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AGRICULTURAL
ENTOMOLOGY
AND
PEST CONTROL
S. PRAOHAN
ttl
leAR
.
" .' .
AGRICULTURAL ENTOMOLOGY
AND
PEST CONTROL
s. PRADHAN
Head of the Division of Entomology
Indian Agricultural Research Institute
New Delhi
leAR
INDIAN COUNCIL OF AGRICULTURAL RESEARCH

NEW DELHl
FIRST PRINTED MAY 1983
Chief Editor P. L. JAISWAL

Editor DR (MRS) A. M. WADHWANI
Assistant Editor RAJINDER SINGH
Chief Production Officer KRISHAN KUMAR

Associate D. S. BEDEKAR
Chief Artist M. K. BARDHAN

Senior Artist A. CHAKRAVARTY
© 1983, by the Indian Council of Agricultural Research, New Delhi
Price: Rs 33.00
Printed in India by Singhai Printing Press, JabaJpur -482002 and published by

Shri P. C. Bedi, Under Secretary for the Indian Council of Agricultural Research,
New Delhi.
FOREWORD
THERE is now a growing feeling that Indian agriculture is again entering
a state of relative slow growth rate. This is attributed partly to the
vulnerability of agriculture to weather conditions and partly to inadequate
progress in improving production and productivity during the south-west
monsoon period when over 80% ot rainfall in India is received. The
south-west monsoon season (May to Octob~r) generalJy referred to a.s
the kharif season is characterised by higher atmospheric huinidity and
cloudiness as compared to the winter or rabi season in most parts df the
country. The major food~rain crops of .the kharif season such as nce, sor-
ghum (iowar), 'maize and millets like the pearlmiIIet (bajra) as well as
industrial crops like cotton are affected by a wide range of pests. The
farmer has to wage It continuous battle against the triple alliance of pests,
pathogens and weeds more during the kharif, months. TIlls is why the
late Dr Pradhan used to repeatedly emphasise during his· life time that
just as human beings require an umbrella to protect themselves against
rain, crops require a plant protection umbrella during the humid season.
We now know that unless greater attention is paid to the health of tbe
plant and the care of the soil it will not be possible to maintain the
momentum of agricultural progress. This book is hence timely.
Dr Pradhan was a scientist of great vision and understanding. He
looked at crops as an integral part of a living system interacting with
the environment on the one hand and beneficial as well as harmful insects
on the other. He hence studied in great detail both host-insect and insect-
environment relationships. He did pioneering work on pesticides of plant
origin and demonstrated the anti-feedant and locust-repellent properties
of extracts from the seeds cif the neem tree.
The present publication brings together many of the papers and ideas
of Dr Pradhan. The Indian Council of Agricultural Research deserves
our gratitude for publishing this book, which will be of immense value
to both scholars and extension workers engaged in the study and control
of pests of agricultural importance. Our gratitude also goes. to Mrs Pradhan
for the deep interest she has taken to ensure that the vast store of valuable
information and ideas left behind by her late husband becomes available
to every one who wishes to derive benefit from it.
M. S. SWAMINATHAN
Director-General
International Rice Research Institute
and Independent Chairman,
April 11. 1983 FAO Council
ABOUT THE AUmOR
DR S. PRADHAN
(13 May 1913 - 5 February 1973)
DR Shyam Sunder Lal Pradhan, the author of this book was a great
teacher. scientist and a tireless crusader for due recognition to entomo-
logy in Indian agriculture. He expired on 5 February 1973 after a brief
illness.
Dr Pradhan was born on 13 May 1913 at Bahraich in Uttar Pradesh.
After being awarded the degree of M.Sc. in Zoology by the Lucknow
University in 1934, he continued to work there on a problem of functional
morphology in insects under the well-known zoologist, Dr K. N. Bahl,
and was awarded the D.Sc. degree in 1938; his thesis was adjudged the
best in the University for that year. Dr Pradhan then worked in an
Indian Council of Agricultural Research (ICAR) scheme on sugarcane
pests at Gorakbpur. In 1940 he joined the IARI as Assistant Entomologist
at its Kamal substation. In 1946 he was selected by the Government of
India to work at Rothamstead Experimental Station, England, to work
on a problem in insect toxicology, and was awarded the Ph.D. degree in
1948. On his return home he established the first school of toxicology in
the country. In 1958, Dr Pradhan became the first Professor of Ento-
mology in the newly created post-graduate school of the IARI, where he
organized the curricula for M.Sc. and Ph.D. degrees in entomology. In
1962 he was appointed Head of the Division of Entomology, and he con-
tinued to hold that office tiII his death.
Dr Pradhan established a section of ecology and toxicology at the
IARI where, besides guiding the research staff, he trained 65 students
in the subject, 45 for the Associateship diploma, one for M.Sc. degree,
and 19 for Ph.D. Today most of them are occupying responsible positions
in India and 'Other countries, a~d some have established toxicology labo-
ratories in their states.
Among the more important of his scientific contributions are Pradhan's
temperature development equation; biometer; biotic circuit; biotic theory
of locust cycle; effect of temperature on insects' susceptibility to insecti-
cides; effect of particle size on the toxicity of insecticides; differentiation
between external and internal resistance of insects to fumigants; corre-
lation between insecticide solubility in cuticular lipoids and insect
resistance to insecticides; correlation between epicuticular structure of
insects and their resistance to insecticides; mode of action of DDT; neem
as an· insect 'repellent; techniques for population studies; and ecological
ABOUT THE AUTHOR v
manipulation of safe storage of foodgrains (Pusa bin). Dr Pradhan
published about 200 papers in various scientific journals and a book
'Insect Pests of Crops'. Keenly feeling the ne~d for communication among
entomological workers in the to un try, he initiated the publication of the
monthly 'Entomologist Newsletter'. The present book was the last one
written by him.
Dr Pradhan was a founder-member of the Entomological Society of
India and heild one or the other office since 1942. He was the president of
the society for 4 terms; and It was during one of these that the society
celebrated its Silver Jubilee in 1964, when a national seminar on various
aspects of entomology was also organized. In 1969, again during his
presidentship and because of his presistent efforts, the first international
seminar on integrated pest control was organized by the society. In reco-
gnition of his contributions to science, he was elected a Fellow of the
Indian National Science Academy in 1966.
Dr Pradhan was widely recognized authority on insect ecology, toxi-
cology. and integrated pest control. In 1956 he was invited by UNESCO
to write a chapter on the 'Ecology of arid zone insects' in the book 'Human
and Animal Ecology'. The honorarium received for this he donated for
instituting an annual award of a gold medal to the most outstanding
student in entomology at the IAR!.
He was a member of the F AO panel of experts on integrated pest
control. For a number of years, as the Chairman of the Entomological
Committee of the lCAR, he guided the planning of national policies on
entomological research. teaching and extension. About a month before his
death he actively participated in preparing plans 'for launching an intensive
drive on integrated control of pests of rice.
Dr Pradhan had travelled extensively to several countries as a visiting
scientist and for participating in international meetings and seminars.
His critical observations on the status of entomology in different coun-
tries are recorded in his article 'Entomology round the globe'. In the
14th International Congress of Entomology. held at Canberra, Australia,
in August 1972, he presided over three important sessions. Immediately
after his return from the congress he was invited to Hawaii to
advise on preparing a syllabus for the special course on integrated pest
control.
During the last few months of his life he worked to cdIlect and
project facts and figures to prove that pest control research has to be
further intensified for increasing yields of different crops. His last publi-
shed paper, 'In tropics protection research more needed than production
research', projects most of his ideas. Another paper completed just before
his death was 'Plant breeding through the window of pest control'. which
vi AGRICULTURAL ENTOMOLOGY AND PEST CONTRoL
he was to have read at a conference of plant breeders in the last week

of February 1973.
Dr Pradhan was due to retire as Head of the Division of Entomology
on 12 May 1973. but to continue utilizing the services of this distinguished
entomologist even after retirement the leAR had announced his appoint.
ment as an Emeritus Scientist more than a year in advance.
CONTENTS
Foreword iii
About the Author iv

1. Place of Entomology in Relation to Man 1
2. Dominance of Insects 8
3. Insects Useful' to Man 18
4. Origin of Insect Pests 40

5. Estimation of Insect Population 52
,
6. Assessment of Losses due to Insect Pests 75
lDS
8. Effect of Temperature on the Dynamics of

Insect Development 123
9. Principles of Insect Control 177

10. Pesticide Hazards 198
11. Revolution of Pest Control 212
12. Importance of Protection Research in Tropics 223
13. Integrated Pest Control 239
Epilogue 260
Index 261
CHAPTER
PLACE OF ENTOMOLOGY IN RELATION TO MAN
THE word insect has its root in the Latin word insecare (to cut into) and
is equated with 'insectum'. Entomology. the science of insects, has its
origin in the Greek word temno (to cut). The Sanskrit word keet may
have a similar sense and root. Thus. the most prominent physical cha-
racteristic of insects which seems to have impressed the early observers
who coined these names, is the division of the insect body into 3 promi-
nent regions, namely, head, thorax, and abdomen; this single feature has
had the most spectacular impact on the etymology of entomology. The
second important character of an insect is its 3 pairs of legs; this is the
basis of the term Hexapoda, meaning 6-legged, coined by Latreile in 1825
for the zoological class Insecta (Fig. 1.1). The corresponding Sanskrit'
Fig. 1.1 A typical representative of class Insecta.

2 AGRICULTURAL ENTOMOLOGY AND PEST CONTROl
term Schatpada, also meaning 6-legged, was there in Amarkosh as eady

as the first century A.D. when that dictionary was compiled. The third
character, which almost completes the definition of this class of animals,
is the chitinous body wall, which also serves as skeleton and is called
exoskeleton or outer skeleton; actually it is a characteristic of the larger
group, the phylum Arthropoda, to which insects also belong. Thus, an
insect inay be defined as an animal which at some stage of its develop-
ment possesses the combination of these 3 major characters, (i) chitInous
exoskeleton, (ii) 6 legs, and (iii) body prominently- divided into head,
thorax and abdomen. In other words, insects are arthropods having a
combination of the last 2 characters.
In zoological classification insects constitute just one of 5 major
classes of the phylum Arthropoda, which in turn is just 1 of 15 major
and several minor phyla of the animal kingdom. From the economic point
. of view there are several groups which are important enough to have led
to the development of different branches of study, such as Ornithology
(birds), Ichthyology (fishes), Helminthology (worms), as well as economic
departments such as those of poultry, fisheries, dairying. There is, how-
ever, hardly any other class of animals or plants which has such vast
potentialities, both for good and for bad. and such remarkable degree of
even purely academic interest, as the class Insecta. The importance of
insects consists in (i) the damage they cause to field crops. orchards,
forests. stored products and household materials; (ii) their interference
with man's health and comfort; (iii) their value as (a) pollinators and
agents of fruit setting in orchards and crops, (b) suppliers of products like
silk. honey and lac. (c) scavengers, and (d) useful parasites and predators;
(iv) their remarkable suitability and utility as material for purely scien-
tific studies. e.g., (a) genetics and inheritance, of which Drosophila is the
hero, (b) sociology and popUlation growth. for which ants, termites. wasps.
and bees have .been in the forefront, and (c) adaptability, mimicry, etc.,
for which the leaf butterfly Kallima and the stick insect are the classical
examples. All these and many others represent the diverse ways in which
insects are intimately associated with man.
Fundamental versus applied Entomology. In practice the term ento-
mology has an applied significance. Hence, it is necessary to understand
at this stage the implications of applied science. The pairs of opposites
generally used are 'fundamental versus applied aspects', 'fundamental
versus developmental aspects', 'academic versus applied approaches'. For
a clea~ understanding, it should be divided into academic and applied
aspects. The motto of the academic approach should be, as it has always
'been, 'knowledge for the sake of knowledge'. Any aspect which is not
known' ought to be known, irrespective of and undeterred by any idea
PLACE OF ENTOMOLOGY IN RELATION to MAN
about the utility, futility, or even harmfulness of the knowledge thus

gained. Any lacuna in knowledge must be filled. This has been rightly
the approach of the traditional universities. The approach of the applied
institutions, such as the agricultural research institutes or the recently
established agricultural universities, has to be rather different; they have
to engage themselves in problem-oriented research activities. They have
to be sure that if they are successful in obtaining an information as they
expect, it will be definitely useful as planned. First they have' to identify
the problems carefully, and then to analyse them in detail so as to expose
clearly the lacunae which must be filled for proper solution ot'the pro-
blems. This may call for only some adaptive research for successful ex-
ploitation of an already known principJ~ under a specific situation, or it
may call for a fundamental probe. Hence, the applied aspect should be
further divided into fundamental and developmental; the former may be
designated as 'fundamentals of applied aspects'. Alternatively, the science
of entomology can first be divided into 'fundamental and: developmental
aspects', and then the fundamental aspect can be subdivided into 'academic
and applied approaches', the former resulting from the motto 'knowledge
for the sake of knowledge' and the latter meant to fill up a fundamental
lacuna for the solution of a well-identified applied problem.
For organizational facilities and for quick scientific solutions of pro-
blems the former dichotomy is more useful, for it provides better co-ordi-
nation between developmental and fundamental aspects necessary for the
solution of any problem.
Science of Entomology
!
I I
Academic aspects Applied aspects
(Motto: Knowledge for the (Motto: Problem-oriented
sake of knowledge) scientific pursuit)
I
Fundamental aspects Developmental aspects
(Aim: To fill up a fundamental (Aim: Adaptive research for
lacuna in knowledge for proper exploitation of a
the solution of a well- known principle or
identified problem) phenomenon)
OR
4 AGRICULTURAL ENTOMOLOGY AND PEST CONTROL
Science of Entomology
I
I
Fundamental aspects Developmental aspects
(Aim: To fill a fundamental (Aim: Adaptive research for
lacuna in knowledge) proper exploitation of a
known principle or
! phenomenon)
I
Applied aspects
Academic aspects
(Motto: Knowledge for the (Motto: To fill a fundamental
sake of knowledge) lacuna for the solution
of a well-identified
problem)
Entomology as an Applied Science

In applied biology, when the aim is economic exploitation of any
species, it has to be studied in almost equal depth whether we want to
exploit it in a positive or a negative way; i.e. to culture it for increasing
its produce as in 'the case of crop plants or to check its multiplication as
in the case of pest species. The number of species of insects is So large
(more than 1 million) that the preliminary item of their identification
itself is one of the most tedius jobs, for which no single organization
in the world-is fully equipped. This is an indication of the work load in-
volved in the solution of pest problems.
The entomologist's interest in various forms of life has to be as varied
as the insects he has to deal with, as will be evident from the following
examples.
1. Class Mammalia. The dairy, wool and livestock industries are
based on this class. Monkeys, jackals, bats, squirrels and rats are
serious pests of agriculture. Insects are so much responsible for dis-
eases of man and livestock that there are separate branches of medical
entomoJogy and veterinary entomology.
2. Class Aves. The poultry industry is based on this class. Fruit-
eating birds such as parrots are serious pests, but some other birds eating in-
sects are useful. Information on their diet and economic importance is
very useful. Entomologists are also interested in insect parasites of pOUltry.
3: Class Reptilia. A number of reptiles including snakes are very
useful in keeping the insect pests under check.
4. Class Amphibia. Some members of this class, e.g. frogs, are use-
ful; they eat insects.
5. Class Pisces. Some fishes eat insects; Haplochilus lineatus
PLACE OF ENTOMOLOGY IN RELATION TO MAN 5
destroys mosquito larvae and has been recognized as definitely useful in

the control of malaria. Ophichthy boro, which is a long. flat eel living in
tidal streams. has been reported to do considerable damage by burrowing
into and making wide passages across the bunds. thus connecting paddy
fields containing fresh water with salt-water channels.
6. Class Chilopoda. Centipedes feed on insects and are beneficial
although sometimes they inflict very painful bites on man.
7. Class Diplopoda. Some species of millipedes attack growing
crops and become serious pests in fields and greenhouses altpough many
feed on decaying vegetable matter. Those that are pests feed on roots or
leaves near the ground.
8. Class Arachinda, The spiders (Order Araneida) feed on several
injurious insects. and have venomous jaws which poison insects. Sometimes
their bites cause serious results. On the whole they are beneficial.
Mites and ticks belong to the order Acarina. Some species of mites
injure plants, some are parasitic on insects (locusts and grasshoppers).
Some are predatory on some insects and mites. While certain species of
ticks are parasitic on men. livestock and other animals., some transmit
diseases.
Scorpions (Order Scorpionida) bear sting at the tip of the abdomen.
They are the best known stingers. Their pedipalps are much more developed
and carry a pair of pincers which are used to grasp the prey. They are
predatory in habit. Despite the sting. they may be considered beneficial.
9. Class Crustacea. Crabs. prawns, lobsters and sow bugs are some-
times injurious in greenhouses. Crabs are known to damage paddy crops.
although several species are carnivorous.
10. Phylum Mollusca. Snails and slugs are often serious pests of
plants in greenhouses and in fields.
11. Class Nematoda: Roundworms are becoming so important as
plant parasites that separate departments of Nematology are coming into
existence.
Colossal Damage by Insects

The one facet of insects' life economy about which we are seriously
concerned is their potentiality for doing immense harm to man's interests.
There is hardly anything that man would like to Ci!lI his own and insects
will not challenge his claim. Taking for example the insect damage to
economic plants and plant products. a rather conservative and commonly
accepted estimate has been that the average loss caused by insect pests
is about 10%. To the nation this means an annual loss of Rs 500 crore-
10% of Rs. 5000 crore, which used to be our average annual income
from agricultural and forest resources. The great harm done to human
and livestock health is incalculable, as also it~ repercussion on the

national activity.
In spite of their causing colossal losses in v:lrious fields of human
activity, the minute size of many insects misleads us to a false sense of
complacency towards insects in general. If a lion ellters a town, the whole
town will get thoroughly shaken up, and every effort will be made to
kill it immediately, although its potentiality of damage may be only a
few death~. But if a house fly carrying cholera germs enters a restaurant,
it can flit about from dish to dish and many people may not even move
their finger to scare< it away, even though its potentiality for harm may be
many deaths through an epidemic of cholera. The dangers from the insect
world arc often underestimated, and it is seldom realized that in spite of
the usefulness of some groups the insects as a class constitute enemy
number 1 of the human race.
It is interesting to note that there are ample references even in
the most ancient records to the 3 main groups of useful insects which
have been providing honey, silk, and lac; and mlln bas preserved tbem
for these valuable commodities. In comparison with such sound study
of useful insects the knowledge about harmful insects at that time was
surprisingly poor. It is amply clear from various hymns in the Vedas
and other ancient literature that, although the pest problems were quite
serious, the knowledge about them and their control was meagre. In
fact, realization of the dangers from the insect world has been extremely
slow all over the wor~d, and the science of entomology came into being
more as a result of the hobby of amateur insect collectors than from the
labours of professional economic scientists. In India this realization has
been even slower, P;;)bably because of the religious beliefs and abhorrence
of destroying life and certainly because of little interest in insects even
as a hobby. In this connection it is instructive to recall the Mahabharata
story of sage Mandavya who developed the hgbit of collecting and
pinning insects; for this crime he had to pay the penalty of his life by
being himself impaled on the point of a crowbllr. Our public is still
unsympathetic to activities of entomologists and expects only simple,
harmless and cheap methods of keeping away frorn their crops and crop
produce what they would like to call mere insect nuisance. Thus, the·re
exists a puzzling, yet interesting, anomaly.
I wonder if any attempt has ever been made to appreciate and
expla1n this anomaly. Careful scrutiny of the wbole situation suggests
2 explanations. The first is somewhat simple. 1he pest problems of
agriculture, for example, arose with the origin of agriculture, and became
aggravated with the intensification of agriculture. Therefore, one can
say that pest problems could not have been so serious when agriculture
PLACE OF ENTOMOLOGY IN RELATION To MAN 7
itself was in its infancy. Similarly. epidemiological studies on insects of

medical and veterinary importance have shown that the incidence of
insect-borne diseases increases with increase in population density of
human beings and livestock. Thus. one can also say that the harm done
even by such insects might not have been so serious in the early days
of human civilization. These arguments only partly explain the' anomaly
but provide full and convincing answer to the query why insect problems
are multiplying with increase in the number of entomologists. The other
subtler explanation is that insects have been exploiting their characte-
ristic minuteness for dodging humanity and their other enemies and
putting them in a sort of complacency or false sense of security against
them. In this way the insects have been ensuring the success, of their
struggle for ex;istence. Generally. however. insects are neither so minute
as to be invisible to thCl naked eye and thus to create a sort of blind awe,
nor large enough to create a visual impression about the magnitude of.
their seriousness. except under certain specia,l circumstances. e.g. locust
invasion.
CHAPTER 2
DOMINANCE OF INSECTS
INSECTS came into existence, according to various estimates, 250-500

million years ago. The antiquity of this can be gauged, from the fact that
man came on the scene only 1 million years ago. Ever since such, an
early geological epoch insects have held their own against all odds and,
as a result of their struggle for existence for so long, today they constitute
the most dominant class of the animal kingdom. They also represent the '
culmination of evolutionary development in terrestrial arthropods.
Size of the class Insecta. Insects constitute the largest class noti only of
the animal kingdom but also of the whole living world. The number of
known species of insects is much more than that of all other species of
the animal kingdom put together. Estimates of both the absolute number
of insect species and their relative proportion vary from 700,000 to 1,500,000
species, and from 70% to 90% of all known species of the animal kingdom.
According to Metcalfe (1940),* the knowledge about animals has been
growing during the last 200 years. In 1758, when Linnaeus established
the binomial nomenclature, he described only 312 genera and 4,203 species
of animals of all classes. Of these, 74 genera were of insects, with 2,102
species. In 1902, Sherborn listed all animals described between 1758 and
1800, and his list included 3,234 genera and 58,833 species. Since 1850 the
annual publication 'Zoological Records' is quite a reliable source of in-
formation. On counting the number of new species published in these
records every 10 years it was found that on an average 10,542 new species
of insects and 7,253 of other animals were described annually. If these
averages are correct, they give a total of 948,780 species of insects des-
cribed between 1850 and 1940, and 652,770 species of animals other than
insects. This appears to be the most reliable of the available estimates,
although Metcalfe tried 2 other methods, and found the average of the
3 estimates to be 1,500,000 species of insects.
Size of Individual Insects. Insects as a class are very small individuals.
In fact, they include the smallest forms of animals with free and active
aerial life. No other animal group has been able to evolve forms so small
in siz~ as some of the insects and yet be capable of leading a free aerial
existence. There are some microscopic spores, of both animal and plant
origin, but they are non-active resting stages which are blown about by
the wind. All the same, the size of insects varies widely. An appealing
• Metcalfe, Z. P. 1940. Ent. News 51 : 219-222.

