Piezodorus hubneri 104, 107 Proboscidocoris tibialis 108, 111
Piezodorus rubrofasciata 236 Proceras africana 396
Pipunculus javanensis 22-23 Proceras indicus 389
Pipunculus mutillatus 23-24 Proceras polychrysa 392
SPECIES INDEX 775
Prociphilus sp. 366 Rhinocoris sp. A 115, 117

Prodenia litura 449 Rhinocoris sp. B 115, 117
Propicroscytus mirificus 277, 279, 281 Rhopalosiphum maidis 83-84, 463
Propylaea japonica 173, 177 Rhopalosiphum nymphaeae 83-84, 463
Proreus simulans 130, 132 Rhopalosiphum padi 463
Prosena siberita 26-27 Rhopalosiphum rufiabdominalis 83-84, 366-367
Prunus padus 463 Rhopalosiphum sp. 463
Prunus sp. 366 Rhyzopertha dominica 527
Prunus spp. 463 Ripersia oryzae 368
Psamotettix striatus 22 Ripersia rehi 368
Pseudagrion pilidorsum pilidorsum 135, 137 Ripersia sacchari 368
Pseudaletia separata 442 Riptortus 126
Pseudaletia unipuncta 443, 643 Riptortus linearis 126, 129
Pseudococcus saccharicola 80, 82 Riptortus pedestris 126, 129
Pseudogonatopus flavifemur 188, 192 Riptortus pilosus 126, 129
Pseudogonatopus fulgori 188, 192 Rivula 570
Pseudogonatopus hospes 191-192 Rivula atimeta 72, 74, 447, 604-605, 626, 634
Pseudogonatopus nudus 188, 192 Rivula basalis 447
Pseudogonatopus sarawaki 188, 190 Rogas narangae 203, 206
Pseudogonatopus spp. 596 Rogas spp. 203, 206
Pseudonapomyza asiatica 34-35 Romanomermis culicivorax 645
Pseudoperichaeta insidiosa 29, 32 Ropalidia cyathiformis 198-199
Psilopa spp. 35-36 Ropalidia fasciata 195, 197
Psix lacunatus 235, 237, 587 Ropalidia flavopicta flavobrunnea 198-199
Pygolampis sp. 115, 117 Ropalidia marginata sundaica 195, 197
Pygomenida 104 Ropalidia nr. philippinensis 195, 197
Pygomenida benghalensis 106-107 Ropalidia sp. A 198-199
Pygomenida varipennis 104, 107 Ropalidia sp. B 198-1 99
Pyracantha coccina 382 Ropalidia sp. C 198-199
Pyrgomorpha conica 409 Rottboellia exaltata 406, 410
Pyricularia oryzae 399 Rumex crispus 382
Runcinia albostriata 291, 294
Quilta 45 Runcinia roonwali 291, 294
Quilta mitrata 46-47 Runcinia sp. 291, 294
Quilta oryzae 46-47 Rupela 740
Rupela albinella 66-67, 388-389, 405, 527, 736
Ranatra diminuta 95, 98
Recilia 280 Saccharum arundinaceum 407
Recilia dorsalis 85–87, 89, 186, 420-421, 526, 529, 618 Saccharum officinarum 390-392, 397, 406-407, 409,
Regillus sp. A 293-294 411-412, 415-416, 420, 432, 439, 460
Regillus sp. B 293-294 Saccharum sp. 450
Rhaconotus oryzae 203, 206 Saccharum spontaneum 368, 407
Rhaconotus schoenobivorus 203, 206 Sacciolepis interrupta 373
Rhoconotus scirpophagae 203, 206 Sagittaria 463
Rhagadotarsus sp. 95, 98 Sagittaria latifolia 452
Rhene sp. 285, 289 Saldula omatula 120-121
Rhinaphe pallidicosta 398 Sastrapada entomophaga 116, 119
Rhinaphe vectiferella 398 Scadra sp. 113-114
Rhinocoris fuscipes 115, 117 Scambus annulitarsis 218-219
776 SPECIES INDEX
Scaphoideus morosus 85, 87-89 Secale cereale 378, 452, 468