DOMINANCE OF INSECTS 9
statement about the size variation in insects is that some are smaller
than the largest Protozoa, the phylum which contains the smallest, singl.e-
celled animals; and some are larger than the smallest Vertebrata, the
phylum which contains the largest animals. The following are some of
the insects which represent these extremes.
1. The largest insects

Coleoptera
Megasoma elephas - 120 mm
Macrodontia cervicornis - 150 mm
Orthoptera ,
Pharnacia 'serratipes - up to 260 mm (length)
Hemiptera ,
Be/ostoma grande - up to 115 mm (length)
Lepidoptera
Erebius agripinna - 280 mm (wing span)
Attacus atlas - 240 mm (wing span)
Odonata
Moganeura (fossil from Upper Carboniferous Strata) -_ more than
600 mm
2. The smallest insects
Coleoptera
Trichopterygidae - 0.25 mm
Hymenoptera
Mymaridae - smaller than Trichopterygidae
Population of Individual Species

Nobody has attempted to assess the population of the various insect
species bi the world. It is, however, interesting to note that whereas
the population of human beings is 3 to 4 thousand million in the world
it is negligible in comparison with the population of any species of insect
pest. For example, the population of aphids in a mustard crop was esti-
mated as more than 40 million of individuals per acre.
Thus, of the 3 criteria, viz. the size, the variety, and the population
density, the insect class can unquestionably claim dominance on the basis
of the last 2. In size of individuals, the insects may seem to be at a dis-
advantage, but small size is highly advantageous for survival in the struggle
for existence.
Secrets of Insects' Inherent Strength and Dominance
Apart from their small size, insects as a class have acquired many
structural, developmental and behaviouristic perfections not found in any
other group of animals or in plants; the significance of these has yet to

be fully appreciated even by entomologists. The following are the cha-
racteristic features that contribute to the insects' dominance as a class.
Structural Perfections
Exoskeleton. This is a characteristic of all arthropods, but insects
seem to have fully exploited the mechanical advantage of their exoskeleton;
for it not onfy provideS! a much larger area for the attachment of muscles
but also protects the muscles from mechanical injury. Moreover. it affords
an excellent mechanism to protect insects against desiccation. But for the
very effective waterproofing efficiency of the exoskeleton, probably such
minute animals as most insects are could not have evolved so successfully
even as terrestrial forms, much less as aerial ones. No other form of life
with such minute size has been able to lead active aerial existence. The
main disadvantage of small size is the comparatively large surface
area per unit body weight; this means greater vulnerability not only to
mechanical injury but also to desiccation. Insects have successfully over- _
come both these difficulties by virtue of their exoskeleton. In addition.
the exoskeleton has turned the appendages into good tools for digging,
preying, and oviposition. Lastly, the exoskeleton maintains the shape of
the body much more efficiently than the endoskeleton; old ~age is unable
to affect the form of adult insects, and even death does not necessarily
deface the beautiful shape of the live insects. How far this advantage
has contributed to insects' success in the struggle for existence is difficult
to assess, but it has certainly made the job of insect collectors easy. Some
important corollaries mentioned below have followed the development of
the waterproof exoskeleton.
(i) Small size : The evolutionary course followed by insects has led to
the development of a large number of smaller individuals rather than a
small number of larger individuals. Undoubtedly, this immensely increases
the chances of survival of the species; for the larger the number,. the
greater the survival chances; and the smaller the size, the easier for indio
viduals to subsist on small quantities of food and to take refuge in small
niches against the vicissitudes of weather and against adverse biotic
agencies. With larger numbers resulting from smaller size the chances of
variability and mutation also increase, thus making it possible for nature
to conduct a larger number of trials in evolution.
Lflstly the smaller size leads to greater efficiency. It is one of the
biokinetic principles that the contractile power of muscles depends on
their cross-section, i.e. area as square of a linear dimension, or a unit with
2 linear dimensions; the weight on the other hand depends on the
cube, i.e. on a linear dimension raised to the power 3, or a unit
DOMINAN_CE OF INSECTS 11
with 3 linear dimensions. Hence, when the size of an animal in-

creases, the power of its muscles increases at a lower rate than its
weight. So, muscular strength is better in smaller animals. It is no
wonder that a flea with a length of _1.3 em can jump a horizontal
length of 33 em and a height of 20 cm. For a man to do a similar
feat it would mean a long jump of 213 metres and a high' jump
of 137 metres. This gives an idea of the relative efficiency of an insect's
small size.
(ii) Quicker speciation: As a result of the exoskeleto~ and small
size there has been quicker speciation. It has been conclu&d that the
peculiar combination of the 2 main characters - the chitinous exoskele-
ton and the at'?rial mode of life - has probably been the l~lfgest contri-
butor to the extraordinarily high degree of speciation in insects. Owing
to the mechanical rigidity of the chitinous exoskeleton, comparatively
minute variations i~ the exte~al genitalia have led to reproductive isola-
tion and consequent fixation of the species. This factor alone, however,
cannot explain the speciation in insects because. other classes of Arthro-
poda having similar exoskeleton do not have as many species. Develop-
ment of the aerial mode of existence, especially the acquisition of wings
leading to life under diverse ecological habitats, has provided the trigger
for variations . It is believed that the chitinous exoskeleton made possible
the evolution of minute insects with aerial mode of life; that life in
diverse habitats resulting from the development of wings triggered a vast
amount of variation; that the higher population of individuals resulting
from the minuteness of the size increased the chances of variability and
mutation; and that quite a large number of these variations and muta-
tions, especially those associated with even slight alterations in the external
genitalia, became fixed and gave rise to a large number of different spe-
cies because of reproductive isolation resulting from the rigidity of the
chitinous exoskeleton of the genital armature. This vast increase in the
number of species immensely increased the economic potentialities of
insects as a class.
These considerations lead one to speculate on the wider aspects of
species and speciation in general. It is generally agreed that the most
definitive criterion for distinguishing species is that they do not interbreed.
Owing to practical difficulties in applying, this criterion the taxonomists,
out of operational necessity. separate species on the basis of other easily
observable characters; but the implication is always there that the parti-
cular characters are reliable guides for assuming that individuals having
those contrasting characters will not interbreed. But at times even this
last criterion fails; for it has been possible to obtain fertile hybrids not
only between different species but also between different genera. though
more SO in the plant kingdom than in the animal kingdom. Such success-
ful hybridization indicates that reproductive isolation is not always attri-
butable to failure of the male gamete of one to fertilize the female gamete
of the other. In other words, reproductive isolation leading to speciation
is not always due to what may be called in a broad sense protoplasmic
incompatibility between 2 species. The possible mechanisms of reproduc-
tive isolation leading to speciation may be as follows.
(a) Geographical: isolation: A variant from the parent popul;ttion
happens to be taken away to: such a distance or across such a barrier that
there is no possibility of intermixture of the characters, although if the
2 were brought together they could interbreed and the variant could be
absorbed back into the parent population.
(b) Simple physiological isolation: Two species do not interbreed
in nature because they come to maturity at different times. This happens
in the case of plants even because of such apparently slight differences as
the anthers dehiscing at different times o~ the day.
(c) Morphological isolation: This appears to be the major cause
of speciation in insects, since comparatively minor variations in the
genital armature may preclude successful copulation.
(d) Protoplasmic incompatibility: This can be said to exist when
all other simple and apparent causes of reproductive isolation are absent
and even then interbreeding cannot be achieved.
There may be several other factors responsible for reproductive
isolation, e.g. mutual repellence due to various causes. However, diffe-
rent species need not be reproductively isolated because of protoplasmic
incompatibility, and it should be easy to overcome other causes of repro-
ductive isolation. If this line of thought is sound, it should open up immense
economic potentialities through hybridization, especially by artificial
insemination and cross-fertilization. For taxonomists too the implica-
tions should be quite significant; for the boundaries between different
species become still more shaky and the life current flowing through
different species becomes more continuous.
Functional wings. This feature the insects do not share with other
arthropods, and it gives them a definite superiority. The power of flight
greatly increased statistical chances of survival and dispersal, except on
wind-swept islands where the reverse adaptation of losing the functional
wings took place. Functional wings increased the feeding and breeding
range: and provided a new means of eluding enemies. attacking a fast-
moving host (running or flying), and finding a mate. Increased feeding
range undoubtedly opened the way for the adoption of specific food,
especia:Ily where the host or breeding medium was sparse. For example,
it would allow a species to adopt carrion as food since the individual
DOMINANCE OF INSECTS 13
with functional wings could seek out and reach carcasses which are not
only isolated but also remain suitable as food for only a short period.
This applies also to mosquitoes, which, but for fUnctional wings, would
not have been able to breed in small pools of water that dry up in a
short time. Functional wings also enabled various species to undertake
intercontinental migrations as in the case of locusts and butterflies. Some
species have perfected this locomotion by developing air sacs, as in bees
tfor carrying honey) and flies (for compensating the loss of one pair of
wings). The structural perfection of these wings can be gauged from the
fact that a hive bee can fly at the rate of 9 km/hr, a hoverfly' 12 km.
a hawk moth F km, ana a butterfly up to 90 km. This is effected by
means of wing' beats in a regular succession, the frequency of which has
been estimated to be 250/second in honey bee, 190 in hoverfly, 85 in
hawk moth, and 12 in large butterflies.
Hexapod locomotion. Jointed nature of the legs is a, characteristic
which insects share with other arthropods, but in insects the number of
legs has reached an ideal stage of evolution. It is instructive to appreciate
that the 6 legs of insects is the optimum number, which no other grolIp
of animals has. The larger number of legs as in millipedes is bound to
make locomotion rather cumbersome, but when the number of walking
legs becomes less than 6 it may create a problem of balancing during
lncomotion. For example, when walking, a biped has to balance his body
on one leg while the other moves forward, and a quadruped has to
balance on 2 legs while the other 2 move. During insect locomotion the
body always rests on 3 legs (a tripod) while the other 3 move forward.
Thus, 6 is the smallest number for a stable equilibrium during all phases
of terrestrial locomotion.
Compound eyes. Insects usually have a pair of compound eyes in
nymphal and adult stages, and sometimes simple eyes too. Insects share
this characteristic only with crustaceans. The compound eyes are of very
complicated structure, not met with in other groups of animals. They are
called compound eyes because each, instead of having a single large
cornea, consists of a number of hexagonal areas each representing the
cornea of a discrete visual organ called the ommatidium, all of which are
compacted together. Thus, ants have 50 to 400 facets or corneas in each
eye, house-fly has 4,000, a swallow-tail butterfly 17,000, and certain
sphinx moths and dragonflies more than 50,000. It is obvious that an
insect will not lose the power of vision completely if a few ommatidia·
are injured.
Scattered sense organs. Except the eyes, none of the sense organs
are invariably concentrated on the head. The organs of hearing are
sometimes associated w~th the antennae. The organs of taste and smell
may be on the antennae, on mouthparts, or even on tarsi or cerci. Such

diffused and scattered nature of sense organs is bound to be advantageous
to the insects; for this arrangement reduces the chances of all being
injured at the same time.
Decentralized nervous system. The central nervous system is a
ladder-like ch~in of ganglia along with ventral side of the body. except
one compl~x nerve centre in the head, i.e., the brain. This system.
despit~ the highly evolved stage of insects as a class. is so decentralized
that brainless insects cun be artificially stimulated to walk, fly or even
feed, and according to some, the abdomen of the living female silkworm
moth when separated from the thorax can be fertilized by the male and
stimulated to lay fertile eggs. How far this decentralization of functions
of the nervous system is useful in preservation of the species in nature
is obvious.
Direct respiration. This important physiological function is inde-
pendent of the mediation of blood, which is not required to transmit the
gases of respiraion. These are taken to and from every minute part of
the system directly through air tubes called tracheae and tracheoles.
which have several openings called spiracles on the body surface. This
mechanism fits in very well with the requirements of terrestrial and
aerial existence of minute creatures liable to desiccation, which can alter
the consistency of liquid blood.
Enteronephric excretion. The excretory function is performed by
malpighian tubules, which open into the front part of the intestine
instead of opening directly to the exterior. The excretory products are
semi-solid rather than liquid urine. This enteronephric system of excre-
tion, regarded at the time of its discovery in earthworms as an unex-
pected phenomenon. was subsequently found to be universal in insects.
Obviously this system is also very well suited for water conservation.
since it makes the excretory function independent of water supply. and
water is not wasted. There are further modifications in the same direction ..
Thus, despite the minute size of the entire internal organization.
thl! insect system is highly complex, quite different from that of other
groups of animals. and admirably adapted to successful life under a
variety of conditions and especially to arid existence.
Developmental Characteristics
• The development of certain groups of insects is unique in the animal
kingdom. The full life-history of the insect consists of 4 well-defined
stages: the egg. the larva. the pupa, and the adult. -Two of these stages,
the mainly feeding larval stage and the mainly reproductive adult stage.
are so different from each other not only in structural details but also in
DOMINANCE OF INSECTS 1S
requirements of food and habitat that competition between the parent

and the offspring for food or shelter is completely eliminated. Further,
by the -interpolation between the larval and adult stages of a quiescent
stage of pupa with great resistance and practically no food require-
ment, the 2 active stages are best fitted into different sessions optimum
for them. The same purpose is served by the eggs in certain cases, e.g.
the phadka egg interposed between the adult and nymph. Thus, the
development of metamorphosis has enabled insects to adapt themselves
to varied situations of existence much better than animals nQt possessing
this mechanism. I
High Fecundity and Controlled ReproductioD

The large' number of eggs laid by the female and the quick rate of
development make insects very prolific. It has been calculated that the
progeny of a single moth can cover the entire dry surfac.e of the earth
to a depth of 25 metres wjthin 1 year if all the 'progeny lives the normal life
span. This calculation is on t~e assumption that a moth lays 200 eggs
and completes the life-cycle in 1 month. The position is about the same
for many other insect species, though there are species which lay thousands
of eggs. Some species have increased their prolificity by polyembryony,
i.c. production of several embryos from a single egg, as in some chalcids;
others reproduce even in the absence of males by parthenogenesis, i.e.
development of the egg without fertilization by male sperm. At the other
ex.treme, there is completely controlled reproduction in some of the
social Hymenoptera, which have learnt to make their surroundings so
safe as not to require wasteful reproduction. A number of social species
of wasps, bees and termites have developed such a high degree of
control on reproduction that all except one or a few females of the
colony are rendered unfit for reproduction and constitute a class of sterile
females. This strict division of labour between reproducing females known
as queens and the labouring ones called workers represents the highest
evolution of a special order in the entire animal kingdom.
Specificity of Feeding
While there is a vast diversity of food habits among insects, there
being often some specific preference 'for one kind of food or the other,
there are insects which have overcome interspecific competition by becom-
ing polyphagous by living on almost any kind of vegetable matter, e.g.,
l~cust and armyworms or by taking to such food for which there is no
competition, e.g., cellulose-eating termites. There is sometimes extreme
degree of specificity not only for different plant species or varieties but
also for different parts of the same plant, as in the case of root-borers, stem-
borers, and top-borers of sugarcane.
Protective Adaptation and Device

Protection against adverse biotic and climatic agencies is as much
essential to normal existence of individuals as to continuance of the spe-
cies. Insects have developed a variety of ingenious ways for ens4ring this
protection.
Morphological adaptations. The protective value of the exoskeleton
in many insects has been supplemented by bristles, spines, hairs, or scales.
These in some cases afford more effective protection by their capacity to
eject venomous liquid, as in many caterpillars and blister beetles. The
minute size of many insects affords protection against large predators. Some
insects are so large and of such grotesque appearance as to frighten away
the predators. Insects as a class have developed a high degree of protective
colouration and adaptations for camouflaging. These have taken various
forms, including protective resemblance with the surroundings in which
insects live (e.g., stick insects, leaf butterfly, bark beetles), and protective
mimicry in which one species mimics some dangerous or distasteful species.
Many butterflies and other insects, themselves edible to birds, frogs, etc.,
resemble in colour, shape or form other species which are poisonous or dis-
tasteful or simulate other insects such as wasps which are known to have
venomous stings. There is another type of camouflage, known as aggressive
mimicry, in which the mimic is able to deceive its prey. Then there is also
what iB known as warning colouration in which venomous or distasteful
insects are so prominently marked and coloured as to be easily distin-
guished from other species.
Behaviouristic adaptations. Some insects, e.g., stinging Hymenoptera,
take up such an offensive attitude when threatened that the predator is
frightened. Some insects feign this attitude even though they have no real
weapon. Others adopt such dodging attitudes as feigning death. Still·
others, e.g., caterpillars and ladybird beetles, eject some kind of offensive
liquid when the enemy approaches them.
Construction of protective structures. The galleries of termites
(white ants) represent air-conditioned pathways wherever even the workers
have to go. There is no parallel in any other group of animals. The hives
of honey bees and wasps, the cocoons or pupae of so many species, the
bag!! of clothes moth and other bag moths, the underground nests of ants,
the nests of so many solitary wasps, are all of this kind.
Selection of safety niches. Many insects use their jumping or flying
capacity to escape danger. Others search favourable niches for tiding over
unfavourable conditions, and sometimes it means long-range migration
as in the case of many butterflies.
DOMINANCE OF INSECtS 17
Zenith of EvolutioD
The zenith of evolution depicted by such social' insects as termites,
ants and bees shows the degree of specialization which the insects have
attained. They have evolved their own agriculture, their own dairying,
th~ir own division of labour and social order, and even their, own
language. This all round efficiency is very effective in the struggle
for existence.
The foregoing paragraphs contain only brief references to ,structural,
developmental, bebaviouristic and organizational perfections of; the insect
world. They are meant to make the reader ponder over the problem and
realize that in the interspecific wars for supremacy man faces tough
challenge from ;,the insect' world. These minute creatures have held their
own for hundreds of millions of years dtlring which many gigantic forms
have come and gone.' Such being the case, it is no wonder that the control
of insect pests is so difficult.
CHApTER 3
lNSECfS USEFUL TO MAN
ALTHOUGH a large number of insects are injurious to crops and are to

be eliminated or effectively kept under check, there are a numb~r of
insects which are of use to man in a number of ways and need protection.
There are many crops, particularly those of fruits and vegetables,
that cannot be successfully grown and harvested without the active and
positive co-operation of insects. Some of these crops are apples, pears,
oranges, peaches, prunes, cherries, plumps, raspberries, blackberries, cran-
berries, grape-fruits, lemons. and figs among the fruit crops; tomatoes me-
lons, pumpkins, cucumbers, peas, beans, squash, egg-plant among the vege-
tables; and clover, alfalfa, soybean, cowpea, sweet clover and cotton among
other crops. These crops depend almost exclusively upon insects for the'
production of their fruits and seeds. This is so because of the phenomenon
known as pollination, insects acting as pollinators. Because pollination
goes on in nature in a rather inconspicuous manner without the farmer
coming into the picture, he is generally unaware of these immensely valu-
able acts of his mute partners. Such friendly insects have not received
adequate investigational attention even from scienti~ts because until
recently it was not sufficiently realized that man can by his own interven-
tion appreciably help this rather incidental but co-operative venture bet-
ween insects and man.
Insect Pollinators
The scientific principal behind insects acting as pollinators is as
follows. Although some plants can continue reproduction by asexual
means, as by bulbs, tubers and cuttings, many plants have essentially the
same type of sexual reproduction as in animals. The reproductive sexual
organs of the so-called flowering plants with which mostly the farmer-s
have to deal with are lodged in the flowers. The principal part of the
male organ is called the anther, which produces a large number of pollen
grains, i.e., the male sex cells, and the principal female- organ is the ovary
containing the female sex celis and the stigma which receives the pollen.
In many plants these male and female organs are lodged in the same
flower; in others the flowers with male organs (male flowers) are separate
from those having female organs (female flowers); in many species the
two kinds of flowers are borne on the same plant, whereas in others the
male and female flowers are borne on separate plants. In the last men-
tioned plants it is quite easy to comprehend that pollen from male flowers
INSECI'S USEFUL TO MAN 19
has to be carried by some agency to the stigma of female flowers. What is

further interesting. however. is that even in many pf those species which
have both male and female organs in the same flower there is some kind
of arrangement that sexual combination, i.e., fertilization, does not take
place between its own pollen and ovary. There are often quite elaborate
arrangements in favour of cross"fertilization or cross-pollination and
against self-fertilization or self-pollination. In other words, there is a kind
of code of sexual conduct even in plants.
Nature is a master geneticist t:}(ploiting the advantages of hybrid
vigour often qui~e effectively. Insects have entered into indispensable
partnership in this game with plants on the one hand and farmers on
I
the other. The pollen of one flower is carried to the stigma of the other.
mainly throug}]: 2 agencies. wind and insects. It is generally easy to make
out wind-pollinated species from insect-pollinated species.: The former
have inconspicuous jnflorescence with small flowers; the latter have deve-
loped a variety of effective contrivances for attracting a particular species
or group of insect species: The insects which are attracted to such flowers
act as agents for the transference of pollen grain from the, anthers of one
flower to the stigma of the other, thus leading to cross-pollination.
In many plants the flowers produce nectar secreted by special glands
situated in hidden niches at the base of the petals, and the petals are
so modified and arranged that the insect seeking the nectar has to come
in contact with anthers and stigma of the flower. In some the pollen
grains are sticky. in others the stigmas are sticky; the insect body parts
are also specially modified for such purposes. Some flowers have charming
colours to attract insects; others tempt them by their perfumes. This is
why some flowers have very spectacular petals but no perfume. and others
have very sweet smell but inconspicuous flowers. Thus, the brilliant tinge
and colour, aesthetic patterns and designs. enticing smell and aroma, and
tasty and nutritious nectar of the flowering flora were originally evolved to
seek thel partnership not of man but of insects. It is in comparatively
recent times that man has begun to act as an effective agent of selection
in the evolution. of the garden flora. The positive correlation between
the evolution of floral structure of certain plant species and the structural
peculiarities of certain insects is so perfect that naturalists have been able
to predict the presence of the one on the basis of observations on the
other. It is difficult to narrate the various corresponding modifications in
insects and plants specifically developed for ensuring cross-pollination, but
a few interesting examples are cited below.
Pollinator bees. Bees in general, and the honey bee in particular.
are the best studied insect pollinators. The honey bee has exploited its
role as nectar and pollen collector to the extent of developing an industry
20 AGRICULTURAL ENTOMOLOGY AND PEST CONTRoL.
of its own. It has perfected not only the art of collecting nectar from
flowers to\ convert and store it as honey but also the art of collecting
pollen on a vast scale for providing proteinaceous food for the bee colony.
In nature there is pollen enough and to spare, and the value of the honey
bee as an efficient pollinator is in no way reduced because it collects
pollen on a large scale for its own use. The hairy body of the bee gets
covered with pollen when it enters one flower after another, and then
the bee collects this pollen from all over its body by means of a spe-
cialized' brush on some of the sub-segment of the tarsus. which is the .
last segment of the hind leg. When this brush becomes fully laden, the
pollen is transferred to a special pollen basket situated on the tibia which
is the last but one segment of the hindleg, and the bee carries this load
of pollen in these baskets on the 2 hindlegs to its hive, where it is stored'
in special cells separately from honey. In this process, the cross-pollina-
tion necessary for seed setting in the flowers is also efficiently ensured.
The value of the honey bee for its pollinating activity can be gauged
from the estimate that during a period in which the honey bee contri-
butes honey worth Rs 5, its contribution to seed setting and fruit produc-
tion is worth Rs 100. In other words, the value of the honey bee as a
pollinator is 20 times as much as a producer of honey. The utilization of
the honey bee colonies has become a commercial undertaking in many
countries and 3 to 6 colonies an acre of a leguminous crop is considered
to be optimum. It is reported that in California 450.000 colonies were
employed for pollination purposes during 1955. Yet the honey bee is not
regarded as the most efficient pollinator. Other species of honey bees
such as Apis florea and A. dorsata are also useful.
There are a number of solitary bees belonging to the genus Nomia
and Megachile which have begun to be exploited on a large commercial
scale solely for purposes of pollination. Their habits have been carefully
studied, and it has been found feasible for man to enter into a kind of
symbiotic partnership with these bees. It is possible to persuade one of
these species to come and rest in an artificial hive, which can be easily
stored under controlled temperature conditions and taken to the field at
the proper time for the progeny to emerge and pollinate the crop. These
genera are well represented in India also, but their study is yet in a very
preliminary stage. There are also a number of other species of the bee
family which do yeoman service in the production of fruit and seed. The
carpenter bee (Xylocopas) in the plains, the bumble bee (Bombus) in the
hills, and Anthophora in both are most spectacular of the flower-visiting
insects. They are large-sized, long-tongued, and quite active. One
example of the value of these pollinators is afforded by the report that it
was impossible to get seed from red clover in New Zealand until bumble
INSECTS USEFUL TO MAN 21
bees were introduced from outside. The total value of insect-pollinated

crops in the USA is reported to be more than 4.5 thousand million doBars
annually.
Other insect pollinators. Besides the bee family there are many other
insects, such as the wasps, some butterflies and moths, some diptera and
beetles, which also carry out pollination; but the balance sheet of the
good and harm that they do is not available and is rather difficult to work
out. Hence it is not safe to call them friends of the farmer, and therefore
they are not being described here. I
I
It is, however, desirable to mention in some detail the iI11Portance
and activities of a chalcid wasp in the production of figs, for which this
wasp is comm9uly know~ as the fig insect. The course of events has been
well studied in the case of Smyrna fig. What is ordinarily known as the
fig fruit is botanically an inflorescence enclosed within a, fleshy pear-
shaped receptacle, the inner surface of which~is lined with innumerable
small flowers; the receptacle opens to the outside through a very small
miftce at the apka\ end. 'the Smym'll fig plouuce-& llf1~'j 1ema\e f!.owet'i>,
whereas the corresponding male-producing plant is known as capri fig.
For proper development of Smyrna fig it is necessary that pollen from
capri fig be carried by some agency to the stigmas of Smyrna fig. The
agent for this cross-pollination is a small chalcid wasp Blastophaga
peenes. The female wasp enters through the apical orifice of both Smyrna
fig and capri fig; but it is able to lay eggs only in the capri fig, and in
this the progeny develops up to the adult stage. The males are wingless,
and fertilize the females within the fig itself. The fertilized females come
out of the fig through the apical opening, with their body covered with
capri fig pollen grains. Subsequently, when these females enter Smyrna
fig they inadvertently deposit the pollen grains on the stiginas, and this
results in cross-pollination and proper fruit development. This absolute de-
pendence of proper production of Smyrna fig on the activity of the fig
insect was spectacularly demonstrated in USA. The Smyrna fig produced
there used to be of very inferior quality to that imported from Asia Minor.
and it was found that this difference was due to the absence of the fig
insect in USA so that fertilization of Smyrna flowers was not taking
place and consequently the fig fruit was not developing the proper flavour
and quality. After the fig insect was introduced, production of normal
figs was achieved.
A still more interesting example of symbiotic partnership is afforded
by what is commonly known as yucca moth (Pronlliba yuccasella), which
pollinates yucca flowers. The mother moth has a sharp-cutting ovipositor
with which it makes an incision in the ovary of the yucca flower and lays
an egg there. Thereafter it shows a very extraordinary behaviour in clim-
bing the stamen, scraping off a quantity of pollen grain, carrying it to

the pistil in which it has laid the egg, and actually thrusting the pollen
grain into the stigma, thus completely ensuring fertilization and develop-
ment of the ovules. The offspring of this moth eats a few of the seeds that
result from the poUination carried out by the mother. This is a very in-
teresting' caSe of symbiosis between an insect and a plant founded on a
well-calculated give-and-take basis.
Parasites and Predators

This is another group of insects which silently do immense servis;e
to humanity. They are the enemies of our enemies - the pests of crops -
and hence they are certainly confirmed friends of the farmer however
unspectacular, inconspicuous, silent and inappreciable their activities may
be. The two terms generally used for insect enemies of insect pests of
crops are parasites and predators; but these are as difficult to define
clearly as the terms "thief' and 'robber', although in most cases there is
no serious difficulty in deciding to which category a particular species
belongs.
The egg parasite Trichogramma. Many moths lay their eggs on the
foliage of plants, and the caterpillars that hatch feed either on the leaves
or by boring into the stem. There is a natural enemy of these insects
which destroys a proportion of their eggs. It is Trichogramma, a minute
cha1cid wasp belonging to the order Hymenoptera. The female wasp lays
its eggs within the moth egg; the wasp egg hatches into a legless larva
or 'grub', devours the contents of the host egg, and develops into adult,
which comes out of the emptied host egg from where otherwise the larva
of the moth would have emerged. The Trichogramma adults fiy about
and mate, and again the females lay eggs in the moth eggs. The principle
involved here is technically known as 'biologicaJ contl'ol' wherein one
living organism is employed to check or control another. The Tricho-
gramma parasitizes the eggs of a large number of moth species, of which
some are pests of crops, e.g., the sugarcane stem-borers, and some of
stored grain, e.g., the rice moth (Corcyra cephalonica). or the paddy grain
moth (Sitotroga cerealella). Moth pests of stored grain can be reared in
huge numbers under indoor conditions where temperature and humidity
can be maintained at the desired level. This makes it possible to rear
Trichogramma also on a very large scale all the year round. What is
actually done is to rear indoors one of the moth species in thousands,
cqlIect the eggs in millions, paste them on semi-stiff cards, expose them
to Trichogramma adults for parasitization (i.e., egg laying), hold the cards
at suitable temperature and humidity, and then hang them at suitable
places in the sugarcane field infested by stem-borers so that the Tricho-
gramma adults emerging from the eggs may fly about, search out, and
parasitize the eggs of sugarcane borer moths laid on sugarcane plants in
the field. In this way the good work which the natural population of
Trichogramma may already be doing in the field can be COIisiderably
supplemented by the Trichogramma population bred under room condi-
tions. The one important point to remember is that this kind of biologi-
cal control is feasible with Trichogramma only because it has a host which
can be reared indoors; otherwise there would be no point in rearing it
on sugarcane borers collected from the field, or in raising an acre of sugar-
cane for raising a population of borers and rearing Trichogramma on their
eggs for saving' just another acre of sugarcane elsewhere. The alternative
approach of ~earing the sugarcane borer itself on cheap artificial diet
indoors is still in experimental stage.
Parasites of aphids. Aphids or plant lic>e are prolific insects which
often densely cover the shoots and tender leaves of plants, suck the sap,
and thus weaken the plants on which they are found. There are some
parasites which exercise a certain degree of control on th~ natural abun-
dance of these lice. The parasite which has done 'the most spectacJ1lar job
in several countries including India is Aphelinus mali, which parasitizes
the woolly aphid Eriosoma lanigerum, a serious pest of apple trees.
A native of USA, it has been successfully introduced into some 30 other
countries in Europe, South America, Australia and Asia. It was imported
into India sometime in the late thirties and it has been so successful
that the woolly aphid has ceased to be a serious problem. The female
parasite lays eggs in the abdomen of the host aphid. The eggs hatch in
a few days, the larval period takes a few weeks, and the pupal period
about a week or so; the whole life cycle takes about a month for its
completion. depending on temperature. All stages are passed within the
body of the host aphid, which dies in due course, but by this time a
sort of liquid oozing out from the aphid body fixes the aphid securely to
the plant surface. After death the aphid turns black and becomes some-
what distended and hardened. The\ adult parasite cuts out a round hole
in this hardened body wall to come, out. This is an important example of
an exotic parasite proving highly successful. When such successes are
obtained in the field the repeated rearing and releases of parasites of
insect pests become unnecessary.
An indigenous example is Trioxys indicus, a wasp belonging to
the family Apidae of the Order Hymenoptera, which is parasitic on
Aphis gossypii attacking brinjal and cotton. The parasite first appears
in the field in November, and an increasingly large proportion of the
aphids are subjected to its attack in the following months. Another
parasite belonging to the same group is Aphidius smithi, which is a
natural enemy of the pea aphid Macrosiphum pisi in many parts of the
country, and is believed to be an important factor in keeping this pest
from becoming more serious. A remarkable control of the same pest
was brought about in some parts of USA when this parasite was intro-
duced and released there in 1959.
Larval parasites. The sugarcane borer has a natural enemy in
Stenobracon deesae, a slender graceful-looking wasp which has a long
sting or ovipositor. The female inserts this ovipositor into the holes
made in the stem of sugarcane (as well as maize and jowar) by the
caterpillars of the borer, stings the caterpillars, and lays eggs in them.
The eggs hatch into tiny parasite grubs, and these feed on nearby full:
grown borer caterpillars by lacerating their integument and imbibing \
the liquid oozing out, grow in size, from pupae, and then become adult.
The sugarcane borer has another parasite. Apanteles flavipes, a small
black insect. The female has a short ovipositor, and deposits its eggs in
only young borer caterpillars. The eggs hatch and 3 well-defined larval
stages are passed in succession before the grubs come out of the nearly
dead host caterpillars, spin cocoons; and then emerge as adults. The
parasite appears to tide over the cold winter of the northern plains as a
second-stage grub inside the hibernating caterpillar of the maize and
jowar stem borer, Chilo zonellus.
Bracon greeni lefroyi is the best known parasite of the spotted boll-
worm of cotton. The adult parasite is a small, brownish wasp with a
prominent dark spot on the upper surface of the abdomen. The female
wasp has a prominent ovipositor with which it probes and paralyses the
host caterpillar and then lays its eggs on the caterpillar's body. It is
capable of laying up to 219 eggs, hatching in 3 to 4 days into tiny,
creamy, legless grubs which lacerate the caterpillar's body wall with
their sharp mandibles and suck the body fluids that ooze out. The grub
period is shorter in summer than in winter, and there is a quiescent'
pupal stage before the adult, stage is reached. The emerging adults mate,
and the females continue the racial job of host-finding and oviposition.
Attempts have been made to utiliZe this species for biological control
of spotted bollworms.
A pupal parasite Trichospilus pupivora parasitizes the pupae of
the black-headed caterpillar Nephantis serinopa, a serious pest of the
coconut palm. The adult parasite is a yellow-brown wasp 1-2 mm
loni, with a small, nearly globular abdomen joined with the rest of the
body by a short, narrow waist or petiole. The female wasp lays a varying
number of minute eggs inside the host pupa by repeated puncturing with
the ovipositor. Each female may lay up to 236 eggs in 5 pupae or so:
and more than 1 parasite may lay eggs in the same pupa; however, up to
INSECTS USEFUL TO MAN 2S
866 individuals have been recorded to emerge frbm 1 host pupa of

Plusia peponis and up to 595 from Prodenia litura pupa. The eggs hatch
into tiny grubs, which feed on the contents of the pupa, develop. and
themselves become pupae in about a week from the time the eggs are
deposited. After 8 to 10 days in the pupal stage, the parasites become
adults; thus, a little more than a fortnight is required for developme~t
from eggs to adults. It is believed that the adult parasites mate within
the hoIIowed-out shell of the host pupa, and the males rarely come out
f
of it; the females emerge from the remains of the host and searcn out fresh
host pupae to carry on their appointed task. This parasite a1so attacks
pupae of some other ~oths, such as Sylepta derogata and Prodenia
litura. It thrives/best at moderate temperature (22°-25°C) and. rather high
humidity (92-94%); thus, it is active in coastal regions of Kerala from
October to February' but becomes rather scarce during the hot and dry
months of March to May; for the same reason it is rather scarce in the
drier coastal regions of Tamil Nadu.
This parasite has been employed in the biological c,bntrol of the
coconut pest Nephantis serinopa. The pupae of the pest' are collected
from the field and kept in cages for the emergence of the parasite-s,
which are then liberated in areas of infestation. When the pupae of
Nephantis are scarce, those of other Lepidoptera, such as Sylepta and
Prodenia, are reared for use as host by the parasite.
The main points about the use of this parasite for biological control
are that (i) each individual parasitizes on an average 2 or 3 pupae; (ii)
the female lays on an average 100 to 150 eggs; (iii) the life-cycle can be
completed in 16 to 17 days-22 generations were actually reared during
1 year at Calicut; (iv) it can fly up to 5 km; and (v) it can breed in a
number o~ host species. Its main defect is that it is very susceptible to
low humidity,
Parasite of adult flea beetle. Microctonus indicus is a parasite
which attacks the adult stage of its host, Phyllotreta cruciferae, a beetle
pest of various cruciferous vegetables. The female parasite inserts its
ovipositor into the thin lower surface of the thorax of the beetle and
deposits eggs in its body cavity. The eggs hatch into grubs, and of the 2
or 3 grubs in a host (each from 1 parasite egg) only 1 successfully
completes its development up to the final larval instar owing to internecine
struggle. The fully grown larva issues out of the beetle by tearing a
hole in the posterior part of its body; the quiescent or pupal stage is
passed outside the host, and the adult parasite emerges from the pupa.
Predator beetles. The most useful group of predators or entomo-
phagous (insect-eating) insects is that of the ladybirds or ladybeetles or
ladybird beetles belonging to the family Coccinellidae of the insect order
Coleoptera. The adults are medium to fairly large-sized insects, round or

oval in shape; they have prominent, brightly spotted, hard, leathery
forewings covering the abdomen. The ladybirds are perhaps the best known
of the beneficial insects; the sobriquets 'ladybirds' and 'ladybeetles' were
probably given them as terms of endearment, possibly as recognition of
their good work as destroyers of plant lice, scales and mealy bugs in-
festing various crops.
Cocdnella septempunctata is a common ladybeetle found in many
couhtries including India. The adults are fairly large, and the leathery
forewings are brownish with 7 black spots on each. They are long-lived,
and females lay up to a few hundred eggs each over a period of d~ys.
The eggs are elongate and orange-yellow and are laid in clusters. each
comprIsing a few eggs, on plants infested with plant lice, mealy bugs and
scale insects. Small, dark, very agile grubs hatch out in about a week and
actively search out and devour aphids and scale insects. In about a month
the grubs lose their activity and transform into dark-coloured pupae which
can be seen sticking to leaves and stems of the plants on which the.
grubs once crawled about. Adult ladybeetles emerge from the pupae in
about a week, and they too devour plant lice, mealy bugs and scale
in&lects. Unlike the hymenopterous parasites discussed earlier, e.g.,
Trichogramma, Apanteles, Aphidius, Microctonus and Trichospilus, whose
immature stages alone feed on other insects, both immature stages and
adults of the coccinellids devour the hosts.
Another important ladybird beetle is Rodolia cardinalis, which saved
the citrus groves of California, and later of many other parts of the world,
from total destruction by the cottony cushion scale, ]cerya purchasi. It
was this beetle, popularly known as vedalia, that first made people take
cognizance of the possibilities of controlling insects biologically. After
it was introduced into California from Australia in the last decade of
the century, the control of the scale was so spectacular that this beetly
was introduced in many other countries. The scale was widespread in
peninsular India, and the lady beetle was imported in 1928 from Australia.
Large-scale rearing and releases of the predator were taken up in 1946,
and by 1954 a very satisfactory contro~ of the scale was obtained. The
female beetle lays on an average 300 brick-red eggs. which hatch in about
4 days into dark little grubs; these moult thrice in about 9 days, then
pass about 4 days in the pre-pupal stage and about 9 days in the pupal
sta_ge belfore emerging as adult beetles. Thus, the life-cycle takes about
25 days. Both adult and grub stages of the beetle feed voraciously on
eggs and young ones of the fluted scale, and it is estimated that on
an average the beetle grub during its existence of about 9 days devours
more than 200' eggs of the scale.
Insect Enemies of Weeds

Some insects are useful to the farmers directly or indirectly in yet
another way. These are the insects that feed on weeds and destroy them and
thereby free arable land for cultivation or pasture. Weed control by
insects came into its own after the splendid results achieved by the in-
troduction of the moth Cactoblastis cactorum into Austra1ia from
Argentina for controlling the cacti (Opuntia spp.) which had threatened
to destroy Australian agriculture (and livestock) through their fantastic
usurpation of nearly 60 mi1lion acres of land.
In India many weeds pose a threat to arable and pasture lands. The
cacti Opuntia vulgaris, Opuntia dilleni and Opuntia elatior "had been
introduced into India in the 18th century for culturing "the cochineal
insects. In du'e course the cochineal dYE! industry died out because of the
advent of synthetiCs, but the' cacti gradually became not only a nuisance
but a real danger to agriculture. A mealy bug named Dactylopius in dicus,
which had been introduced perhaps inadvertel]tly along with the cacti,
was found to exercise a certain degree of control on Opuntia vulgaris.
In 1924-1926 another mealy bug, Dactylopius tomentosus, a native of
Mexico already imported into Sri Lanka, was introduced in Tamil Nadu,
and within a few years it cleared more than 100,000 acres of land en-
croached upon by Opuntia, dilleni; it also brought about some control
of Opuntia elatiQr.
The mealy bugs (Dactylopius spp.) are small sessile insects, en-
crusted in a white meal or powder (whence the name mealy bug); they
suck the juices of the cacti by inserting their sharp mouthparts into
the fleshy parts of the plants, which eventually wither away.
Insects Yielding Products of Commercial Value

The three universally appreciated insects are the honey-bee, the lac
insect, and the silkworm. Honey extraction from wild hives is definitely
a case of exploitation, but modern bee-keeping provides an illustrious
example of science enabling man to enter into a just and ethically sound
partnership at least with one species of honey-bee. The relationship
with the lac insect also need not De necessarily abhorrent, but that with
the mulberry silkworm particularly is a simple case of a bio-industry
in which, as in many other cases, an animal species is reared and cultured
only to be ultimately killed and utilized.
The Roney-bee
There are 3 species of honey-bee in India, viz., the large rock-bee
(Apis dorsata), the little bee (Apis florea), and the medium-sized Indian
honey-bee (Apis cerana indica). An gather nectar and pollen from
numerous -species of flowering plants and in this process bring about

pollination of the flowers. Man' has been consciously interested since
time immemorial in the nectar collected by these insects and later on
appreciated the yeoman serVice they do as pollinators. Therefore, scienti-
fically minded people and later on professional scientists have tried to
study and. ,befriend these insects and enter into a mutually beneficial
partnershIp with them; but so far they have succeeded in doing so only
with the medium-sized honey-bee, Apis indica, the European counter-
part of which is Apis melli/era. The rock-bee is too ferocious to deal
with in a friendly way. It builds large open single combs measuring up
to 1.5 or even 2 metres from side to side and 60 to 120 cm in depth.
These combs are suspended from rocks, ceilings of tall buildings, and '.
branches of tall trees. It is the combs of this species with which most
people are familiar. There are records of fatal attacks by rock bees on
man and animals, and even armies are said to have been repulsed by
swarms of these bees.
The little bee builds very small combs which are suspended from
bushes, trees, roof corners, etc. People have not considered it worth-
while to devot~ much attention to this species. The medium-sized bee
is reasonably docile in its habits and, therefore, this species has been
carefully studied. Before these studies, the honey collectors used to deal
with these bees also as mercilessly as they do even today with the
rock-bee, i.e., they used to practically overpower the bees with smoke,
etc., particularly during night, and cut away the combs for collecting
honey; this caused much wanton destruction of the bees, their young
ones, and their combs.
Modern partnership between man and medium-sized honey-bee.
This partnership is the basis of the modern industry of apiculture. and
its scientific principles are based on the following information about the
habits and life economy of the medium-sized honey-bee. The Indian'
species is found all over the country. Unlike the other 2 Indian species,
it builds several parallel combs, generally in the hollow of tree trunks,
hollows in rocks, and other common closed and covered places. The
bee colony is a highly sophisticated social community with rigorous
laws, particularly about the division of labour between groups or castes
of individuals. The division of labour is so perfect that it has its imprint
eve)1 on the structural peculiarities of different castes. There are 3 main
castes, (i) the queen, which is usually a single individual in a colony,
(ii) the workers, numbering 20,000 to 30,000, and (iii) the drones, gene-
rally not more than a few hundred, and at times none.
Queen. The queen is the onlv fully developed female; it is the mother
of the colony, readily distinguishable from the other members. From
the workers it differs in being bigger in size and having an elongated

shining black abdomen, which is striped and triangular in shape in the
workers; because of the prop<lrtionately longer abdomen, the wings of
the queen appear shorter than those of the worker. The queen does
nothing but reproduce; during the period of highest activity it may lay
as many as 1500 eggs a day.
Drones. The drone is male; its sole function is to fertilize the queen;
it dies after mating. It has a black and rectangular abdomen with a
blunt free end without a sting. It has large eyes, which meet; over the
head. Drones are reared and tolerated only during the breeding season
in spring and autumn when new queens are there to be fertilized. When
not required, before mosoon and during winter, they are driven out of
the hive to sta~ve and die; hence at certain times there may be no
drone in the colony. ,
Workers. T4e worker-bee is an imperfectly developed female, unable
to reproduce but possessing all the maternal instincts. It ~s smaller in
size than' the queen and the drone. The workers do all the, work of the
colony, build combs, gather nectar and pollen and certain' other subst-
ances used in the hive, and care for the brood. They keep the, hive
clean, and regulate its temperature by fanning their wings. From nectar
they make honey. To perform these duties satisfactorily, the workers are
provided with the necessary structural adaptations. Each worker-bee per-
forms different types of work in its life-time, and its body becomes
equipped for various duties in succession as it advances in age, i.e., it
passes through a series of professions, changing its activity as it grows
older. Since new broods of bees const~ntly arise, at any time there are
bees engaged in different tasks of running the hive.
Just after emergence as adult, and for a period of about 2 weeks
thereafter, the worker's principal task is to act as a nurse. It brings honey
and pollen from the storage cells to feed the queen, the drones, and the
larvae. As it grows older, it becomes able to produce wax from glands
on the underside of its abdomen and begins to work on the construction
of the combs. In addition, it performs the job of house cleaning, and of
standing guard at the entrance of the 'hive against any intruders. About 3
weeks after emergence it begins to attend to all outdoor duties and collects
and brings into the hive nectar, pollen, water and propolis (bee-glue).
When on outdoor duty, the workers divide into searchers and gatherers.
The former fly around and bring back news of available food and commu-
nicate it to the latter by performing a dance indicating the exact distance
and direction of the source. The gatherers then rush to the indicated spot
and bring back loads of pollen and nectar. The variety of the dance
forms the language of the bees. The worker-bee labours hard for the
colony till it gets exhausted and dies.