Scarites sp. A 152, 155 Serratia marcescens 642
Scarites sp. B 152, 155 Sesamia 551, 562
Scelio muraii 246 Sesamia botanephaga 78-79, 389
Scelio tsuruokensis 246-247 Sesamia calamistis 78-79, 211, 388-389, 405-406, 493,
Scelio zolotarevskyi 246, 249 496, 527
Scelipron modraspatanum conspidam 193, 196 Sesamia cretica 211
Schistocerca americana gregaria 48, 52 Sesamia inferens 78-79, 201, 203, 215, 217, 221, 227, 270,
Schizapis graminum 81, 84, 463 389, 407, 640, 643
Schoenobius incertulas 9, 201, 400 Sesamia sp. 394
Scipinia horrida 113-114 Sesamia vuteria 405
Scirpophaga 494, 551 Sesselia pusilla 179, 183
Scirpophaga albinella 405 Setaria 740
Scirpophaga auriflua 238 Setaria glauca 368, 452
Scirpophaga gilviberbis 66, 69 Setaria italico 407, 409, 443
Scirpophaga incertulas 10, 66-67, 203, 207, 215, 217, 221, Setaria pallide-fusca 392
223, 227, 238, 388-389, 400-403, 494, 506, 520, 526, Setaria sphacelata 410
530, 563, 565, 587, 590-591, 640, 646, 668, 682, 687, Setaria splenldida 406
689, 715 Singa hamata 335, 338
Scirpophaga innotata 9, 66-67, 215, 217, 221, 227, 238, Sipha 740
389, 403, 489, 555, 562, 564, 566-567, 687 Sipha flava 81-82
Scirpophaga melanoclysta 240 Sipha glyceriae 463
Scirpophaga nivella 66-67, 217, 220 Sirthenea flavipes 116, 118-119
Scirpophaga occidentella 68-69 Sitobion akebiae 463
Scirpophaga spp. 225 Sitobion avenae 463
Scirpophaga virginia 66-67 Sitobion fragariae 463
Scirpus articulatus 412 Sitobion graminis 463
Scirpus grossus 370, 392, 452 Sitobion miscanthi 463
Scirpus maritimus 407 Sitophilus oryzae 527, 542
Scleria sumatrensis 370 Sitophilus zeamais 527
Sclerostachya fusca 394, 407 Sitotroga cerealella 527
Scolia nr. scutellaris 229-230 Snellenius manilae 211, 214
Scolia sp. 229-230 Sogata distincta 426
Scotinophara 101, 242, 245, 311, 633 Sogata furcifera 184, 186, 188, 191, 426, 521, 627, 645,
Scotinophara bispinosa 104, 107 682-684, 717-718
Scotinophara coarctata 103, 105, 235, 243, 369-371, 587, Sogata pallescens 426
619, 626, 632 Sogatello 434
Scotinophara horvathi 103, 105 Sogatella furcifera 90-92, 426-428, 526, 529, 596-597,
Scotinophara inermiceps 103, 105 626, 659, 718, 720
Scotinophara latiuscula 104-105, 235, 243, 369-371, 527, Sogatella kolophon 426
587 Sogatella nigeriensis 426
Scotinophara lurida 103, 105, 243, 369-371, 464, Sogatello vibix 426
632-633, 715 Solanum torvum 470
Scotinophara obscura 104-105 Solenopsis germinata 193, 196, 282, 284
Scotinophara ochracea 104, 107 Sorghum 443
Scotinophara parva 103, 105 Sorghum arundinaceum 396
Scotinophara scotti 103, 105 Sorghum bicolor 379, 390-391, 398, 406-407, 410, 412,
Scotinophara serrata 101, 105 432, 443, 446, 468, 470
Scotinophara spp. 618, 632 Sorghum halepense 397, 406, 409, 441
SPECIES INDEX 777
Sorghum verticilliflorum 439 Telenomus alecto 240-241