Life 'history. The queen lays a small white egg, slightly curved on one
side. at the bottom of a cell. In appearance and· size the eggs of different
castes are alike, but they may be either fertilized or unfertilized; for the
queen is capable of laying both the types. The unfertilized egg produces
a male (drpne); the fertilized egg produces a female, which may become
either a, worker or a queen depending on the food available during the
laryal stage. The eggs are, however, laid in separate cells provided for
rearing different castes. The egg hatches in 3 days and a tiny larva emerges.
Remaining in the cell, the larva feeds voraciously on the food supplied
by nurse-bees. When fully fed, the larva spins a cocoon in the cell which
is then closed with a waxen cap; inside which the larva transforms' jnto
a pupa'l In the case of larvae which are to give rise to workers, the larval
period lasts for 4 to 5 days. After 11 to 12 days of pupal period the
worker-bee emerges. cutting open the cocoon and the wax cap of the
cell. The total period of development in the worker-bee is about 18 to
20 days; it varies in the other two castes.
The young larvae are fed a special food called 'royal jelly', a secre-
tion of special glands located in the head region ()If the nurse-bee; it is
supplied to young larvae of the 3 castes up to the third day of their
development. Thereafter the composition of food given to the larvae
destined to become queens, workers and drones differs. It has been esta-
blished that differentiation into worker or queen depends on the quality
and quantity of food fed to the larva. Partial starvation from about the
third day onwards results in the production of worker. The larva destined
to form the queen has constant access to abundant food ,and is reared in
large cells specially made for the purpose at certain times of the year
along the lower margin ~f the comb.
When a new queen emerges in a hive she may be killed by the old
queen or may herself kill the old queen. Frequently, however. the old
queen leaves the hive and takes with her a large number of workers to
found a new colony. The division of a colony in this manner is called
'swarming'. The new queen mates with a drone in the air. After some
period it starts laying eggs, and routine activity in the colony starts all
over again. A queen may live for 2 or 3 years.
Comb. The comb is made of wax secreted from a set of 8 glands, 4 on
each side of the underside of the worker's abdomen. It comprises numerous
. hexagonal chambers or cells built on the sides of a central midrib. The
midribs of adjacent combs are at a distance of 3.2 to 3.5 cm depending
on the variety of the honey-bee. The cells are utilized for rearing the
young and as store-rooms for pollen and honey. The cells meant for
rearing individuals of different castes are different, those for the workers
being the smallest, those for the queen the largest, and those for the
drones intermediate in size. Both worker and drone cells can b~ used
for storing honey and pollen. The arrangement of cells in the colony
is quite orderly. The brood area consisting of cells in which the workers
are being reared is generally in the lower part of the comb. In the top
and the sides of the brood area is a band of cells stuffed with pollen,
and above this is the band of cells containing honey. The comb in the
middle of the hive contains the largest brood area and on the sides the
brood area becomes progressively small. Thus, the brood space of the
colony is a .kind of sphere in the lower part of the hive.
Apiculture. A good working knowledge of the honey-bee was acquired
by man right,in the beginning of human history. Even neqlithic man is
saia to have/been acquainted with its value. Early Egyptians practised
commercial bee-keeping, even migratory bee-keeping. The Greek also
practised this art. So did the Romans. Aristotle wrote on bee-keeping. In
India there is mention of the use of honey right from the Vedic period.
To primitive man the honey, the pollen, the brood, and:even the young
bees meant food. It was only during the last 100 years that bee-keeping
became sophisticated and civil, after Langstroth invented his beehive
with moveable frames. Later, it became practicable to keep boxes over
the hives in which surplus honey was stored by the bees. This led to the
construction of the beehive in 2 storeys, so that honey could be stored
by bees in the upper storey (super) and brood-rearing could be carried
on in the lower storey (brood chamber). This further made it possible to
remove the movable combs containing honey in the super without
disturbing the brood chamber. The question of killing the brood or the
bees for extracting honey became a thing of the past. A further improve-
ment was to extract the honey in a special extractor without destroying
the comb, which could be put back in the hive to be filled with honey
again. This meant a lot of saving of labour for the worker-bee, which
could devote the energy thus saved to collection of nectar and pollen.
It has been estimated that making 1 kg of comb is equivalent to pro-
duction of 6 to 10 kg of honey.
All this improvement and sophistication has been possible by care-
fully studying the habits and requi~ements of bee. These studies have
now made it possible even to understand the language of the bees. It is
established that the honey-bee has its own language, which is not arti-
culate but akin to the language of scouts. This is actually called the
dance language of the honey-bees. The patterns of dance have definite
meaning which can be decoded now by man.
The Lac Insect

Many of us do not connect a stick of sealing wax, a gramophone
record, or French polish (and other varnishes of the non-synthetic kind)
wi~h insects. Yet, all these in reality come from an insect known popularly
as the lac insect and technically as Laccifer lacca. Where and how this
insect lives -and produces lac makes a fascinating story. While lac c.ul ti -
vation is an industry, being increasingly run on highly commercialized
scale, the basic limit of it is the lac insect; hence the entomologist is
directly interested in lac cultivation.
Lac is produced by the insect as a brown resinous encrustation. The
reddish encrustations one sees on such trees as kusum (Schleichera o[eosa),
palas (Butea monosperma), and ber (Zizyphus mauritiana), and occa-
sionally on Ficus spp., are actually the secretions of the lac insects. The
first 3 plant species are the most favoured hosts of the lac insect, although
it is also found on a variety of other plant species.
The female insect lays eggs inside the lac cell with which it is
covered, and the eggs hatch immediately after laying. Each female pro-
duces about 300 minute. soft-bodied, crimson larvae. Larval emergence
in large numbers takes place at certain times of the year. Immediately
after emergence the larva crawls over the host plant and rests only when
it finds a suitable place on a shoot. It pierces the plant tissue with its
needle-like mouthparts and sucks up the juice of the plant. Once settled,
the larva does not move about and goes on depositing resin around itself
except about the mouthparts, breathing pore, and anus. The larvae grow
and moult 3 times before becoming sexually mature males and females.
Female larvae have larger and thicker cells than male larvae. Both kinds
lose their legs, antennae and eyes after the first moult, but the males
eventually regain these organs after they turn into pupae. The duration
of the development varies with the prevailing climatic conditions. The
adult males are of 2 types, winged and wingless. They move about in
search of females, which remain fixed to their original sites. The males
live for a short period (62-92 hours) during which they mate and fertilize
the females. From this time onwards lac is secreted by the female at a
faster rate, and the size of the insect as well as that of the enveloping
lac cell increases rapidly to several times the size of the male lac cell;
therefore the female is the chief source of lac. This state of activity
)asts for a varying period according to climatic conditions. As the time
for egg laying approaches. the body of the female contracts on one side,
gradually vacating space inside the cell in which eggs are laid.
The foregoing account of the life-history of the lac insect constitutes
the scientific basis of the lac industry. As the lac insects settle in close
proximity on the branches of the host tree, their profuse encrustations
make a sort of continuous sheet; and when the young larvae of the next
generation have crawled out to new and younger shoots the old encru-
station, which is practically lifeless, is scraped off and processed. How-
ever, in systematic large-scale lac cultivation the young larvae are helped
by human agency to reach twigs of plants specially made suitable for
fresh infestation. Host trees are pruned so that they may throw out new
shoots at the proper time. Then the twigs of the old host plant are cut
a few days before larvae of the next generation are expected to emerge,
and these twigs (carrying seed lac) are hung on fresh plants processed
as above for the young ones to come out and infest fresh shoots.;
Lac cultivation is practised in Bihar, Madhya Pradesh, Uttar Pradesh,
West Bengal, Assam, Orissa and Maharashtra, the first 3 states produc-
ing the largest qU:;lntities 6£ commercial lac. The major lac factories are
located in Biha~, Madhya Pradesh and Calcutta. Lac export~' earn about'
Rs 14 crore of foreign exchange, and hence the cultivation of lac has a
direct impact on our trade balance. There i;; scope for considerable
improvement in this bio-industry. Research on. scientific fines can do
much to make lac cultivation a profitable business.
Lac cultivation in India is of great antiquity, references to it being
found in the Vedas. In the Mahabharata is recounted the story .of the
abortive attempt by the Kauravas to burn alive the Pandavas in a man-
sion built of lac as denoted by its name laksha griha. The Sanskrit word
for a hundred thousand and for lac is the same, viz. laksha, possibly
to indicate that lac is produced by large or countless numbers (of insects).
The Silkworm
We do not give a humane treatment to the silkworm, although we
rear it on industrial scale and use its product for a prosperous industry.
The silkworm from which we obtain silk, the king of fabrics, is the
caterpillar of the moth Bombyx mori. The moth is medium-sized, with
a soft body and creamy white wings. In spite of the wings being present
the moth has practically lost the power of flight because it has been
completely domesticated for ages. The knowledge of the production of
silk from the silkworm is from time, immemorial; it is on record that an
Indian ruler sent silk fabric to an Iranian ruler mOre than 6000 years ago.
Since then this insect has been reared in captivity.
Silk is secreted by the salivary glands of fully grown caterpill;!'fs.
and this secretion on exposure to air at once hardens into a fine delicate
thread. The caterpillar, which feeds exclusively on mulberry leaves (hence
known as mulberry silkworm), spins out this fine thread to make a cocoon
for its own protection in the pupal stage. Man has, however, found means
to utilize this thread for his own purpose.
Life history. The female moth laysl 300 to 400 whitish eggs. The cater-
pillars on hatching start feeding on fresh mulberry leaves and begin to
grow. In 4 to 5 weeks they become fully mature and are yellowish white
with a cylindrical elongated body about 5 em long and having a small
horn at the hind end. When ready to pupate, each caterpillar spins a
yellowish-white cocoon inside which it pupates. About 10 to 12 days
thereafter the moth is ready to emerge. Just before emergence, the moth
secretes a fluid that softens one end of the cocoon from where it squeezes
out breaking the silken strands. The moths live for only 23 days and do
not take any food. They mate and lay eggs; and thus the life-cycle is
repeated.
Depending on the number of life cycles passed in a year, there are
2 main races of the silkworm, univoltine and multivoltine. In the univol-
tine silkworm there is only 1 life-cycle in a year, i.e., the eggs remain
in diapause for 5 to 7 months. This race is reared in Italy, France, J~pan,
China, and India (mainly Kashmir). In the multivoltine silkworm, on
the other hand, there are many life-cycles in a year, there being no diapa-
use in the egg stage. The multivoltine silkworm found in India and
commonly reared in Karnataka, West Bengal, Assam, and Tamil Nadu
has 4 to 7 life-cycles in a year.
How silk is collected. Each cocoon is composed of a single continuous
thread, which may be 200 to 350 metres long in the case of multivoltine
races. If the moths are allowed to emerge, this single unbroken thread
would be dissolved and broken into tiny pieces and become useless.
Therefore, about 10 days after the cocoon is made, the pupae are killed
by dropping them into hot water, by steaming, by exposure to the heat
of the sun, or by fumigation. The cocoons are sorted according to
colour and texture and soaked in warm water to wften. When the silk
thread becomes loose, it is skilfully unwound. The threads from several
cocoons are wound together to form reels of raw silk, which is then
processed to bring out the lustre; finally it is spun. It has been estimated
that about 25,000 cocoons (consume about a ton of mulberry kaves) give
1 Ib of silk.
Some oth~r silkworms. Silk in nature is produced by caterpillars of
several species of moths, but there are only a few from wh:ch it can be
utilized commercially. The important ones in India besides the mulberry
silkworm are the eri silkworm, the tusser silkworm, and the muga
silkworm. Mulberry silkworm is the most important because of the
superior quality of silk obtained from it and the ease with which it can
be handled. Tusser and muga silkworms are found wild in nature.
Silk industry in India. It is an important industry in India producing
la:r:ge quantities of silk for. our own use and for export to many countri-
es in the form of silk fabrics. The total production in 1962 was 17.80
lakh kilograms, Silkworm-rearing thrives primarily as a cottage industry.
CONSERVATION AND UTILIZATION OF
INSECT FRIENDS OF MAN
Obviously, 'it does not stand in need of any serious pleading to
state that every effort should be made to conservCi the friendly fauna of
the insect world, and that special precautions are Icalled for to avoid as
much as possible any harm to this section of' the class Insecta.""
This consideration makes the task of pest-control scientists a very
difficult and delicate affair. Selective killing of the insect pests and pre-
serving the friendly insect when the two are rather inextricably mixed
together is at present only an ideal, to strive for. This is particularly true
in the case of chemical control of insect pests. All the same, scientific
research during req:ht years has been revealing a number of promising
avenues for realistic approaches to the ideal we have in view. Informa-
tion on all these aspects is readily available in liter~ture. - Here only a
few such ideas are recorded as do not seem to be quite common in
literature or do not seem to be adequately appreciated.
Pollinators
Pollinators are different from parasites and predators of insect pests
in certain respects. First the activities of pollinators are more restricted
in time, i.e., to the flowering period of the crop. Hence, the general re-
commendation that as far as possible chemical control operation should
not be undertaken during the, flowering phase of an insect-pollinated
crop. No 'such general statement is possible in the case of parasites
and predators. Secondly, the activities of pollinators are in favour
of the other party-the flowering plants which they deal with-where-
as the reverse is the case with parasites and predators whose acti-
vities are against the host and prey whom they parasitize and kill. The
result is that flowers have evolved, and are evolving, various mechanisms
of colour, design, scent, ete. to attract pollinators even from a distance,
but the reverSe is the case with parasite hosts and preys whose survival
depends on their capacity to evade and hide. The degree of specificity in
insect-plant relationship as far as cross-pollination is concerned is of a
low order. Generally speaking, a plant can be cross-pollinated by more
than 1 species of insect, and an insect species can gather nectar from more
than 1 species of plant. In this regard this relationship is more akin to that
between a prey and a general predator than to that between a host and
a specific parasite. The implication of these seemingly insignificant
differences is that whereas the efficiency of a specific parasite'S activities
is highly density-dependent and is reduced with reduction in host popula-

tion, the activities of pollinators are likely to be much less dependent
on population density of flowers. However, since scientific investigation
so far carried out on pollinators is very inadequate, our knowledge
about them is meagre and imperfect.
Parasites and Predators

Much work has been 90ne bn parasites and predators and on their
conservation and use for biological control of insect pests and weeds; yet
there are some very significant surprises in the development of this branch
of pest control science.
(a) Inadequate appreciation of biological control. Paul De Bach, a
noted authority on the subject has recently edited an exhaustive and
useful treatise, 'Biological Control of Insect Pests and Weeds'. His
opening satatement, which depicts the factual pDsition of. the moment,
is: "The biological control of insects, mites and weeds has received
great and enthusiastic acclaim during the past 70 years with highly
successful practical results achieved in over 60 countries around the
world. In spite of this, some few scientists have looked upon the method
with incredulity." This certainly does not depict a very happy situation
in a scientific field. On the theoretical side there exist thought-provoking
contributions of great scientific value on the basic phenomenon of insect
parasitism, host parasite popUlation oscillation, etc., but they have not
been put to actual use in the practical field. On the other hand, in the
field of practical biological control there exist a large number of classical
and spectacular successes, and a much larger number of disappointing
failures, but little effort has been made, if at all, to adequately analyse
these successes and failures. Under these condi~ions it is no wonder that
De Bach was reminded of the following warning of an earlier scientist:
Reasoning unwarranted by facts, and facts not correctly and sufficiently
reasoned out, are equally worthless and dangerous for practical use."
However. De Bach's treatise will go a long way in correcting the situa-
tion, and those interested in the subject will be well advised to study
his book.
(b) Inadequate appreciation of the role of predators. Insect predators
were the first to attract the attention of growers and orchardists. The
Chinese had started since very ancient time purchasing and utilizing the
predaceous red ant, Oecophylla smaragdina, for the control of the pest in
citrus gardens. This ant is very common in Indian mango gardens. A
similar practice is reported to be common in the Middle East where
predaceous ants are transported from mountain regions to the plains
for control of date-palm. pests. Coccinellids have been known for their
good work for centuries, and it was the introduction of the predator
coccinellid vedalia for the control of the cottony-cushion scale that for
the first time established biological control as a valid branch of pest
control science. The first discovery of insect parasitism is credited to
Aldrovandi, who in 1602 mistook the cocoons of the parasite Apanteles
glomeratus on the larvae of the cabbage butterfly to be- eggs of some
insect, and it took another century before I
Vallisnieri in 1706 correctly
interpreted this as' a caSe of insect par,asitism. Today the situation is
that parasites have received much more investigational attention than
predators. In the field of practical utilization, too, it is reporte{l that of
95 species of entomophagous insects iinported and established in the
USA 81 are parasites and 14 predators; and that two-thirds of the
successful cases of biological control are due to parasites. Obviously, these
lop-sided statistics are the result of inadequate appreciation of the
role of pr~dators as compared with that of parasites. The reason appears
to be 'that parasites remain associated with the pest, which is collected
in parasitized stage, and often the parasites can be seen to emerge there-
from. On the other hand, the predator eats away the pest and is rarely
observed in the actual process of eating or harming it; hence the pre-
dator does not attract enough attention. This is all the more true in the
case of nocturnal predators. The extent to which predators bring about
reduction in the population of harmful species has to be appreciated
only through circumstantial evidence, as was done in the case of the
biotic theory of locust periodicity, and the tentative ideas thus deve-
loped have to be tested by specially planned critical experiments.
(c) Dosage of parasites in experiments on biological control. Many ex-
periments have been carried out in various parts of the world to control
particular pests by releasing parasites specially bred for the purpose,
but the success has not been spectacular. It is now being slowly realized
that the USe of indigenous parasites can have more chances of success
on an inundative instead of inoculative basis, and that one of the major
causes of frequent failures is insufficient number of parasites released.
Few efforts haye been made to determine the number of parasites
needed to bring about the desired degree of control at different popula-
tion densities of the host. In other words, there does not exist adequate
information on dosage requirements in the field of biological control,
and it is not surprising that the results have been erratic or discouraging.
Planned experiments should be carried out to determine the number of
parasites necessary to control a particular pest at its different population
densities, and the information thus obtained should be utilized in deter-
mining the number of parasites to be released under any particular situa-
tion. In fact, we should go on increasing these numbers as the host
population goes down so that the efficiency of the parasite population

to bring about effective control of the host is not impaired by lowering
the host population density. With improved techniques for mass rearing
of parasites, we should aim at making their efficiency independent of
host densities by flooding the areas with parasites, just as the screw-
worm was eradicated by flooding with sterilized males. The rearing of
screw-worm on animal tissues, on a large industrialized scale has led to
parasite rearing also on a commercial scale.
From the foregoing it will appear that there remains much to be
improved in the methodology of experiments on biological control, and
the gloomy picture that often emerges from inconsistent results recorded
in literature is in most cases probably due to some defect or the ~th:er
inherent in the experimeur. For example, experiments with Trichogramma
against borers of sugarcane are reported to have given inconsistent results
in India; but actually these experiments have shown that Trichogramma
releases resulted in significant increase in parasitization in certain areas
but not in others. Adequate attempt was not made to interpret these
results against the ecological background of the areas, otherwise most
probably the interpretation would have been that the performance of the
parasite depended on the climatic conditions of the region where it was
released. It seems to be almost certain that if ecological aspects are pro-
perly kept in view the percentage of success in biological control projects
can be significantly raised. The failures so far appear to be failures of
those engaged in these operations rather than of the principles involved.
Utilization of parasites and predators for the control of insect pests
and weeds has to be tackled in one or more of the following ways.
1. Introduction of ,exotic parasites and predators. This consists of im-
porting into an ecological region all such parasites and predators as are
not present in the region and are likely to be of use on prima facie basis.
For this purpose there now exists a world-wide organization, the Common-
wealth Institute of Biological Control, through which such insects can be
obtained and tried. They can also be obtained directly on exchange basis.
2. Utilization of indigenous parasites and predators. This requires
a study of the indigenous fauna to understand how the efficiency of each
individual species can be increased. This can be attempted in a number
of ways.
(i) Conservation. This requires such efforts as are necessary during
various operations of crop husbandry in order to ensure that parasites
and predators are not adversely affected. An example from mechanical
control is that the egg-masses when collected may not be destroyed but
kept in the field itself in a wire-gauze cage from which the minute para-
sites can fly out but not the various stages of the pest host. Similarly, there
are various precautions possible in the caSe of chemical control. Such

examples will be found under the sections dealing with integrated control.
(ii) Releasing the parasite in suitable dosage. The parasite releases
may be (a) on inundative basis, i.e., in such large numbers as
may be expected to attack almost every individual of the pest pop,lliation.
or (b) on inoculative basis, i.e., in numbers just enough to overpower the
pest. The success of such operations would naturally depend on the amount
of care taken to ascertain the correct strain of the parasite. the optimum
number to be released, the correct timing of releases, and the suitability
of the habitat where release has to be made. '
(iii) Release of parasites along with alternative hosts. In. the case
of parasites, p~rticularly-,those that are rather specific. it is likely that the
parasite may -get a setback due to the low population of the host. In such
a case it is worth-while trying to increase the host population by providing
an alternate host. In the case of Trichogramma it should be feasible to
increase the host density by hanging in a fie!d of sugarcane a few cards
containing Corcyra eggs so tt~at those parasites that are not able to find
the borer eggs may lay their eggs in Corcyra eggs. This was tried earlier
with the primary host, i.e., the pest itself, but the farmers were reluctant
to allow additional pest population merely to give a fillip to the parasite.
There will be no objection to the use of eggs of an alternate host which
is not a pest in the field; the larvae emerging from unparasitized eggs are
bound to die without doing any harm.
(iv) Genetic improvement. Genetic improvement is as much possible
in parasites and predators as in crops and livestock. One fear has been
that the improved strains developed in the laboratory are likely to be
competed out when released in nature. This is quite a sound argument,
but We can overcome the difficulty once we decide to maintain the im-
proved strains in the laboratory and use them on inundative basis in areas
and against pests for which they have been specially developed and bred.
We can easily maintain a number of strains of each useful species spe-
cially developed for various situations that are likely to arise.
These are' only some of the ways in which parasites and predators
can be fruitfully utilized.
CHAPTER 4
ORIGIN OF INSECT PESTS
THE earliest fossil remains of insects are traceable up to the Devonian

deposits of the Palaeozoic epoch, and the extent of evolution which this
earliest-preserved insect fauna obviously had already undergone has led
scientists to conclude that they must have originated latest by the Silurian
period, i.e., about 500 million years ago. Even the oldest fossil insects yet
discovered were fundamentally not very different from these insects (11-
though their organs were simpler and less specialized. Hence, conside!_irig
the rate of change that insects have undergone from the Devonian period
up to the present day it is quite rational to conclude that the origin of
insects as land animals coincided with the rise of land plants from their
marine ancestors sometime during Silurian or Ordovician period. On these
considerations the general view is that the ancestors. or at least the grand"-
ancestors of insects were marine forms and were feeding on marine plants.
Later, when land masseS emerged from under the seas, the marine plants
on such land masses had to respond to this major change in their en-
vironment by developing stomata on their vegetative parts. A similar
adaptation is supposed to have taken place in the marine ancestors of
insects which, after a preliminary period of cutaneous respiration through
the general body, developed spiracles and tracheae for aerial respiration.
From such simple beginnings the evolution of insects seems to have
closely followed the evolution of their food plants, and the first record
of winged insects almost coincided with the record of tall forest trees in
the upper Carboniferous period, about 270 million years ago. In other
words, the development of insect wings was most probably triggered by
the increasing tallness of their food plants, i.e., the height from the ground
of the foliage on which they tried to continue feeding.
Many of the world's rocks reveal on explorative digging the presence
of a vast variety of animals and plants which got buried during the natural
process of sedimentation in aquatic environment or through a few other
specialized processes, and have been preserved in what is known as
fossilized form.
Modern science has made it possible to determine fairly accurately
tne time when these plants and animals got buried. This is done with
the help of a new radio-geological technique which is based on the fact
that all animals and plants have a certain proportion of radio-active
carbon in their bodies. This proportion of radioactive carbon remains
fairly constant during life, but it begins to decrease after death at a
I
ORIGIN OF INSECT PESTS 41
known rate. Hence, by determining the proportion of radioactive carbon

in these fossili2:ed forms it is possible to determine the time when they
lived and died. The time scale on which these determinations are made
is in millions of years. According to these determinations, the oldest sedi-
mentary rocks are about 3,000 million years old, and the oldest fossil-
bearing rocks are about 500 million years old.
Population Density
It is not necessary to discuss here the details of the various theories
regarding the course which the evolution of insects has follo:.ved; but in
the present context it- WQuid be well to consider how the insects in their
I ~
evolutionary march changed their status.

The first basic point which needs to be properly appreciated is that
it is mostly the insect's capacity to increase its number rather than the
nature of the damage caused by it that makeS' all the difference between
a major and a minor pest. The loss caused is a function of insect popu-
·lation density, and any insect species that causes some kind of damage
to the crop or crop produce can become a serious pest once its popula-
tion reaches a critical level. This critical level can be much lower in the
case of insect vectors of virus diseases in which a very small population
of infected individuals caU spread the infection to the whole crop than in
the case of a simple leaf eater in which the amount of leaf eaten is a
nearer index of the loss caused.
The second point is that the losses caused vary in time and space from
o to 100 %; it is, therefore, the frequency of the insect incidence both in
time and in space that determines the seriousness of the pest. For example,
an insect pest that persists year after year is more serious than one that
attains serious proportions at long intervals. Similarly, a serious pest con-
fined to a few scattered fields is of less significance from the national
point of view than the one which is more widespread.
The third point is that the loss may be either quantitative as in the
case of reduced yield, or qualitative as in the case of apples with spoilt
look owing to even slight infestation by scale insects. The wheat bug,
Eurygaster integriceps, is known to affect adversely the baking quality of
wheat.
The fourth point is that attempts to express insect losses in terms of
money are at times objected to. Besides the invariably fluctuating nature
of the price factor due to various other causes, the objection is sometimes
raised that the selling price of the commodity would be reduced if insect
infestations were controlled and greater quantities of the produce came
to the market; this aspect should also be taken into consideration if the
loss is to be computed in terms of money.
Despite all the above considerations in deciding whether a particular

species is a serious. moderate. or minor pest. the one point which clearly
stands out is that it is the insect's capacity to increase in number which
is most important for deciding harmfulness and economic status. Hence.
in order to understand properly the origin of insect pests. it is necessary
first to understand the forces which determine insect populations; and for
this purpOSe we have to study carefully the interaction between 2 opposite
forces, viz. the forces of reproduction and the forces of annihilation. The
techni.cal 'term for the forces of reproduction is 'reproductive or biotic
potential'; for the forces of annihilation it is 'environmental resistance'.
Balance in Nature
Biotic potential. Biotic potential denotes the highest potential' ?r
capacity of an insect to reproduce itself under ideally optimum condi-
tions. This potential is generally high and constitutes the main motive '
force of population growth. It tends to raise the population level indefi-
nitely.
Prodigiousness of reproduction. Let us take the case of an insect
laying only 200 eggs and completing its life cycle in 1 month, and con-
sider the possible number of its descendants, if all attain maturity.
Commencing, for the sake of convenience, on 1 January, we have 1 fer-
tilized female which lays 200 eggs. All eggs hatch and mature by the end
of the month. On an average, half of these will be females, each of which
will lay 200 eggs on 1 February, so that by the end of February we shall
have 100 x 200 = 20,000 mature insects. Of these, half again will be
females laying 10,000 x 200 = 2,000,000 eggs on I March. It would be
tedious to follow this increase month by month, but simple calculation
will show that by the end of the year the number of descendants of a
small female insect will reach the prodigious total of 2 trillion.
The human mind is quite incapable of grasping the significance of
such figures. but a few calculations may assist imagination. If 1,000 in-
sects weigh only 1 oz, the total weight of 2 trillion will be about 6'00
billion tons. Again. if 1,000 insects are packed in 1 cubic inch boxes, they
would cover an area of 50,000 million square miles with a uniform layer
1 inch deep. or the dry surface of the earth being 51 million square miles
they would cover the whole earth to a depth of 81 ft.
The hypothetical case considered above is by no means extraordinary
as regards either the number of eggs deposited or the shortness of the life
cycle. A single female of cotton bollworm (Earias labia) lays on an ave-
rage 235 eggs at 25°C, and 12 generations are possible in 1 year. Thus. if
conditions remain optimum, the descendants of a single female can cover
th~ entire earth up to a height of about ] 00 ft in 1 year.
Let us also see some examples of reproductive potential in other

animals. A shad is said to lay from 30,000 to 100,,000 eggs, and a carp
from 2 to 4 million eggs in the season. The tapeworm Taenia will have
at least 8,800 eggs in 1 proglottid and may daily discharge 13 or 14 prog-
lottids. The phenomenon of polyembryony found in certain parasitic in-
sects increases the reproductive potential in a peculiar way. The female
lays eggs within the host, and each egg so securely laid produces more
than 1 individual. Many of the social insects have made reproduction a
social affair rather than an individual affair. Thus, the prodigiousness of
reproduction is a common phenomenon, and all theoretical calculations
show that almost anyone of the thousands of species of insects should
be able to multiply its number within a short time to limit~ yyrging on
absurdity, and ,that the earth surface is too small for the progeny of any
single pair. Fortunately, however, nothing of this kind happens in nature;
and going back to Qur imaginary insect we find, as a rule, only 2 descen-
dants, including 1 female capable of laying 200 eggs which will hatch
and complete the cycle in I month. This enormous difference between
potential and actual rates of increase is the result of prodigiousness of
destruction. This leads to the Idea of Balance in Nature, i,e., balance bet-
ween the forces of reproduction and those of annihilation.
Prodigiousness of destruction. A simple calculation will show that
destruction must be enormous to keep the population at the average level.
In the case of an insect laying 100 eggs, as many as 98% of the eggs
must be regularly eliminated to prevent the numbers from increasing, i.e.,
only 2 individuals of each pair (1 male and 1 female) should reach matu-
rity. In more prolific species the percentage of survival is found to be even
smaller. In order to arrive at a, generalized formula, let us suppose :
Number of eggs laid per female (~) = a
Total number of eggs of females ( ~ )
producing eggs = b
Number of generations in a year - c
aC
Now 1 female (~) at the end of 1 generation will produce b if all
eggs reach maturity. But in order to keep the population at the average
C C
\ a ( a -1 )
level all except 1 should die, i.e. of be females bC should die
aC
b C -1
the proportion of destruction
1 be
1-
be be
This proportion of destruction is called 'co-efficient of destruction',
which represents the fraction of progeny that must normally be eliminated
in order to keep the population at the same level.
Thus, in case of E. labia laying 235 eggs and completing under optimal
conditions 12 generations a year the co-efficient of destruction should be:
total number,
(supposin~ : = 50: 50, i.e. or b = 2)
number of
qc (1 - 212 ) 100
235 12
212 X 100
100 -
23512
- 100-0.000,000,000,000.000,000,000,01441
So, % destruction each year = 99, 999, 999, 999,999. 999, 999, 999,
98559.
Destruction % during each generation :
2
q = (1 - - - ) 100
235
(235-2) 235 x 100
100 = 99.14%
235 235
I.e. 99.14% of each generation should die.
Students of evolution say that this huge destruction is the result of
struggle for existence, whereas ecologists say that it is caused by environ-
mental resistance. The two expressions are not exclusive of each other.
The environment gives rise to a force of annihilation, which leads to the .
struggle for existence.
Anyway the; practical point which has to be realized in this connection
is that when such a huge destruction is necessary to keep the popUlation
near the average level one must be very careful not to underestimate the
value of control the nature exerts. In a species of which the female lays
200 eggs, 99% destruction is needed in each generation if the population
is to be maintained at a constant level, and if nature relaxes even so
little' as to reduce the destruction by just 1 %, i.e., the destruction becomes
98% instead of 99%, then the population will get doubled after each
generation. In other words, even a slight relaxation in environmental re-
sistance can lead to an epidemic. Surprisingly, however, this ·fact is easily
overlooked. Some experimenters could rear only 8 % of the E. fabia eggs,

i.e., about 18 moths out of about 235 eggs, even by providing optimum
conditions. They wrote that they had not been able to reproduce natural
conditions in the laboratory because there could not be such a high mor-
tality in nature. The fact is just the opposite. The production of 18 moths
from 1 pair at the end of 1 generation completed in about 3 weeks means
that in 21 days the population should increase 9 times. Such increase
occurs only rarely if ever. Thus, it was undue diffidence to feel that nature
provides better environment than the laboratory.
It seems to be a general law in natur« that if the reproductive potential
is high the survival potential is low, and vice versa. Many species pro-
duce relatively few young but exercise parental care to see that'the few
produced actually survive. On the other hand, many species produce enor-
mous numbers of young and leave them to take their own chance in
nature; in them the survival potential is low. Nature has seen to it that
organisms which live in environments that offer great resistance possess
a high biotic potential and those which live in relatively constant and
secure environment produce very few young at each reproduction. The
history of economic biology gives examples of numerous species which
have been moved accidentally from one environment to another and in
the new environment the population level has been entirely different from
that in the natural environment because of the difference in the resistance
of the 2 environments. It also contains records of many 'biotic explosions'
in nature when there has been a sudden change in environmental resistance,
the biotic potential has been more fully expressed, and populations have
shot up to enormous proportions. Distribution of different species shows
that tropical species go north until the resistance of the environment over-
comes their biotic potential. Similarly, species from low lands move up
the mountains until their biotic potential is overcome by environmental
resistance.
Biotic formula. Some scientists have expressed the above idea by
the formula:
p B,
in which 'P' represents the population level; 'B' repre-
R
sents the biotic potential, i.e., the maximum capacity of reproduction
under optimum conditions when theoretically speaking there is no envi-
ronmental resistance; 'R' represents the environmental resistance, which
is a highly complex and composite factor consisting of all agencies which
bring about the death of individuals before they are able to complete
their reproductive phase .
.Evidently, the above equation is based on the analogy of the equa-
46 AGRJCULTUR~L ENTOMOLOGY AND PEST CONTROL
tion in the field of electricity: C =:!_ , in which C is the electric

R
current, V the electric potential, and R the electric resistance. A little
reflection will show that the analogy is useful though not exact. Fig. 4.1
illustrates a biotic circuit analogous to an electric circuit in diagrammatic
form. Obviously, the analogue of the electric current in our case is what
we may call biotic (vital) current 'P' measured in terms of population den-
sity. The electric potential and the biotic potential depend on the inherent
constitution of the electric cell and the animal species, respectively. The
BIOT~C CIRCUIT
Fig. 4. L A diagrammatic sketch illustrating the decrease in the number of

progeny of a single female from egg (E) to adult stage (A) in a static population.
L 1 - L 5 = larval instar, P = pupa.
resistance factors in both cases are complex combinations of a number

of components.
Let us take another analogy from the passage of heat through a con-
ductor. A conductor reaches a steady state when the amount of heat com-
ing in at any point becomes equal to that leaving the same point .at any
moment, without affecting the temperature at that point. Similarly, in a
steady population the number of births, i.e., the number of vital units
entering the environment has to be equal to the number of deaths, i.e.,
the number of vital units leaving the environment in a unit time. Yet
another analogy can be taken from the flow of a river where the level at
any point is determined by the amount of-water reaching and th~t leaving
it at any moment.
Analysis of biotic potential and environmental resistance. The biotic
potential is divisible into (i) reproductive potential and (ii) survival poten-
tial; the latter is further divisible into (a) nutritive potential, and (b) pro-
tective potential. ~utritive potential is the ability of the organism to uti-
lize environmental materials for the support of Its own metabolism. Pro-
tective potential is the ability t9 protect oneself against e~vironmental
forces. Social insects have made protection also a social fun~tion.
Further, it has been convenient to divide environmental resistance into.
(i) climatic resistance, due to any sort of adverse climatic fluctuation, and
(ii) biotic resistance, due to parasites, predators and diseases. Of late,
these 2 categories have found strong supporters forming 2 opposite schools
of thought known as the 'biotic school' and the 'climatic school', each
unnecessarily considering its own viewpoint most important and exclusive
of the other. For applied aspects, a thorough knowledge of biotic resistance
is required for what is called biological control, whereas an equally good
knowledge of climatic resistance is needed for forecasting epidemics. Fur-
ther, it is interesting to find that generally those interested in higher warm-
blooded animals are strong supporters of the biotic school of thought,
whereas those interested in smaller cold-blooded animals like insects are
strongly in favour of the climatic school of thought. What the biotic school
does not seem to realize is that higher animals are generally long-lived,
and the effects of seasonal changes in weather etc. get averaged so that
the climatic effect is masked. On the other hand, insects are so short-lived
that different generations live, so to' speak, in different climates. The
winter generation faces an absolutely different climate from the summer
generation; hence, climatic factors foreshadow biotic factors.
There are several other systems of dividing the environmental resis-
tance. One system divides it into 3 factors. viz. food. safety. and sex. Food
is resolved into presence of food (host in the case of parasite) and its
approachability (mobility). Safety is resolved into climate (humidity. tem-
perature. light) and enemies (parasites, predators. etc.). Sex is resolved

into relative abundance of males and females. Another system divides the
environment into (i) media - water. soil, air; (ii) factors - light. heat,
humidity. etc. (both biotic and climatic); and (iii) controls - topographical
latitude, longitude. altitude, barriers. etc. Yet another system classifies the
environmental factors into (i) density-dependent factors (sex mating. food
finding, etc.) and (ii) density-independent factors (climate). Of course, it is
difficult to accept this classification on strict basis; for density does change
the micro-climate. i.e .• both temperature and humidity.
Origin of Insect Pests with Origin of Agriculture

Having studied the interaction between biotic potential and environ-.
mental resistance. it is now easy to visualize the origin of insect pests. It
is obvious that any change in environment that brings about a relaxation
in the intensity of environmental resistance will change the balance in
favour of the population of the species concerned. One can visualize that
before the origin of agriculture all types of plants were growing in a
mixed population depending upon the survival value of different plant
species. In such a state of wild growth different species of plant-feeding
insects had to search out their own food plant from time to time. For
example. the moth of the paddy stem-borer, which lays eggs only on
paddy plant on which alone its larvae can feed, had to search a paddy
plant among the wild growth before it could ensure the continuance of
its progeny, and in this search it had to face the dangers of annihilation
by various climatic and biotic agencies. This state of affairs materially
changed when man came into the picture and selected economic plant
species like paddy, wheat, sugarcane, beans. etc., for organized cultivation.
The origin of agriculture took place with this decision of man to main-
tain monoculture of individual economic species of plants. Thus, when
man ploughed down the wild growth and began to maintain in its place
a monoculture of a particular species of plant,_say paddy, all the insect.
species which could not feed on paddy had to leave that area. At the
same time, the conditions became extremely favourable for those species
of insects that could feed on the paddy plant. The monoculture of paddy
meant that the insects that fed on it were saved from the dangers in-
volved in flying about in search of paddy plant in a wild vegetation. This
meant considerable relaxation in environmental resistance to these species
of insects, the population of which could rise much more than was possible
in wild vegetation. The paddy plants, which during this process of mono-
culture were saved from competition with other plant species, began to
face a much more serious threat from the insect species than before. The
insect species which formerly were only minor pests of paddy became
ORIGIN OF I!,!SECT PESTS 49
more serious pests, and in this way the origin of insect pests of paddy
took place.
Not only did the insect pests originate with agriculture as explained
above, but the intensity of pests continued to increase with the intensifi-
cation of agriculture. In the early days of agriculture the fields used to be
small and different crops were grown in different fields. This was so in
India till quite rec!!ntly and one could see a field of wheat surrounded
by fields of gram, or gram fields surrounded by those of mustard. and
so on. In such situations the pests of wheat would have to travrl quite
a distance to get to another field of wheat, and those of gram similarly
would have to fly quite a distance to get to another field of gram; and
in this travel the insects had to face dangers from various sources. Later
on. as larger farm~ appeared and we began to have continuous' fields of
the same crop, environmental resistance relaxed and pest intensity in-
creased .. Continuity of the same crop for miles together meant that the
insects obtained unlimited supply of food near at hand. So, a change
from discontinuous cropping to continuous cropping- in space, inevitably
resulted in increase of pest intensity.
With further intensification of agriculture, there has been continuous
cropping not only in space but also in time. For example, in some part~
of India there are as many as 3 crops of paddy every year. This continuity
of cropping in time is very favourable for increase in pest intensity. In
areas where there is only one crop of paddy the pest population ordi-
narily becomes almost annihilated during the off-season. On the other
hand. in areas where there is repeated cropping during the year this
annihilatory process with the harvesting of the crop does not take place.
There is a third way in which intensification of agriculture leads to
intensification of pest infestation. It is well known that when a crop is
heavily manured. particularly with nitrogenous fertilizers, it grows more
luxuriant and succulent. Luxuriant growth generally makes the micro-
climate more favourable for the pests. Succulent plants, in general, are
easier for insects to feed on.
Thus. from every angle it is clear that intensity of pest infestation is
a natural corollary of intensification of agriculture. Hence. it is no wonder
that the pest problems have been increasing in complexity and serious-
ness. In future, too, the pest problems will continue to increase in their
seriousness with further intensification of agriculture. The more we try
to grow the crop. the more we have to increase the efforts to protect it.
In this context it will be useful to understand and appreciate the concept
of the accentuation of pest problems with the intensification of agricul-
ture (Fig. 4.2).
(i) This diagram depicts aU production efforts in agriculture being
50 AGRICULTURAL ENTOMOLOGY AND PESl CONTROL
WEAKER SPOTS REP'RE~iEN·r-l

ING POTENTIAL
WHICH WILL
SERIOllS WITH
CATION OF
....
Z
'"
II>
'"a:
Q.
INSECT FRIENDS OF MAN

Fig. 4.2. Diagrammatic representation of the accentuation of pest problems with
intensification of agriculture.
aimed at filling up a cistern.

(ii) The size of the cistern is related to the yield potential of the
crop concerned.
(iii) The cistern has a number of inlet pipes representing the various
inputs. such as (a) irrigation, (b) manuring, (c) improved varieties, (d) im-
proved agricultural practices.
(iv) The cistern also has a number of leaks and weak spots in its
walls.
(v) As the level of the produce increases inside the cistern, it begins
to leak out through fresh holes.
,(vi) As the pressure inside increases, the existing holes become bigger
and there is greater leakage through them.
(vii) In other words, as we are taking to more and more intensive
agriculture, the pest problems are increasing both in number and in in-
tensity.
OE,IGIN OF' iNSECT PESTS 51
(viii) The land area being limited, there is no way but to intensify
agriculture to feed the increasing population.
(ix) Hence, we have to be prepared to face more and more serious
problems of plant protection.
To sum up it may be stated that the origin of insects almost coincided
with the origin of land plants, that the origin of insect pest in the hoary
past also coincided with the origin of agriculture, that ever since then the
problem has been getting accentuated with the intensification and sophi-
stication of agriculture, that we should take this phenomenon as the law
of nature, and that we should make adequate provision for meeting the
challenge.
The policy implication or what has been discussed above is ,as follows.
Pest control. / It is not the production alone but the production pro-
tection axis that constitutes the backbo~e of the nation's efforts to
ensure proper food supply for its teeming milliqns. The land is limited,
and the population is rising. Hence, in ultimate a1}alysis increasingly in-
tensive cultivation alone can provide a rational solution of ~)Ur national
problem in the field of food self-sufficiency. But the inevitable corollary
of intensive cultivation is a corresponding accentuation of the intensity
of pest infestation.
Fig. 4.2 provides a good illustration of the relationship between
productive and destructive forces in agriculture. The job of production
specialist is proper management of the inlet pipes, while that of protection
specialists is to keep the exit holes as narrow and few as possible. The
exit holes in the cistern of Indian agriculture are not only numerous but
also distributed at different heights in the wall of the cistern. The result
is that, as better management of inlet pipes by production specialists tries
to increase the level of agricultural production, leakage automatically
increases. Also, as if with greater pressure inside the cistern, the size of
individual holes increases and weaker spots in the wall give way resulting
in fresh points of leakage. This analogy is not merely conjectural; it has
a sound basis in the scientific analysis of the whole ecology of agriculture,
particularly the analysis of the causes of ,origin of insect pests as discussed
in the foregoing paragraphs. The more one wants to produce, the more
one has to protect. This simple fact has not been adequately appreciated
in the past. The result is that the provision for increasing the know-how
about food protection has been much lower than that for increasing the
know-how about food production. This imbalance has to be rectified if
the country is to enjoy the fruits of its labour and expense on production.
~.~~T-;,a~~o·~
1". .
retE
i
/ q;_....
lI"lR'~
ACCf,ssion
,.
tJo..;,.\.3 ~
.
CHAPTER S
ESTIMATION OF INSECT POPULATION
SINCE the damage caused by insect pests is generally a function of . the

population density of the species, estimation of insect populations is the
most important aspect of economic entomology. The techniques of popu-
lation study, however, are far from satisfactory (Konakov, 1939). Criticisms
have been made that population studies tend to stop with the mere' accu-
mulation of masses of undigested data, and that often they are not suffi-
ciently backed or followed by laboratory experimentation. If these Crlti-
cisms are taken care of, the advantage will be that once an investigatof'
has overcome the initial difficulties of dealing quantitatively with insect
population and of interpreting the results in relation to the environment,
there will be no fear of his ever losing completely the broader viewpoint
so necessary for an eCQnomic entomologist.
Three excellent reviews on insect popUlations have appeared in the
past few years (Morris, 1960; Richards, 1961; Strickland, 1961). This is
an index of the importance this subject has assumed.
For a correct and scientific understanding of the population problems
it is of fundamental importance to develop sound methods for estimation
of populations. Two basic considerations are (i) the stage (egg, larva, pupa,
or imago) at which the counting and estimating is to be done most ad-
vantageously, and (ii) the actual process of counting and estimating. The
ideal method would be to count all the individuals, but since this is often
impracticable, the process of sampling has to be resorted to. For a satis-
factory and successful sampling one has to be clear in mind about (i) the
nature of the sample, (ii) the mode of sampling, (iii) the size and number
of samples, and (iv) population estimation by computation and analysis of
the sample counts.
Stage to be Counted
Choice of the stage to be counted has been mostly on convenience
rather than scientific desirability. Most population records are based on
counts of either the most prominent or the most injurious stage, be it is only
1 stage or a mixure of several stages available during the examination.
As a scientific principle, conscious exclusive counting of only 1 stage (while
ignoring other stages equally or more prominent) has been perfected pro-
bably to the fullest satisfaction only in the honey bee (Bodenheimer, 1937)
in which estimates at other stages are based on those of closed brood
cells; this is possible on the assumptions that after the eggs hatch there
ESTIMATION OF INSECT POPULATION 53
is no appreciable mortality in the brood and that all the instars are com-
pleted within specified periods - egg, 31 days; larva, 6 days: sealed brood.
12 days; nurse bee, 10 days; house bee, 10 days; and field bee, -72 days
in the equitable hydro thermic conditions in the beehive.
Population records based on different stages are not comparable so as
to give any idea of the relative efficiency of any stage. Theoretically, th~
egg sta~e is considered ideal for population estimation because it is in this
stage that the highest potential population of the coming generations is
represented and it is possible to count eggs without killing them. The' dis-
advantages are the practical difficulties arising from the generally minute
size of the eggs, and their being laid in the case of some insects in. con-
fused clusters. In the/stages aft~r hatching the advantage of gradual in-
crease in size is nullified by the corresponding increase in movement" which
ultimately makes it necessary to kill them before counting and thus affects
the density of population during each examination - a great disadvantage
in certain types of population studies. In such circumstances it appears
that in selecting the stage or stages to oe counted we have to strike ,a mean
for each species for each purpose in view. -but the point to be kept in
mind is that the population of every species remains continually changing
from the highest potential at the egg stage to the lowest potential of
gravid females (Fig. 5.1). Therefore, estimation, and comparison in
subsequent generations, of population levels at a single point in the biotic
circuit can give only a total of the effects of factors operating on all stages
A w
C B a:
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:g~ 3·f, ii .e...... ..:f, •.•~. I . 3 ~
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w£U~S~~~1~2-3~4~5~6-7--8~9~lO-1~1-'2~13-'~4-'~5-'~6-'7~1~8-'-9-2~O~2~'~22-2~3-2~4-25~26-2~7~ '"
JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER JANUARY
Fig. 5. L Weekly fluctuation in population of the spotted bollworm of cotton at