Spartina cynosuroides 391 Telenomus applanatus 240, 242, 244
Spathius helle 201, 204 Telenomus chloropus 243, 247
Spicaria densa 625 Telenomus cirphivorus 242, 244
Spicaria rubido-purpurea 633 Telenomus cristatus 243, 247
Spodoptera 73, 236, 242, 551, 734-735, 737, 740-741, 748 Telenomus cyrus 243-244, 587
Spodoptera abyssinia 75, 77 Telenomus dignoides 238-239
Spodoptera exempta 76-77 Telenomus dignus 238-239, 591
Spodoptera exigua 75, 77 Telenomus gifuensis 245, 247
Spodoptera frugiperda 73-74, 447-448, 734, 736-737 Telenomus globosus 240-241
Spodoptera litura 73-74, 209, 211, 213, 218, 242, 449, 643 Telenomus lucullus 242, 244
Spodoptera mauritia 227, 450-452 Telenomus nakagawaii 245, 247
Spodoptera mauritia acronyctoides 76-77, 242, 643 Telenomus podisi 245, 247
Spodoptera mauritia matuitia 76, 79 Telenomus remus 242, 244
Spodoptera pecten 75, 77 Telenomus rowani 236, 238-239, 588, 590-591
Spodoptera spp. 495, 634, 669, 700 Telenomus spp. 590
Spodoptera triturata 75, 77 Telenomus transversus 238, 241
Staccia diluta 116, 119 Telenomus triptus 243-244
Stariodes degenerus 106, 109 Telenomus versicolor 240-241
Steinemema feltiae 645-646 Teleogryllus mitratus 39-40
Stellaria media 452 Teleogyllus occipitalis 39-40
Stenchaetothrips biformis 55, 58, 60, 430-432, 527 Telicota augias 59, 62, 215
Stenobracon nicevillei 201, 204, 591 Temelucha biguttula 225-226
Stenocatantops splendens 50, 52 Temelucha philippinemis 225, 228, 591, 594
Stenocoris 464 Temelucha stangli 225-226
Stenodema sibiricum 464 Tetragnatha javana 324-325
Stenomesius japonicus 274, 276 Tetragnatha mandibulata 326-327, 329
Stenonabis tagalicus 120-121 Tetragnatha maxillosa 326, 329
Stephanodes imbricatus 263, 265 Tetragnatha nitens 324, 327-328, 330
Sternolophus rufipes 170-171 Tetragnatha praedonia 326, 329-330
Sternolophus sp. 170-171 Tetragnatha sp. 671
Stratiotes aloides 452 Tetragnatha spp. 596
Streblocera okadai 205-206, 208 Tetragnatha squamata 324-325, 327
Sturmiopsis inferens 29-30, 588 Tetragnatha virescens 324, 327
Submera latifrons 154, 157 Tetramorium pacificum 282, 284
Susumia exigua 435, 586-587 Tetraneura akinire 366
Sympiesis spp. 274, 278 Tetraneura basui 366
Syngamia 735, 737, 740 Tetraneura nigriabdominalis 80, 82, 366-367
Syngamia sp. 737-738 Tetraneura radcicola 366
Synharmonia octomaculata 594-596, 602 Tetraneura sp. 366
Tetrastichus ayyari 591, 594
Tagasta marginella 43-44 Tetrastichus confusus 272, 276
Tagosodes 735, 737, 740-741 Tetrastichus diopsisi 274, 276
Tagosodes cubanus 90, 92-93 Tetrastichus flavobrunneus 272-273
Tagosodes orizicolus 90, 92-93, 428-429, 526, 530, 734, Tetrastichus howardi 270, 273
736-739, 742, 748 Tetrastichus inferens 270, 273
Tanymeus indicus 165, 167 Tetrastichus pachydiplosisae 270, 272-273
Tanymecus nr. boencheri 164-165, 167 Tetrastichus risbeci 272, 276
Tapinoma melanocephalum 282, 286 Tetrastichus schoenobii 270, 273, 588, 590-591
778 SPECIES INDEX
Tetrastichus sp. 210, 213 Trichogrammatoidea 248

Tetrastichus spp. 590 Trichogrammatoidea armigera 248-449, 252
Tetrastichus variabilis 212, 214, 216 Trichogrammatoidea bactrae 250, 252
Tetrodontocheyls lucens 186, 189 Trichogrammatoidea lutea 250, 252
Tetrodontocheyls sakaii 186, 189 Trichogrammatoidea nana 248-249
Tettigella spectra 412 Trichogrammatoidea simmondsi 248-249
Tettigoniella spectra 412 Trichomalopsis apanteloctena 279, 281
Thaia oryzivora 14, 85, 88 Trichomalopsis deplanata 279, 284
Thaia spp. 81, 89 Trichomalopsis oryzae 219, 281
Thaia subrufa 85, 88 Trichomalopsis shirakii 279, 281
Thanatus sp. 290, 292 Trichomma (Trichomella) cnaphalocrosis 225, 223, 594
Thecocarcelia oculata 28, 30 Trifolia 463
Theridion octomaculatum 304, 306 Trifolium pratense 452
Theridiosoma sp. 322-323 Tripnotylus coelestialium 464
Thioptera 135, 131 Trilophida annulata 43, 47
Thomisus cherapunjeus 290-291, 294 Tripsacum laxum 406
Thomisus nr. labefactus 291, 294 Trissolcus basalis 236-231
Thomisus okirawensis 291, 294 Trissolcus hullensis 236, 239
Thrips dobrogensis 430 Trissolcus mitsukurii 235-237
Thrips holorphnus 430 Trissolcus solocis 236, 239
Thrips oryzae 430 Trissolcus thyantae 236-231
Tibellus oblongus 290, 292 Trithemis aurora 131, 134
Tibellus sp. 290, 292 Triticum aestivum 378, 391, 406-407, 420, 432, 443, 452,
Tibraca 134, 131 468
Tibraca limbativentris 136 Triticum vulgare 410
Tipula aino 17, 19 Trochosa sp. A 313, 316
Togo hemipterus 464 Trochosa sp. E 315-316
Tomosvaryella oryzaetora 22-23 Tropobracon schoenobii 201, 204, 591
Tomosvaryella spp. 596 Tryporyza incertulas 400
Tomosvaryella subvirescens 22-23 Tryporyza innotata 403
Tomosvaryella sylvatica 22-23 Tychea rehi 368
Trathala flavoorbitalis 225-226, 586-581 Tylorida striata 328, 330
Trichispa sericea 116, 180, 459-460 Typha latifolia 394, 452
Trichogramma 248, 673 Tytthus chinensis 110-111
Trichogramma chilonis 251, 254
Trichogramma dendrolimi 251, 254, 689 Ulmus sp. 366
Trichogramma evanescens 251, 254 Ummeliata (=Oedothorax) insecticeps 319, 323
Trichogramma fasciatum 253, 256
Trichogramma hesperidis 250, 252 Valanga nigricomis 50, 52
Trichogramma ivelae 251, 254, 256 Venturia sp. 223, 226
Trichogramma japonicum 250, 252, 586-591, 594, 689 Veronica officinalis 452
Trichogramma leucaniae 251, 254 Vetiveria zizanioides 391, 451, 460
Trichogramma nanum 588 Vigna unguiculanta 310
Trichogramma mwanzai 250, 252 Vitis vinifera 382
Trichogramma pallidiventris 250, 252 Vossia cuspidata 406
Trichogramma perkinsi 253,256
Trichogramma semifumatum 251, 254 Wendilgarda sp. 322-323
Trichogramma sp. 501
Trichogramma spp. 590, 593 Xanthopimpla enderleini 215-217
SPECIES INDEX 779
Xanthopimpla flavolineata 215–216, 594 Zeadiatraea lineolata 389

Xanthopimpla modesta modesta 216–217 Zeuxippa cawxantha 588
Xanthopimpla pedator 217, 219 Zincrona caerulea 106, 109
Xanthopimpla punctata 216–217 Zizania aquatica 391, 395, 452, 457
Xanthopimpla stemmator 217, 219 Zizania latifolia 461
Xenocotantops humilis 50, 52 Zizaniopsis miliacea 391
Xysticus sp. 293, 296 Zoophthora radicans 637, 639
Zoysia japonica 430
Zea mays 370, 379, 391–392, 394, 397, 406–407, 409, Zulia 740
411–412, 415–416, 430, 432, 441, 443, 450, 452, 468 Zygobothria ciliata 29, 32