Delhi during 1941-42 (After Pruthi and Pradhan, 1945).
of the generation, and often this total may be too confusing to allow
analysis. Consequently, it is well to estimate the population level at
many suitable points in the circuit as practicable or desirable for a parti-
cular purpose. In the case of the fruit tree leaf roller Paradis and Leroux
(1962) assessed the population at 4 points (i) egg stage, (ii) May larvae
(ins tars I and 11), (iii) June larvae (instars III and IV), and (iv) pupae.
Morris (1959) did it in the larval stage in each generation.
Nature of Sample
The nature of the sample depends on the insect and the stage to be
sampled. The nature of samples used by various workers is described b~low
under different categories. \
1. Net sweepings. There are many complexities in the estimation of
insect population by the sweeping method. Evening catches were found \
to be 300 to 800% greater than those made during midday (York and
Prescott, 1951). Poisoned insects do not come in the sweep (Hoerner and
Longford, 1925). The catch varies with the density and height of the grasses
and also with species (Rubtzov, 1932). The number of sweeps necessary for
a fair degree of accuracy is generally too large for the method to be of
much practical value (Gray and Treloar, 1933). Khanna et ai. (1957) con-
cluded that this method provided no idea of any of the indices used in
judging intensity of infestation. The shape of the net, however, does not
seem to be of much significance, and individual net counts are considered
better than continued sweepings (Beal, 1935). Ail the same, because it is
easy, the method is recommended by some (Mohammed Afzal et al., 1944).
2. Sudden trapping. For such minute insects as Collembola in pas-
tures, Davidson and Swan (1933) tried the method of sudden trapping.
They took a cylinder of fixed cross-section and with its bottom be levelled
to form a cutting edge, put it quickly on the surface of the pasture with-
out causing any disturbance, and thus trapped all the insects present in
the area enclosed. Then they cut out the earth below the cylinder to a
depth of a few centimetres and raised the whole thing covered 'from below'.
They separated the insect fauna from the earth by floating or washing.
Hills (1933) suddenly trapped insects as above and then collected them
by means of a vacuum apparatus. Smith and Stewart (1946) used a cage
for sudden trapping and sampling of grasshoppers.
3. Screen trap. Gaines (1932), working on migration and population
sh,Idies of the cotton bollworm (Heliothis obsoleta) devised a rather useful
cart-type screen trap which was pushed along over cotton rows in the
field. The moths thus disturbed flew upwards and were caught in the trap.
This trap proved quite successful in obtaining information on the extent
of local migrations and the abundance of bollworm moths in the field.
ESTIMATION OF· INSECT POPULATION 55
4. Narcotized collections. Delong (1931) and Collyer (1951) .des-

scribed what they called a successful method for sampUng popUlation of
quick-moving insects; they dusted the plant with an insecticide having
quick knockdown effect, and collected the insects falling on a sheet below
the plant. Gray and Schuh (1941) used a sampling can consisting of a
funnel covered with a coarse screen mounted below a gas chamber con-
taining methyl-iso-butyl ketone, the fumes of which made the aphids to
drop. The can was then given 30 shakes to make the aphids fall through
the screen into the carton below the funnel. Dempster (1961) used a
sampler consisting of a box in which the insects were anaesthetized for
estimating population of active insects on vegetation. .'
5. Light-trap. Light-trap catch has been so generally used to indi-
cate the time of ~mergence, of positively heliotactic insects, especially
moths, after periods of hibernation or aestivation, that its use as a
sampler for quantitative, estimation of simultaneous seasonal abundance
of several species began to attract attention; but Cook (1928) and Williams
(1935) gave verdicts against this kind of sampling'. Cook's findings indi-
cated that light-trap catch is not reliable for any estimate of comparative
abundance of different species, although it is fairly reliable:' for inter-
annual fluctuations, especially in stationary traps for long periods. Williams
found that different species came at different parts of the night wjth the
greatest influx in the earlier part. These studies are sufficient to indicate
that the light-trap catch for the same species taken at 2 different times
can be compared only under exceptional conditions. Evidently, long-period
stationary traps may be taken to indicate inter-annual fluctuations because
the average of environmental conditions, especially weather conditions.
may tend to be averaged out in several years. Nevertheless, light-trap catch
has been used by many to indicate comparative abundance of insects.
Bogush (1951) and Broadbent (1948) carried out an analysis of light-trap
catches of aphids. Williams (1940) published a comprehensive analysis of
4-year captures of insects in light-traps. Nagel and Granovsky (1947) used
a turntable light-trap for making insect collections at regulated periods.
and Glick and Hollingsworth (1953) studied the response of pink boll-
worm to certain ultraviolet and visible radiations; they caught 50,000
insects with mercury-vapour light-trap, and 112,000 with ultraviolet trap.
6. Water trap. Some workers used 'a floatation pan trap for insects
associated with water surface (Grigarick, 1959). An aluminium pan, 20 em
in diameter and 3 em deep, was set in a-wooden float. The pan contained
water with a little wetting agent to lower surface tension and cause the
alighting insect to sink. A correlation has been found by some (Harper
and Story, 1962) between the number of adults caught in water traps and
the number found in emergence cages in the dipterous root maggot pest
56 AGRICULTURAL ENTOMOLOGy AND PEST CONTROL
of sugar-beet. Experiments have also been carried out with water traps
of different colours (Lewis, 1959).
7. Suction' trap. A number of workers have developed various kinds
of suction traps in which the suction apparatus is operated either by hand
or by means of an electric motor as in powerful electric suction pumps
of standard pattern (Johnson, 1950; Ristich and Lockwood, 1953; J ohn-
son et al., 1957;- Dietrick et al., 1960; Dietrick, 1962; Southwood and
Pleasance, 19,62).
8. Adhesive trap. Adhesive traps consist essentially of some suit-
able persistent adhesive material coated on a suitable surface so that in-
sects that come in touch with the surface get stuck to it. Some workers,
have used such simple types as strands consisting of midribs of palm
leaves coated with adhesives and hung round the stacks (Reiley, 1957);
others have used the adhesive surface fitted into a wind-vane type appa-
ratus (Staples and Allington, 1959), although the efficiency did not im-
prove into the funnel in front of the trapping surface. Another improve-
ment has been the use of a transparent plastic sheet adhesive wrapped
round a suitable circular surface; this sheet can be removed and examined
at suitable intervals (Broadbent, 1948). In the case of Aphis migration
the colour of the trap was found to make a difference in the catch; brilliant
yellow was more attractive than white, and white more than black.
'9. Bait trap. Bait-trap catch has been used for estimation purposes,
although the samples have more serious complexities than those with the
light-trap. Legner and Davis (1962) used a simple trap for the European
earwig using wheat flakes as attractant. Beckham and Dupree (1952) used
bait-trap for the green June beetle. Saunders and Krueger (1957) based
their technique for counting larvae of the confused flour beetle on the
ability of the larvae to cling to rough paper. A number of workers have
used a very peculiar bait-trap in which a female of the same species kept
in a cage was used to attract the males by its sex scent (Ambros, 1937;
Farsky, 1938; Komarek and Pfeiffer, 1938; Maksimovic, 1960). Davidson
and Andrewartha (1948) studied annual trends in natural population of
Thrips imaginis on the basis that roses were highly attractive to adults
and gave a satisfactory index of the levels of the population of this pest
in the area.
10. Sight counting. Sight counts from measured areas have been
used in estimates of grasshoppers and locusts. Lockwood (1924) used field
glasses for counting grasshoppers from a distance; and Shotwell (1935)
described a method for making grasshopper survey in which the counter's
sight was first trained to estimate without measurement an area 1 sq yd
and then to count the hoppers in that area. The limitations of such
methods are obvious. VinQkurQV (1938) used a specially designed frame
to disturb and count grasshoppers from a measured area; and Smith and
Townsend (1952) used an apparatus which restricted the area of the
visible crop to 1 sq yd in which the insect could be directly counted.
Richards (1953) and Scheepers and Gunn (1958) used the technique of
counting the red locust jumping up at every 100 paces and for 2-yd strip,
resp'ectively. Hartzell's (1946) technique of counting by means of a net
micrometer disc attached to a low-power microscope is also in this
category. Carpenter (1935) studied fluctuations in biotic communities of
prairie forest of central Illinois using as the unit the number of birds
and mammals that could be seen by cruising through the fort;st edge
over a route about 1 mile long.
11. Fixed volume or area of earth. For such soil insects as beetle
grubs, earth dug put from 'small fixed areas to fixed depths has been
successfully used as sample. Fleming and Baker (1936) used it for eluci-
dating the general principles of determining the size and number of sam-
ples and also the mode of sampling. Many work~rs have tried to make
improvemen_ts both for taking out samples (Wallace, 1956) and, for sifting
the insects from the sample by washing (Bennett and Ke~rns, 1943;
D'Aguilar et al., 1957; Pickles, 1946) or floatation (Daniels, 1933; Ladell,
1936; Murachev, 1938; Cockbill et aI., 1945; Henderson, 1960). Barnes
(1937) estimated the population of crane fly Tipula paludosa, by obtaining
the larvae from a fixed area of earth not by digging but by watering it
with an emulsion of orthodichlorobenzene; the larvae came to the surface
withlll a few minutes without serious injury. Milne et al. (1958) used heat
for forcing tipulid larvae and pupae out 'of the soil. Miller and Martyn
(1952) developed a sampling technique for underground grass grubs in
which they removed the turf and soil to a depth of about 1 inch from
I sq ft sampling area; they loosely plugged the openings of the tunnels
made by the grass grubs, and after 24 hours counted the holes made by
the live larvae.
12. Crop samples. For estimating the population of most of the
internal crop feeders, the affected crop itself is sampled and the number
of insects found in each sample is taken as the sample count. Innumerable
examples of this kind of sampling can, be quoted from literature. Most
of the important pests, such as the European corn borer, rice borer
(Tsai and Tong, 1937), codling moth (Barrett, 1933), blackheaded bud-
worm (Silver, 1959), various species of American and Indian cotton boll-
worms, sugarcane borers, gall midges, and even aphids, thrips and mites,
which are actually not internal feeders but remain only entangled within
foliage and flower petals have been studied with this kind of sample.
Improvements in techniques involving crop samples have been made for
taking samples from suitable parts of the plant and for detecting insects in
58 ,'AGRICULTURAL ENTOMOLOGY AND PEST CONTROL
the samples and extracting them from it. Thus. for example. Madsen et al.
(1961) found that sampling from terminal. central and basal portion of an
apple shoot gave a better indication of aphid numbers than a sample res-
tricted to the terminal leaves. Summers and Baker (1952) found that almond
mites feed on leaves but rest on barks; hence only a small portion of
the mite population remains on the leaf at anyone moment. necessitating
that the foliage should be cut along with the woody portion. Wilson (1962)
developed a portable device for sampling foliage-inhabiting insects; it
entailed such crude operations as beating the sampled branch with a pole
to make the insects fall into a specially designed cloth bag; but others
developed methods for quite delicate handling. Thus. Henderson and
McBurnic (1943) and Morgan et al. (1955) designed special machines for
carefully brushing mites and their eggs from leaves and fruits; Shrick
(1948) used a modified Berlese funnel for collecting and counting onion
thrips; Nielson and Bleak (1961) developed an insect-separator-collection
box'; Newel (1947). Hartzell and Horsfall (1944) and Morgan et al. (1955)
developed techniques for removing mites and their eggs by soaking and
shaking the leaves in various chemicals; and Le Pelly (1942) removed
thrips by dipping the leaves in 70% alcohol and filtering the alcohol
through filter paper treated with blueprint solution so that the thrips could
be easily seen against the dark background. Prasad (1953) and Pielon
(1961) determined aphid numbers by volumetric methods. A number of
workers (Venables and Dennys. 1941; Summers and Bakers. 1952; Muller,
1959) used the imprint method for estimating the number of thrips; the
sample leaf is gently pressed between 2 sheets of paper and the stains
left on them by individual insects are counted. Baten and Huston (1943)
counted mites on a fixed area of the leaf. A comparison of imprint and
brushing techniques showed the latter to be superior (Chant and Muir.
1955). Milner et al. (1950) used x-ray to detect insect infestation inside
the grain. Flanders (1932) used the heat of infestation as a rough measure
of grain infestation.
13. Emergence cages. Emergence cages with a sampling area of
2 sq ft were used to trap adults of the lac sawfly as they emerged from
the soil (Tumock. 1960). The technique used for the adults of Ephestia
elutella in which muslin bags were fixed over strips of ceIling crevices
about 2 ft long so as to catch the moths emerging therefrom also comes
in this category (Richards and Waloff. 1946).
14. ,Amount of damage. The satisfactory nature of the counts from
crop samples, coupled with the economic importance of the crops, has led
workers to take the amount of damage done as the unit in estimating
population fluctuation of pests. For example. Barrett (1933) worked out
a general method for measuring insect population by using the weight
ESTIMATION. OF INSECT POPULATION 59
of wormy nuts for estimating the population of codling moths; but a

serious consideration in almost every case will reveal the unexpected com-
plexities of this procedure. Ballard (1921). working on pink bollworm in
south India. discussed how the knowledge of the number of boBs attacked
is insufficient to give accurate data. He pointed out that damage is done
,by (i) seed being destroyed. (ii) lint development being retarded and lint
weakened, (iii) premature opening of the boll and invasion by saprophytic
fungi. (iv) staining of lint. and (v) lowering of the germination of un-
attacked seeds in an attacked boll. Hence. the number of attacked bolls
on any given date does not give an accurate'idea of either !he cash value
of the damage or the extent of infestation. Reference may be ~ade to an
excellent pa~er by Gough (1919) on this aspect of pink bollworm damage.
Ossowski (1956) used coloured plates enabling 7 degrees of infestation
to be judged by the appearance of the trees. A somewhat similar match-
ing method Ylas found suitable for rapid estimation of foliage iniury in
potato crop (Granovsky and Peterson. 1954), in paddy crop by Hispa
(Basu and Banerjee. 1957). and in grape by leafhopper (Hartzell. 1946).
A highly significant coefficient of correlation was obtain~d between visual
rating of infestation and average number of borers per maize plant (Sinm
et al., 1962). It has been shown that uric acid content of garin provides
a good index of the damage (Venkatrao et al.. 1957).
15. Colonies of social insects. In the case of social insects the colony
has been invariably taken as the unit. but the problem that has been con-
fronting the workers is how to count the number in each colonv. In the
case of ants and termites it apnears that counting has been done either
in the cold weather when the insects are inactive (Cory and Haviland.
1938) or after fumigation. Fairly ingenious methods have been used in
the case of the honey bee. According to Bodenheimer (I937) the old
method of estimating the ponulation of a beehive was to esflmate the
number of the various sections by senarate methods. The hive was weighed
in the morning before anv bee had gone out. and al!ain in the afternoon
when all bees were out in the field: the return of the bees was indicated
bv an increase in the weight of the hive, which was keot constantly on
the balance. The difference between the maxim 11m (morning) and minimum
(afternoon) weights gave the weight of the field hees_ The numher cOlllrl
he cl1lculated from the weil!ht. The difference in the weil!ht of the hive of
field and house bees and its weij!ht with field bees !!ave the weight of
the house bees.
16. Collection of marked insects. Release a number of marked in-
sects in the field; allow them to mix thoroul!hlv with the insects of the
field to become a homo!!;eneous part of the general pooulation: then make
an unbiased catch of the insects including the marked ones. and calculate
the population by using the formula :

No. of insects marked and released x No. of total catch
Population ='
No. of marked insects recaught
This principle was first applied by Lincoln (1930) for calculating water-
fowl abundance on the basis of banding returns. The same principle was
perhaps independently initiated, developed and improved by Jackson
(1936, 1939) and Lloyd (1936), who tried to make allowance for the decrease
from death, migration, etc. in the number of marked insects during the
period between release and re-catch. Jackson utilized 2 methods which he
called 'positive' and 'negative'. In the positive method he released a number
of marked insects and made catches on several successive days, and then
from the decrease in the proportion of marked insects in the successive
catches he calculated the rate of decrease in marked insects and, thus, the
actual number of marked insects on any day of re-catch. In the negative
method he continued to release marked insects for a number of days using
different colours on different days, and then made a re-catch and calculated
the rate of decrease from the proportions of marked insects of the previous
releases. Lloyd released a number of marked insects at intervals, using a
different colour each time, made a random collection on each release day
after the first, and released back the re-catch after calculating the propor-
tions of those marked on previous occasions. Thus, while essentially using
the negative method of Jac Lloyd, he was able to collect larger. and con-
sequently more reliable, data than Jackson.
It may be pointed out that there is a serious objection to Jackson's
positive method. He does not take into consideration that fluctuation in
the proportion of marked insects in successive catches can be ascribed to
fluctuation in the number not only of marked insects but also of unmark-
ed insects whose population it is desired to know. This defect does not
remain in his negative method or in Lloyd's method because different
proportions of marked insects released on past occasions are calcu-
lated with reference to the same catch. Thus. the difference in the
proportions of the different days' releases recaught on the same day and
referred to the same day's total catch cannot be attributed to any other
cause than the actual difference in the number of marked insects within
the general population. Both Lloyd and Jackson had used the method
with tsetse flies. Several workers have tried this method with other insects,
such as Pyrilla pest of sugarcane (Pradhan, 1945) and orchard population
of the heteropteran Blepharidopterus angulatus (Muir, 1958). Richards
(1953) used the method for the red locust. and Richards and Waloff (1946)
for Ephestia larvae. Melville (1957) used an essentiallv similar principle
in determining the density of mite eggs in flour; he mixed a known quan-
tity of Lycopodium spores in the flour containing mite eggs; there after
he separated the bulk of the flour by chemical methods, which did not
affect either Lycopodium spores or mite eggs; then he examined the sus-
pension of the residue under the microscope, determined the ratio between
the numbers of eggs and spores, and finally calculated the number of eggs.
A somewhat similar principle is discernible in the technique of Bean (1958).
according to which the knowledge of the percentage parasitism of spruce
budworm by techinids and the number of techinid larvae on 1 unit area
of ground beneath the tree can be used for estimating the population of
the spruce budworm. I
17. Human records and nature's preservations. In studyipg the fluc-

tuations in numbers of insect communities through long periods workers
have had to resort to various special kinds of samples. Carpenter (1940)
studied fluctuations in insect numbers in Europe by using as the unit
the number of outbreak records in each year. Despite his effort to give
some weightage to the records according to importance, there remain seri-
ous limitations in his method, All the same, his work has revealed some
interesting indications. Brues (1933), being impressed by the relation beween
the number of various insects becoming entrapped in the'resin flowing out
of trees to their actual number in the forest, studied the progressive change
in insect populations of forests since the early Tertiary by a numerical
study of the) various groups preserved in amber.
18. Comparison of different types of samples. A number of workers
have tried to compare experimentally the relative efficiency of different
types of traps for sampling insect populations (Mohammed Afzal et al.,
1944; Kretzschmar, 1948; Johnson, 1950; Groves, 1955; Taylor and Smith,
1956; Heathcote, 1957; Lewis, 1959; Race, 1960). The relative efficiency
of different types varied with different species of insects studied.
, ..
Mode' of Sampling
The process of sampling is a child of necessity arising out of the prac-
tical difficulty or impossibility of counting the whole population. Evidently.
the larger the percentage of the whole population it is practicable to sample.
the more reliable the result. However, modern statistical researches have
made it possible to reduce considerably the total cost of sampling without
much sacrifice of accuracy. In fact, the requirement of the degree of
accuracy being different for different aims in view, at times it is a waste to
sample more than a certain percentage of the whole population. Sampling
resources being limited, it is always advisable to consult the statistician for
their proper allocation. For a general understanding, however, it should be
easy to appreciate that most decisions regarding sampling techniques have
to depend on the nature of the insect distribution. If the distribution is
ideally uniform (which is rarely the case), even a single sample should
give the information about the whole population. If the distribution is very
heterogeneous. as in the case of social Hymenoptera it may be extremely
difficult to develop a proper sampling procedure. Hence, it is necessary for
the entomologist to study in collaboration with the statistician the nature
of distribution of the insect species and of the stage in which he
is interested. Population density and degree of infestation seem to
have a consider:able effect on uniformity of distribution, and the rela-
tive magnitude of sampling errors varies inversely with the population
mean (Morris, 1955). In one case the level of infestation accounted for
63% of the total variation (Krause and Prederson, 1960). Therefore, infor-
mation regarding the distribution must be available before a suitable
statistical method can be developed (Upholt and CraIg, 1940; Larrimer.
1924). Insect distribution is rarely normal, and it has been found to
approximate to the Boisson series in the case of wireworms (Jones, 1937),
to negative binomial in the case of it "number of species (Waters, 1960;
Nielson, 1957; Geier, 1956; Burrage and Gyrisce, 1954; Connola et aZ.,
1957), and to distribution in the case of European cornborer (Neyman,
1939). As a result of such studies various types of sampling techniques
have been suggested, e.g., double sampling (Wadley, 1949), sequential
sampling in the case of winter moth (Reeks, 1956) and lodgepole needle
miner (Stevens and Stark, 1962; Stark, 1952), and stratified sampling in
a number of cases including spruce budworm (Wilson, 1959; Morris, 1955),
potato aphid (Anscombe, 1948), bean aphid (Banks, 1954), and large
sawfly (Turnock, 1960). In the case of potato aphid the advantage of using
the leaflet or even one-half of it as sub-unit of potato leaf sample has been
reported by Shands et aZ. (1954). In some cases quite exclusive type of
sampling has been suggested; e.g., in the case of pests of cotton sown in
rows it was suggested that a certain number of plants should be examined
in each row along a diagonal from comer to comer of the field (Yakhontov,
1931).
Number and Size of Samples
After deciding the method of sampling and striking a mean between
the minimum percentage that must be counted and the maximum that it
is possible to count, one has to adjust that percentage between suitable
size and number of samples. These matters, especially in the study of
insect populations, are generally decided merely by common sense, and
it is only,in recent years that a number of workers have tried to settle
questions experimentally or at least systematically.
Livermore and Neely (1933) were probably the first to try determining
the number of samples necessary to measure the difference with varying
degrees of precision. They developed the formula:

PEs
PE J\I = PE !If = Probable error of mean
vI-n- PEs = -do- single determination
PEn = -<10- difference
PEA = -do- mean A
PE B = -<10- mean B
vi (PEA )2 .+ n = number of samples
D = difference between means
R = ratio DjPE D
Since probable errors themselves are small quantities. the difference
between them must be still smaller and negligible. Thus. PEA may be
taken as equal to PE B ,
.. PE D = vi 2(PE111)'
••
=
vi 2.PE ...... Now. the degree
of accuracy depends on ratio DjPE D • for which let us put
D D D
R
PED -
vi 2PEy: V 2PEs
vi n
Dv'T (R v' 2PEs
or R =, orn =
D
vi 2PEs
In order to use the formula we have to determine PEs by a prelimi-
nary examination of about 20 samples. and the value of R (ratio D/PE D )
can be taken as desired. Generally. a value of R higher than 3.2 is taken
as significant.
Fleming and Baker (1936) worked out a method in which populations
of larvae of the Japanese beetle in the field were estimated by examining
completely a plot of 2500 sq ft with least units of 1 sq ft. Then these
initial units were combined in various ways and the efficiency of various
sizes. numbers and percentage value of sample was compared by the
criterion of relative standard error. 'It was found that 1 sq ft was the
most suitable size and that larger units increased the standard error. It
was also concluded that 1 % area sampled was sufficient. although it was
quite evident that the standard error increased rapidly as the percentage
of the area sampled became small. It was the size and not the shape of
the unit that affected accuracy. Marshall (1936) independently tried to
solve similar questions in regard to the American bollworm Heliothis
obsoleta. He conducted a precision experiment in which he counted the
bollworm eggs on every plant of a maize plot of 0.125 acre, and then
tried to study the frequency distribution of the eggs in the field by means
of various units formed by combination of the initial I-plant unit. The
units he tried were I-plant, I-yard row length, 3-yard row length, and
finally a composite unit of 4 random I-yard row lengths. He found that
the frequency distribution of none of these units except the last was nor-
mal, and that even this gave a rather skewed curve.
Beal (19.39) tried to evolve a method for estimating the population of
insects in a field and used Colorado potato beetle as the experimental
medium. He got all the beetles of a plot collected and counted, using the
smallest units of 2-ft row length, and then by combination of these initial
units he studied the efficiency of various shapes, sizes and numbers, as
well as direction in the case of longitudinal units. This experiment showed'
that without impairing accuracy a marked reduction in percentage of the
crop sampled could be obtained by stratification and by making the
number of sampling units examined in a particular stratum proportional
to the standard deviation therein. It also indicated that the smallest
sampling units gave the least variability in estimation from a given amount
of sampling. BeaI's experiments, unlike Fleming and Baker's showed that
the shape and even the direction in the case of long-plot units do influence
variability, for there was greater variation between the rows than within
the rows, and consequently long narrow samples running in the direction
of greater changes were more efficient than those running in the direc-
tion of lesser change.
Pradhan and Menon (1945) carried out precision experiments for deve-
loping the sampling techniques for the spotted bollworm of cotton. They
examined the whole of a 0.025-acre cotton crop on the basis of the smallest
sample unit of 1 plant, then with the help of their basic data formed 23
different types of sampling units and analysed these statistically in a
variety of ways. They concluded that (i) the sample units should be
formed on random plant basis, and (ii) the co-efficient of variation de-
creases with increase in the size of the sample units, but this effect is at
times more than nullified by the corresponding decrease in the number
of sample units (keeping the total number of plants examined as constant).
Owing to these 2 opposite effects the relative standard error fluctuates
within rather negligible limits. This should mean that the size of the
sample units is immaterial; but if we look back to the nature of fre-
quency distribution we find that from absolute skewness with the smallest
unit the frequency distribution tends towards normalcy with increase in
the size of sample units. Since the usual statistical tests are based on the
assumption of a normal distribution, and the harm done. if any, by the
larger' units is not much in raising the relative standard error of the mean,
the balance again is in favoUr of larger units. But this increasing of

size must not be continued beyond the limit which is absolutely necessary
for satisfying the conditions of normal distribution, because with increase
in size the number of classes goes on increasing, and the concentration
about the mean goes on decreasing; and, although the co-efficient of varia-
tion remains decreasing with increase in size, the actual range of variation
increases. Thus. for example, with 2-plant random unit there are 9 classes
(0 to 8) with 0 to 8 bollworms unit, but with 10-plant random unit there
are 1 to 19 bollworms unit. So it is concluded that a unit of 5 to 7 ran-
dom plants is a suitable size.
Because 'the analysis of variance shows generally greater variation
between rows than within rows, care should be taken during ,sampling to
include in the observations as many rows as practicable.
Since there is rather significant correlation between the bollworm
numbers and the condition of the plants as indicated by the number of
buds and bolls un 'them, it may be worthwhile to study the population
fluctuation with the help of adjusted population.
These and a few other precision experiments carried out during the
late thirties and early forties were followed by a large number of con-
tributions on similar lines, particularly on spruce bud worm in Canada
(Morris, 1955) and sugarcane pests in India (Acharya et ai., 1958; Rahman
and Singh, 1944; Pradhan and Prasad, 1963), and procedures for deter-
mining optimum allocation of sampling resources are found even in a
number of textbooks (e.g., Cochran, 1953; Snedecor and Cochran, 1956;
Beal, 1954). These procedures have also been used for estimating the
optimum number of samples per tree as well as the optimum number of
trees required to be sampled for population estimation of bud moth
(Le Roux and Reimer, 1959) and fruit tree leafroller (Paradis and Leroax,
1962). Sreenivasan (1956) carried out a sort of precision experiment fOJ
sampling the visible amount of damage caused presumably by jassids on
the leaves of cotton.
Besides the experimental results quoted above, certain considered views
of King (1939) based on enlightened experience should be kept in mind.
According to him, "the unit size sqould be such that the frequency of
both zero and high counts will be minimized for each common form.· An
area of about I sq ft is ideal for many population studies of invertebrates.
A smaller area should not be accepted unless it reduces the time per
sample unit sufficiently to permit a corresponding increase in ·the number
of units without material reduction in the aggregate area. If the size of
sample is cut to one-fourth the number must be at least doubled to compen-
sate for the lesser aggregate area included. Small samples are definitely
worthwhile only when there is a very dense population."
From the foregoing review it seems fairly reasonable to infer the

following for general guidance of the entomological worker.
1. The higher the percentage of the population sampled. the better.
2. The larger the number of samples. the better.
3. The smaller the size of the sample. the better. until the size be-
comes so small that (i) the majority of the samples are ilkely not
to represent the mean characteristics of the population. or (ii) it
begins to exercise a harmful effect on the accuracy of the result by
rendering the examination so tedious that the percentage of the
population sampled has to be materially reduced, or (iii) it intra'
duces to much error.
4. All the 3 quantities, viz.. percentage value. size, and number of
samples. have to be increased with in heterogeneity of the populat-
tion distribution and with decrease in population density of the
insect.
Method of Computing Pest Incidence
The incidence of both pests and parasites is generally recorded as
percentage values. The pest incidence is recorded as percentage of the
material liable to be damaged; for example, in the case of cotton boll-
worm it is common to record the incidence as, say, 50%, meaning that
50% of the bolls and buds examined were attacked by bollworm. Similarly,
parasite incidence is recorded as percentage of the host examined. Some
authors make slight deviations from this general practice. Thus, the inci-
dence of cotton bollworms has been expressed as percentage of the number
of plants instead of buds and bolls. Workers on leafminers and Ieafsuckers
often express incidence as number of insects per leaf (which is the same
thing as percentage incidence with respect to the number of leaves), as
number of insects per unit area of the leaf, and so on.
The chief flaw in all these methods is that they overlook the funda-
mental principle that the scale used for measurement must be constant if
the measurement is to be reliable (Pruthi and Pradhan. 1945). When the
incidence of a pest is recorded as percentage of the material liable to be
damaged, the quantity of the damaged material becomes the scale against
which the pest population is measured, i.e .• the number of buds and bolls
becomes the scale in the case of bollworms, the number of canes in the
case of cane-borers. the number of leaves or area of leaf surface in the
case of leafsuckers like aphids and white flies. Similarly, when we express
the incidence of the parasite against the density of the host, the number
of the host becomes the scale against which the parasite population is
being measured. Now these scales are obviously not constant. In nature,
the quantity of material damaged by pests and that of the parasitizabl~
host material may vary from zero to infinity from season to season and
in the same season. There are also variations due to several other factors.
Thus, the scales with which we measure the incidence of pests and para-
sites are subject to such wide fluctuations that the measurements~ i.e., per-
centage incidence values, cannot be expected to be reliable or comparable
except under very well-defined conditions. Let us take a concrete example
of the bollworm. Suppose at the time of the first observation there are
100,000 buds and bolls and 10,000 bollworms in the field. If we conduct
sampling in the usual way we shall record 10% incidence. If ithe same
field is examined after some time, when the number of buds 'and bolls
has increased to , 200,000 ,but the number of bollworms has re~ained
'
stationary at 10,000, we shall record only 5 % incidence. These observa-

tions will lead us to erroneous ~onclusion simply because the 'scales used
for measuring the incidence has not been constant. That such erroneous
conclusions are actually arrived at in practice will be clear from Fig. 5.1 in
which 3 different kinds of data, vi~., the number of buds and bolls per acre,
number of bollworms per acre, and percentage incidence of bollworms are
graphed together. Starting from E, the number of bollworms rises up to
the point F or at least remains constant; but since the number of buds
and bolls has considerably risen from E to F the percentage incidence
~hows a fall. Again, at the point N, as the number of buds and bolls
rapidly decreases the percentage incidence shows rapid rise though there
is little rise in the actual number of bollworms. These are the discrepan-
cies met with even when we compare the data from week to week;
they become much more glaring when we compare the data from year to
year. There will be some improvements if the incidence of the bollworm
is recorded, say, for every 25 plants because the plant density in the field
is more constant than the number of buds and bolls; but this percentage
incidence will not be comparable if it is recorded in different fields with
different plant densities or in the same field in different years when
germination has been different. Incidence expressed in terms of number
of insects per leaf has the same flaw as the percentage incidence of boll-
worms. The incidence as number of in:~ects per unit area of leaf appears
at first sight to be quite precise, but actually it is subiect to even greater
fluctuations than the number per leaf, because not only the number of
leaves but also the area of the leaf is variable from time to time.
Thus, the incidence of pests and narasites should always be calculated
with reference to some fixed scale. In the case of agricultural pests, and
of parasites of these nests, the unit area of the land on which the croP
is grown provides a fairly good constant scale. The nopulation per acre
can be easily computed by simultaneously estimating, with ordinary
sampling procedure the number of insects per plant as well as the
number of plants per acre. And if the nature of the insect under study
allows, population per acre can be calculated by direct counting of insects
on small units of land area. Where it is considered desirable to record
the percentage incidence, it is advisable to record also the average popu-
lation per acre of the host material so that the population per acre can
be calculated whenever a more precise comparison is called for.
Reference may also be made to some papers of special interest ih
calculating and interpreting the population determinations. Barrett (1933),
working out a general method for measuring insect populations, studied
the effect on population density by comparing not the population density
itself but the percentage' increase over the initial population. Morrison
(1940), working on hope pests, considered the number of red spiders per,
leaf as 'measure of population distribution'. Greenslade and Pearce (1940)
studied field sampling for oomparison of infestations of strawberry crops
by an aphid species and reported that the quantityv n-f enables better
comparison of different samples than the actual number (n) of the aphids
on a leaf. Evidence is given to show that the standard deviation of v' n + ~
rarely exceeds i.3. According to Bliss (1941) it is advantageous to transfer
the number of Japanese beetIe larvae into square roots. Henson and
Stark (1959) suggest that instead of describing insects in absolute numbers
their infestation should be classified as (i) tolerable - populations that
do not utilize the entire 'excess of the biological productivity of the host.
(ii) critical - populations that utilize the excess biological activity of the
host. but less than the total productivity. and (iii) intolerable - popula-
tions that are depleting the host at a rate greater than the current rate
-.
of production.
CumuJative Effect on Insect PopuJation

It is often more scientific and convenient to estimate insect population
on the basis of counts of a single stage, i.e .• egg. larva. pupa. or (rarely)
adult. Since the duration of each stage varies with changes in environ-'
mental factors. it is necessary to understand the effect of these changes
on the census observations in any particular stage. For example. let us
take at case in which the egg stage is to be counted under 4 different
environmental conditions, A. B, C, D (Fi!!. 5.2). under which the in-
cubation period of the eggs is 2. 3, 4 and 5 days. respectively. Further.
suppose as a theoretical possibility that 3 eggs a day are being laid
unoer each of the 4 conditions. At the end of the 1st day (I) when the
eggs have begun to be laid, there will be only 3 eggs (F.E.) in each con-
dition. Qn the 2nd day (IJ), 3 more eggs (F.E.) will be laid in each. and
at the end of the 2nd day (II) there will be 6 eggs each. On the 3rd day
o
o <
-0--- 0
t• __ . .8_1_E-,..4
•
• : FE i
o
o
o
---
o - o
o g 2E
o
o 0 o <
o 0
8 1.E
o 0
-
• - - •
• •• FE
•
o _ _ JL _ _
_lL g ! 2E
0 0 0
0 0 0 o 0 " 1E
-
• -- -
• ---
• - - ......•----1
•• •• •• •• FE
o
t--i.--
r•• •• ••g -- g
•
: FE
1E
,.•• • •
• ••
• I :• FE
1
A
1
B
1
c
1
o
Fig. 5.2. Cumulative effect on the population of Earias sp.
(III) again 3 fresh eggs (F.E.) will be laid in each. but in A wherein the
incubation period is only 2 days the 3 eggs laid on the 1st day will have
70 AGRICULTURAL ENTOMOLOGY AND PESl' CONTROL
hatched (F.L.) by the time the 3 eggs of the 3rd day are laid, and the
population will remain only 6 in A, whereas in each of the other 3 there
will be 9 eggs. On the 4th day (IV) 3 fresh eggs will be laid under each
condition, but in A the 3 eggs laid on the 2nd day and in B the 3 eggs
laid on the 1st day will have hatched on the 4th day and there will be
only 6 eggs in A, 9 in B, and 12 in C and D eacq. Similarly, proceeding
on the 5th day (V), there will be 6 eggs in A, 9 in B, 12 in C, and 15
in D; and the population at this ratio and at these levels will continue
indefinitely till the same number of eggs (3) continue to be laid daily in
each and the incubation period remains unchanged. Undoubtedly, this
great variation in population from 6 to 15 as shown is not a real ope,
and it has been caused by the difference in the i.ncubation. period of the
eggs under different environmental conditions. That this virtual difference
is at times very serious in nature will be clear from the fact that the in-
cubation period of Earias eggs, for example, varies from about 3 days in
summer to 18 days in winter, i.e., in an extreme case the same number
of eggs wifi be found in the nefd even if the daiiy addition of fresh eggs
decreases in winter to one-sixth of what it was in summer. Thus, it is
quite necessary to make some correction in order to make allowance for
this virtual difference. One correction that suggest!; itself (Pradhan, 1947)
is to divide the egg counts by the incubation period. Thus, 15 divided
by 5, 12 by. 4, 9 by 3, and 6 by 2 will remove the virtual difference and
bring the index to the comparable level of 3 eggs in each case. The gene-
ralized correction equation may be written as Y-X/Z, where X is the
actual count, Y the corrected index, and Z the incubation period, larval
period, or pupal period.
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14 AGRIculTURAL ENTOMOLOGY AND PEST CONTRoL
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CHAPTER 6
ASSESSMENr OF LOSSES DUE TO INSECT PESTS
A large number of entomological laboratories and departments are being

maintained by governmental and other agencies mainly because of the
immense harm done by insects to human interest. The maintenance of a
department of agricultural entomology. for example, can be rationally
justified only on the basis of the estimated losses caused by insect pests
of crops and the proportion that can be expected to be saved with the help
of the results achieved by these departments. Naturally, ther_efore, the
question about the average annual loss suffered by the nation in various
spheres as a result of the damage caused by insect pests has been raised
a number of times by various bodies including Parliament A number of
committees have been constituted in the past from time to time for initiat-
ing work to provide an answer to this basic pertinent question. The leAR
constituted committees for this, purpose at least twice during the last three
decades. Both entomologists and statisticians have been fomulating elab~
rate research schemes on the as_sessment of losses. In 1958, the Entomolo-
gical Society of India circulated a questionnaire for obtaining information
on what was called the felt loss due to insect pests. Unfortunately, however,
the question, simple at first sight, has proved far too difficult to find an
answer. The committees ultimately suspended their work in a sort of silent
despair.
In the' absence of any correct estimate of losses caused by insects
guesstimate for example, is that the insect pests cause about 10% loss
annually, and on this basis the annual loss sustained by India from insect
pests amounts to about Rs 500 crore, simply because the national income
of India from agricultural and forest resources has been fluctuating about
an average of Rs 5000 crore. This calculation gives only a very rough idea
of magnitude of the problem and makes out a prima facie case for devoting
very serious attention to it. Like this, there are a number of statements
made by various scientists and administrators (e.g., Teotia, 1956) which
one often feels tempted to quote, but since none of them is backed by
adequate data collected on a sufficiently large scale, either in time or in
space, it is no wonder that these few individual opinions. even of persons
who ought to know, do not satisfy the public.
The position even in the scientifically advanced and rich countries is
not very different. The president of the American Association of Economic
Entomologists devoted his whole address to the annual meeting in 1942
to the needs for a sound system of 'Annual Insect-damage Appraisal', how
it could be organized and operated, and the benefits which might result
from its use (Parker, 1942). It is interesting to note from this address that
the round figure of 10% loss due to insect pests dates back to 1891 when
James Fletcher in his presidential address (quoted by Parker) said: "The
amount of damage done to crops every year is so vast that the figures
excite incredulity from those who do not study crop statistics. The agricul-
tural products of the United States are estimated at about $ 3,800,Ooo,odo,
of this it is thought that one-tenth is lost by ravages of insects." This
guesstimate seems to have gone round the world, and India's Bainbrigge
Fletcher about 3 decades later repeated the same round figure of 10%
loss to sugarcane in India due to insect pests (Hussain, 1938). This 'guess'
is current even today as the most popular estimate of loss due to insect pests.
James Flecher's fear of incredulity regarding the colossal figures of
losses due to insect pests persists. Parker in his 1942 address says: "Now
we find upon investigation that accurate estimates of damage done by
insects are exceedingly difficult to arrive at, and the figures are so large
that we are rather afraid to quote them ourselves lest we should prevent
rather than encourage investigation, and it has been the custom of ento-
mologists to minimize the estimate for fear they should not be believed".
The position has changed materially during the past 4 decades, but the
entomologists still find it difficult to convince the public about the losses
caused by insecets. It is, therefore, not surprising that another president
of the American Association of Economic Entomologists (Strong) should
choose for his 1946 address a subject entitled "Stabilizing Entomology"
and state: "Throughout the presidential addresses delivered to the Asso-
ciation almost invariably there has run, probably without definition but
none the less clear, one thought, in some cases a hope, in others a belief,
in still others the admonition that for the welfare of humanity entomology
must be stabilized".
Techniques of Estimating Losses Caused by Insect Pests