Biology MGMT of Rice Insects PDF

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BIOLOGY AND

PUBLISHING FOR ONE WORLD

WILEY EASTERN LIMITED • NEW AGE INTERNATIONAL LIMITED

This book or any part thereof may not be

This book is not to be sold outside the

Production: M.I. Thomas

Baton Rouge, Louisiana E.A. HEINRICHS

II. BIOLOGY AND ECOLOGY

III. CONTROL TACTICS AND STRATEGIES

10. Integration of Rice Insect Control Strategies and Tactics . . . . . . . . . . . . . . . . . . . . . . . . . . . 681

IV. IMPLEMENTATION OF RICE IPM SYSTEMS

Remedios M. Aguda Michael E. Loevinsohn

Species name x =12 Distribution

4. DAMAGE CAUSED BY RICE INSECT PESTS

4.1 YIELD LOSSES

Region yield loss

4.2 FACTORS AFFECTING EXTENT OF INSECT-CAUSED YIELD LOSSES

5. DEVELOPMENT OF INTEGRATED PEST

Taxonomy of Rice Insect Pests and their

ALBERTO T. BARRION AND JAMES A. LITSINGER

2. KEY TO INSECTS AND SPIDERS

2(1) Wings present (with few veins or well developed) ............................ 3

3(2) With only 1 pair of wings ................................................. 4

4(3) Forewings reduced to a pseudohaltere, leather-like and without venation; functional

31' Face not as above, without such facial keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32' Not entirely as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

34' Hindcoxae bare posterodorsally; vein R1 setulose and vein R4 + 5 setulose up to or

53' Anal cell present; ocellar triangle less conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

60' Leaf miner; midtibiae with 3 extensor bristles (Fig. 112) . . . . . . . . . . . . . . . . . . . . . . 61

78' Pronotum rather smooth, non-tuberculated; wings of different colors . . . . . . . . . . 79

84' Pronotum without tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

108(107') Apices of male cerci notched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

ovipositor dissected ventrally; narrow wings pointed apically; antennae 6- to 9-seg-

126(125') Midtibiae smooth; wings with yellow markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

128' Midtibiae smooth and not spined ........................................ 130

131' Discal cells without prominent spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132

133(120') Maxillary palpi 4-segmented (sometimes 2-3); haustellum reduced; chaetosemata

134'' Wing coloration not as above; stem borers ................................ 139

137' Costa without strigulae ................................................. 138

164' Valves broad, not as slender as in Heliothis; vesica different . . . . . . . . . . . . . . . . . . . 166

169' Indentation present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171

182' Tarsi 3-segmented; abdomen never with siphunculi . . . . . . . . . . . . . . . . . . . . . . . . . 194

199' Anterior margin of pronotum with black markings .......................... 201

261' Postocular anterior margins of pronotum straight to slightly sinuate; anterolateral

277' Rostrum reaching hind-coxae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279

287 ' Ocelli present, without combination of above characters . . . . . . . . . . . . . . . . . . . . 291

291' Not as above: abdominal margins without spines . . . . . . . . . . . . . . . . . . . . . . . . . . . 292

292' Apex of scutellum triangular or subtriangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294

310' Arolia present; front legs and hemelytra variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315

337(335') Endocorium and clavus black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338

344' Cerci normal; wings and abdomen not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348

351' Distal ante-nodal nervure of fore-wings incomplete . . . . . . . . . . . . . . . . . . . . . . . . . 353

373(367') Small to moderately small tiger beetles, 7-12 mm long . . . . . . . . . . . . . . . . . . . . . . . 374

373' Large tiger beetles, 13-20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380

462(460') Tarsal segmentation 5-5-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463

493' Pronotum with a prominent transverse depression . . . . . . . . . . . . . . . . . . . . . . . . . . 495

535' Forewings with 3 cubital cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 551

538' Antennae of both sexes normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539

545' Areola and costulae absent ............................................. 547

583' Areolet absent on forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 587

612' Tergite I of abdomen with sublateral setae; notauli absent . . . . . . . . . . . . . . . . . . . . 614

627' Non-oriental (Palearctic, Nearctic and Neo-tropical) species of Telenomus attacking a

631' Not as above .......................................................... 632