The problem of estimation of losses concerns the techniques to be
followed for these estimates as well as the organizational aspects. Accord-
ingly, the following methods are suggested on the basis of the various
techniques developed so far for estimating the losses caused by insect pests.
1. To grow a, crop as free from insect infestation as possible and then
to compare its yield with that of the check crop in which the insect activity
has been normal.
(i) Mechanical protection of the crop from pest damage. Efforts have
been made to grow various crops under wire-gauze or cotton-gauze enclo-
sures to keep out the pest, and then to compare the yield with that obtained
from infested crop under similar conditions. This principle has been used
ASSESSMENT OF LOSSES DUE TO INSECT ",ESTS 77
.with various modifications for estimating the losses caused by insect pests
of small grains (Borodin, 1926), by Hessian fly to wheat (Hill et al., 1943),
by jassids to potato (Peterson and Granovsky, 1950) and cotton (Mohammad
Afzal et al., 1944), by white fly to cotton (Hussain and Trehan, 1942), by
one or more species of insects to alfalfa (Stitt, 1948), by bulb fly to wheat
(Raw and Lofty. 1957). and by flea beetle to potato (Prasad, 1960). It
was observed in the case of jassid injury to potato that yield reduction was
more from low leaf hopper densities and 'proportionately less with further
increase in population. The obvious flaw in this kind of technique is that
the growth of the plant under such enclosures is seldom qu'ite normal;
sometimes the crop gets etiolated because of change in the environmental
conditions. Sucp flaws dQ not arise in the case of storage pest~ ana the loss
from these cart be determined more directly.
(ii) Chemical protection of crops tram the pests under investigation.
An effort is made to protect the experimental crop by the pest control
schedule known for a particular pest, and the yield is compared with that
under nbrmal insect infestation. This technique has been in vogue for some
time. Boll weevil losses were 'estimated by controlling the pest with the
application of calcium arsenate (Hunter, 1924). Other examples include the
experimental estimation of losses caused by spider mite to growth and yield
of cotton (Rousel et al., 1951), greenbug damage to small grains (Dahms
and Wood, 1957) in which infestation was produced with greenhouse-
reared insects and the insects were controlled by insecticide; crop losses
following insect attack on cotton in East Africa (Mckinley and Geering,
1957); damage caused by cabbage aphid (Strickland, 1957); reduction in
the grade and yield of com caused by sugarcane borer (Floyd et at., 1960);
the effect of infestation by pea aphid and lygus bugs on the yield of alfalfa
(Klostermeyer, 1962); and losses caused by aphid infestation of mustard
(Pradhan et al., 1960). Even in this widely used technique there is a the-
oretical flaw that the crop treated for chemical protection can also be
physiologically affected, for better or for worse, by the chemical.
2. To make use of the differential infestation under natural conditions
in estimating the loss caused by pests.
(i) Comparison of the yields in different fields having different degrees
of pest infestation. In view of the flaws pointed out above in mechanical
and chemical control of pests, many workers have tried to determine the
yield per unit area in different fields which have had different degrees of
infestation with a particular pest and then to work 6ut a correlation
equation between the yield of the crop and the degree of infestation. Thus,
an early cooperative estimate of the loss caused by sugarcane borers was
based on the correlation that a 100% infestation. i.e .. with every staJk
bored, represented a loss of one-third of the crop and a ]0% infestation
implied one-tenth of that one-third (Holloway and Haley, 1928). A

highly significant correlation was found between population estimates of
the meadow spittlebug and yield of red clover forage, and there was a
reduction of 12.7 kg/ha cured hay for each additional nymph/ft. Large-
scale tests in the field of mixed forage showed a reduction of 17 kg cured
hay/ha for each adult bug/sweep (Everly, 1959). Another study by
means of multiple regression analysis showed that an average of 1.5
borers/roof resulted in a reduction of 5.5% in yield of red clover (Pruess
and Weaver, 1958). Significant correlations have also been worked out
between flea beetle infestation and potato yield (Granovsky and Peterson,
1954) and between stem-borer infestation and paddy yield (Israel and Veda-
moorthy, 1955). Many similar examples exist in literature. The apparatus fot .
measuring insect injury to corn (Connell, 1956; Ditman and Ditman, 1957) \
appears to be quite a handy tool. From the practical point of view also
this technique is fairly sound although there are often various initial diffi-
culties in working out correlations such as those due to the changing nature
and extent of damage from stage to stage of the crop (Cheema, 1953).
Moreover, the very fact that fields grown under practically identical con-
ditions have shown different degrees of infestation is proof that unknown
factors can also affect yield.
(ii) Comparison of the average yield of individual pest-free and in-
fested plants. Individual plants in the field are examined for the degree
of infestation and their yields are determined. Thereafter, the average
yield of healthy plants in the same field is compared with that of plants
showing different degrees of infestation. The same data can be used for
working out a correlation equation between yield and infestation on the
basis of individual plants. Essentially the same technique has been used
with various modifications in a number of studies. For example, it has been
reported that 71 + 102 y number of earworms "I.. kernels are destroyed by
per sorghum head (Buckley and Burkhardt, 1962). The correlation between
damage by Chilo and yield of jowar grain is given by the equation .
Xl = 6.6204"/4 - 0.9257' Y:l - 27.17, wherein "1.1 is yield of jowar
grain per plant, X!l is percentage of length infested, and "1..4 number of
ears per plant (Pradhan ond Prasad, 1955). The damage caused by the
pod caterpillar to pigeon-pea seed was estimated by separately weighing
bored and sound grains (Argikar and Thobbi, 1957), and similarly the
loss caused by sugarcane borers (Haldane 1937; Pruthi and Narayanan,
]939; Kulshreshtha and Avasthy, ]957; Rajani, 1961; Siddiqi, 1961).
Reference should be made also to Borodin (1926) for different types of
calculations. The advantage of this technique is that the soil heterogeneity
factor is- expected to be considerably less in the same field than in different
fields. All the same, it is subject to the theoretical flaw indicated under
ASSESSMENT OF LOSSES DUE TO INSECT PESTS 79
(i) above i.e., the very fact that different plants sho:-v different degrees of
infestation is itself proof of some unknown factor leading to different
degrees of infestation. This factor may be genetic or physiological in
nature, or it may be mere soil heterogeneity from point to point in the
same field.
One flaw common to these 4 techniques to varying degrees is what
may be called the compensatory effect of the insect damage to a plant bn
the yield of an uninfested or a ll<ss infested plant in the neighbourhood,
or sometimes even on undamaged parts of the same plant, as ,a result of
ecological changes brought about in the environment (Lyubishchev, 1932).
In extreme cases one can find that in a check plot in which no control
measures were adopted many plants have succumbed to inseCt injury but
the few that ha~e esChped injury are in vigorous growth (Pradhan et al.,
1960). Moreover, if the yields in treated and check plots are compared on
the basis of yiel_d per plant, the average yield may be found to be more
in the 'latter than in the former. Such apparently anomalous situations are
obviously attributable to the fact that the few plants which escape insect
injury, while their neighbours succumb, have somewhat better ecological
environment, more space for development of their roots and shoots and
for their roots to draw nutrients from, and more light and air for their
shoots, etc. The compensatory effect on the undamaged part of the same
plant is mainly a physiological response to insect injury. For example,
insect injury to growing plants in early stages of sugarcane crop infested
by different species of borers stimulates tillering unless the seedling com-
pletely succumbs to the injury (Khanna, 1956). Cutworm injury to gram
crop is also known to stimulate branching, which in certain circumstances
is quite desirable. A striking example of some such type of compensatory
effect is afforded by the observations of Pruthi and Narayanan (1939)
that sugarcane stalks, infested by root-borer weighed more than normal
uninfested stalks, although this idea of compensatory effects does not seem
to have occurred to these workers.
The last 3 techniques often suffer from another serious defect from the
fact that 2 fields or 2 individual plants of the same field rarely differ only
in respect of 1 species of insect pest. Often more than 1 species is involved,
and it is extremely difficult to distinguish the effects of different pest
species'on the yield.
Unless definite care is taken to ensure that all other factors are kept
constant in the fields under differential insect infestation one may be faced
with apparently unusual positive correlation between insect infestation and
crop yield. For example, if the manurial status of 2 fields be different. we
may find that the one that has received more manure gives more yield
and also shows higher insect infestation. Manurial treatment which imporv-
ed the yield also increased the attack by Xyleborus lornicatus, and this
reduced the benefit of applying the manure (Gadd, 1944a, b). A physiolo-
gically healthy and well-fed crop is no insurance against pest infestation.
3. The average amount of damage caused by an individual insect.
In view of the various kinds of flaws pointed out above, another
approach which suggests itself is to determine the average amount, of
damage caused by an individual insect. This information can be collected
during primary studies on the biology of each species. Details can also
be worked out about the nature and amount of damage caused by the
various stages and ages of the insects. For example, leaf-eating species
may merely consume the foliage and it may be easy to determine the
average amount consumed. In the case of the phadka grasshopper (Hierog-
lyphus nigrorepletus) it was estimated that each individual consumes Oli'
an average 42 g of green leaf of maize during its life (Pradhan and Peswani,
1961). Somewhat similar work carried out in Russia on 5 species of
grasshoppers showed that an adult grasshopper devours 30 to 50% of
its own weight of grass in a day, and that during its whole development
an individual eats 20 times its weight in the adult stage (Rubtzov, 1932a, b).
In USA a single Mormon cricket was found to require 3.518 mg feed
from hatching till 20 days after reaching the adult stage; on this basis the
amount eaten by a known population was calculated (Cowan and Ship-
man, 1947). Lepidopterous larvae feeding on graminaceous plants were
found to require 20 times their own dry weight of plant, but only 7 times
when feeding on other plants. The same is true of coleopterous larvae with
similar habits. Parasitic insects on the other hand require only 5 times
(or less) -.t_heir own weight (Wolcott, 1925). Insects that produce a few
punctures and spoil the quality of fruit, fibre, etc., and those that carry
disease infections may require more complicated studies to determine the
average amount of damage caused by an individual. These studies, how-
ever, are comparatively easy and can be carried out in detail, and once
the basic information is there the loss caused by the pest in the field
can be easily assessed by estimating its population, preferably during its
different broods in the crop season. In the case of phadka the loss was
calculated as 18%, presuming that during normal years in Delhi the
population in the maize field goes up to 10/sq yd. The question of brood
did not arise in this case because the pest has only 1 generation in the
year. Similarly, in Russia it was found that a grasshopper population of
10 individuals/sq m would cause a loss of 607 Ib of grass per acre, and
that the total annual loss in eastern Siberia would be about 43,300 tons
of hay. This technique does not take into account the compensatory effect
discussed earlier which might be minimizing the loss to some extent. The
view that justifies the ignoring of these compensatory effects is that the
ASSESSMENT' OF LqSSES DUE TO INSECT PESTS 81
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ASSESSMENT OF LqS$ES DUE TO INSECT PESTS 91
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ASSESSMENT OF LO~SES DUE TO INSECT PESTS 93
amount which the insect consumes or damages should be estimated irres-

pective of the extent to which the injury is compensated for by ecological
or physiological factors. and that these should be studied separately.
The few exceptions are plants having an inbuilt tolerance to a certain
population of a particular pest, such as those that produce a particular
part in quantities much more than needed for the normal physiology
of the plant. For example anthers are produced in much greater quantity
than nt::eded, and the plants can tolerate a certain population of anther-
feeders. The flowers and fruits set are generally much mor~ than the
plant can bear; hence quite a high percentage of these flowers can be
destroyed by a particular pest without causing appreciable loss.
Table 6.1. lists
,
the various types of losses assessed by various means.
'
Percentage Increase in Yield and Loss Percentage

Owing to t~e great increase in losses from pests in recent years the
usual methods of calculation have become unsuitable. Agricultural ex-
periments are generally designep to determine the percentage increase in
yield due to different treatments, and the formula used for 'this purpose is
Percentage increase in yield due to treatment =
Yield in treated plot - yield in untreated plot x lOO
Yield in untreated plot
The same formula used to be applied in determining the efficiency of
different pest control agents or pesticides, and since the value of such
increase in yield was seldom more than 10 to 20% the same figure was
taken as percentage loss due to the pest infestation. But when the same
formula is now applied to the data given in Table 6.2, the increases due to
pest control work out to several hundred per cent, and these figures
cannot be taken as the percentage loss as well because loss obviously can-
not go beyond 100%. To overcome this difficulty the formula for calcul-
ating percentage loss has been modified as follows:
Percentage loss due to pests =
(Yield in plots treated (Yield in Plots not treated
for pest control) for pest control) x 100
Yield in plots treated for pest control
Even this modified formula is not quite correct because it erroneously
presumes control of pests to be complete in the treated plots and the yield
to be the real potential of the crop. In fact, with improved technique of
pest control even greater increase in the yield of treated plots is likely;
so the percentage loss will also increase (vide column 6 of Table 6.2).
Thus, the percentage loss worked out by this modified formula is always
likely to be an underestimate, Hence, a new term 'avoidable loss' has been
introduced to overcome this difficulty.

A study of the data in Table 6.2 will also show how, as the increase
in the yield .:>f treated plots goes higher and higher with more
effective control of pests with more effective use of pesticides, the increase
in percentage loss shown in column 6 does not show a commensurate
increase. Hence, care should be taken to compare the efficiency of different
pesticides on the basis of the figures not in this column but in column 5,
i.e., on the basis of percentage increase in yield due to pest control. Co-
lumn 6 is meant only to illustrate that the loss estimates and figures are
theoretically always underestimates and that the highest value available at
any time should be used as the estimate of the Joss for the time being.
However, care must also be taken to ensure that the experiments from
which such data are obtained are not so designed as to increase pest in':.
festation deliberately and give the percentage loss the crop suffers under
unnatural exper:mental conditions. What we should be interested to know
is what actually occurs in nature, not what can occur. Practically any pest
can produce a 100 o{, loss under specially designed experimental conditions.
Organizational Aspects
It will be seen that all the possible techniques for estimating losses
caused by different species of insect pests suffer from some flaw or the
other. All the same, for practical purposes any of these basic techniques
can be suitably perfected for a particular crop. The redeeming feature is
-that often the degree of accuracy needed in the assessment of damage is
not of a high order. Insect infestation and the damage caused by it differ
not only from field to field, or even from place to place, in the same season
b:It also in the same field and on the same spot from season to
season, and these variations range from 0 to 100%. Information re-
garding normal annual losses from insect pests in any locality has
to be based on averages of such wide variations. Hence, all that
is needed is to work out as accurately as practically possible a
relation between the insect population and the damage caused by it and
then to determine the population of the insect from time to time and from
place to place. The main point to be appreciated is that reliability of an
estimate like the annual loss caused by insect pests is not likely to be
affected so much by the fineness of the technique used as by the number
of places at which the observations are recorded and the number of years
for; which the data are collected. The best analogy is afforded by the
mateorological departments, which have existed in the country for a long
time. Temperature is a much more variable factor than insect popula-
tion; it varies considerably not only from time to time at the same place
but also from place to place at the same time. All the same, the meteo-
rological departments have collected very useful data of the average tem-
perature at different places, on which global isotherms are drawn;
reliability of these isotherms is based not on fineness of measurement but
on the large number of places at which temperature has been recorded for
so many years. The same is the case with humidity, rainfall, and other
weather records. The task of working out average annual losses caused
by' insect pests of various crops is of a similar kind; the bottlent;ck is the
absence not, of suitable techniques for assessment but of the organizatfon
needed for the purpose. The solution, therefore, lies in working, out some
kind of organizational liaison between the departments of agricUlture and
the meteorological departments of the country so that simple observations
on insect population qm be recorded at all those places where meteorolo-
gical observatio~s are being recorded regularly. Alternatively; each state
department of agrIculture should take steps to start estimating the losses
caused by major pests at a number of centres, so that after some years they
may be in a position to plan their pest control 9perati6ns with full know-
ledge of the losses they are seeking to avoid.
Felt Loss
The term 'felt need' is quite common in the field of economics. On the
analogy of this term the author, in 1959, as the President of the Entomo-
logical Society of India, circulated a questionnaire (reproduced herein) for
informa~ion regarding the 'felt loss' due to insect pests of different crops
in different regions of the country. Although the returns obtained in this
first attempt were too few to consolidate, they did prove the feasibility of
the step taken and the desirability of following the technique for the time
being. It will take years of patient observations at a large number of cen-
tres before average annual losses can be worked out on a really scientific
basis. In the meantime, therefore, it is necessary to crystallize our ideas
and consolidate whatever little experience is there about the losses which
people bear or at least believe they bear. The need right now is to carry
out an economic survey rather than an entomological survey of the losses
caused by insect pests. This economic survey will provide an index of the
average loss experienced by the cultivator during crop husbandry, by
various agencies during storage, and So on. It is certainly advisable that
a survey like this be carried out by economic entomologists rather than
by pure economists.
The idea suggested here is not new. The economists publish what they
call wastage rates during storage, reported to be for paddy 1.1 %, wheat
3%, barley 2%, jowar, bajra, maize and ragi 5%, small millets
2.5%, gram 2%, other pulses 2.5% potato 17%, and fruit 25% of the
total annual production in the country (Indian Agriculture in Brief, issued
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TABLE 6.7. SOME RECORDS OF FELT LOSS DUE TO INSECT PESTS IN INDIA
Crop or crop product Pest Loss% Reference No.

Sugarcane All pests 10
--<10-
(as raw sugar) Pyrilla, borers, termites 16 2
Foodgrains All pests 10-15 4
Wheat Wheat weevils 1.6 5
Pests and other causes 3.0 3
Rice Pests and other causes 1.1 3
Swarming caterpillar, rice
stem borer, rice caseworm,
rice bug, rice grasshopper,
rice hispa. 10 2
Other cereals including Grasshoppers, termites
jowar, bajra. ragi, cutworms, hairy caterpillar,
-small millets, wheat stem borers and armyworm 7 2
and barley
Jowar Pests and other causes 50 3
Bajra -do- 5.0 3
Maize -do- 5.0 3
Ragi -do- 5.0 3
Barley -do- 2.0 3
Small millets --<10- 2.5 3
Pulses (including Red hairy caterpillar, pod borer,
gram) gram caterpillar and cutworms 5 2
Gram Pests and other causes 2.0 3
Other pulses -do- 2..5 3
Potatoes -do- 17 3
Jassids, aphids, cutworms,
white grubs s 2
Tobacco Tobacco caterpillar, stem borer,
aphids, cutworms s 2
Groundnut (nuts in Hairy caterpillar, termites
shell) 5 2
Other oilseeds Mustard aphids, hairy caterpillar,
including mustard, semilooper, capsule borer,
castor, sesamum and leaf and pod caterpillar
lins,eed s 2
Jute Semi-looper, hairy caterpillars,
Jute apion 5 2
Cotton Bollworms, jassids, cotton
leafriller, cotton stem borer,
aphids, whitefly 18 2
Tea Tea mosquito, looper caterpillar,

leaf eaters, aphids, termites, thrips 5 2
Coffee Green bug, borers, mealy bugs,
cutworm 8 2
Coconut Black headed caterpillar,
red palm weevil, rhinoceros beetle 5 2
Chillies Chilli thrips 10 2
Pepper Mealy bug, tiel! beetle 5 2
Miscellaneous Aphids, caterpillars, borers,
non-forecast grasshoppers, termites,
crops, vegetables, cutworms, bugs, fruit flies, ,
fruits etc. thrips 6 2
Fruits Pests and other causes 25 3
"I. Fletcher, T. B. (1920) quoted by Hussain, M. A .. (1938).

2. Agricultural production in India and estimated losses due to pest damage
issued_ by Burmah Shell (1959).
3. Indian Agriculture in Brief. Directorate of Economics and Statjstics, Ministry
of Food and Agriculture, Govt. of India, 3rd Edition, 1957.
4. Krislll Sansar 1963. 1 (4).
5. Reports on Marketing of Wheat in India 1937, quoted by Hussain, M.A. (1938).
by Economic and Statistical Adviser, Ministry of Food and Agriculture,

Government of India, 1957, p 62). Obviously, a large portion of this wastage
is due to insect infestation. If economic entomologists carry out surveys
like this it should be possible to get a more reliable index from the ento-
mological viewpoint than mere individual impressions. It is true that
figures like those mentioned above, in the present stage of our knowle!lge,
can hardly be called estimates~ but even as 'guesstimates' it is better if
they are based on collective rather than individual guessing.
In fact, practically all the statements made by various scientists as well
as administrators regarding losses from insects, right from those of James
Fletcher in America and Bainbrigge Fletcher in India, should be rationally
considered as records of felt loss (Table 6.7). In comparatively recent years
when people became rather conscious of the limitations of such state-
ments they began to devote attention to the techniques of estimating losses.
and they became so much entangled in the mesh of the methodological
details that the over-all view was lost. Thus, many of the Indian workers
whose techniques have been reviewed in the foregoing pages, for estimat-
ing losss due to pests of cotton, sugarcane, paddy, jowar, potato, mustard.
etc., have also recorded their estimates obtained during rather small-scale
experimentation (Table 6.8). Although these estimates are based on definite
experiments, they represent only what happened in particular studies, and
TABLE 6.8. SOME LOSS ESTIMATIONS CARRIED OUT IN INDIA
(Although these estimations are based on definite experiments they only represent
what happened in particular studies and give no idea about the average loss
due to any pests, to any crop, at any place or in any year)
Crop or crop Pest Loss Reference

product
Sugarcane Top borer

(2 years) 5.7% and 2.4% Rajani (1961)
(Shoots) 12.7% (Var. 'Co 321') Siddiqi (1961)
10.7% (Var. 'Co 421')
Chilotraea 16% (about 105 md. Kulshreshtha,
auric;lia of cane per acre) et al. (1957)
Stem borer
(2 years) 2.3% and 5.8% Rajani (1961)
Stem and root 15% (Var. 'Co 321') Siddiqi (1961)
borers (shoots) 22.5% (Var. 'Co 421')
Root borer 3.1% and 4.2% Rajani (1961)
(2 years)
All borers 27.7% (Var. 'Co 321')
(Shoots) 33.2% (Var. 'Co 421')
(Net loss at 18.0% (Var. 'Co 321')
shoot stage) 22.2% (Var. 'Co 421') Siddiqi (1961)
(Millable canes) 19.0% (Var. 'Co 321')
14.2% (Var. 'Co 421')
(Total loss due 37.0% (Var. 'Co 321')
to borers) 36.4% (Var. 'Co 421')
Borers 4% in weight of canes Haldane (1937)
Termites 1.3% and 0.8% Rajani (1961)
(2 years)
All pests 7.90% (Var. 'Co 331')
12.81% (Var. 'Co 210')
9.10% (Var. 'Co 231')
11.62% (Var. 'Co 313') Pruthi et al ..
3.92% (Var. 'Co 299') (1939)
Mustard Lipophis eryslmi 51.6%, 71.5% and_ Pradhan et al.
(3 years) 87.9% (1960)
Coconut Oryctes rhinoceros 4.9% nuts Ramachandran
(1961)
they are too few to give any idea about the average loss due to any pest,
to any crop, at any place. or in any year. Therefore. it is advisable to send
round a questionnaire somewhat like the one reproduced here and to
collect as many ideas. experiences and guesses as possible. and then to
subject the returns to as rigorous an analysis as practicable so as to pool
and consolidate the information for the different regions. Some such app-
roaches have been made use of in various countries with great advantage
(Hyslop. 1938; Smith, 1938).
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Rubtzov, I. A. 1932 a. PI. Prol. Leningrad 1 : 69-80.
Rubtzov, I. A, 1932 b. PI. Prot. Leningrad :z : 31-40.

Siddiqi, Z. A. 1961. Proc. 4th All India Congo Sug. Res. Dev. Work India 541-546.
Smith, R. C. 1938. J. eeOll. Ent. 31 (5): 618-622.
STitt. L. L. 1948. J. eeon. Enl. 41 (5): 739-741.
Strickland, A. H. 1957. PI. Path. 6 (1): 1-9.
Teotia, T. P. S. 1956. Agric. Anim. Hush. (U.P.) India 6 (10): 7-10.
Weiss, H. B. 1940. J.N.Y. ent. Soc. 48 (2): 195-199.
Wolcott, G. N. 1925. J. Dep. Agric. Puerto Rico. 9 (1): 47-58.
CHAPTER 7
INSECTS' ADAPT ATIONS TO ARID CONDITIONS
MORPHOLOGICAL ADAPTATIONS
ADAPTIVE colouration in animals has been interpreted as serving 3

main purposes: concealment, advertisement, and disguise. "The biological
function of these elusive, attractive or deceptive devices varies widely
according to circumstance", but there is little doubt that Nature has ex-
ploited this principle in both arid and non-arid Lones, probably more so
in the former, where conditions of life are harder. There are innumerable
examples of insects, as of ~ther groups of animals, which illu,strate diffe-
rent categories 6f adaptive colouration, but the subject has not been dealt
with on any kind of zonal basis-not even on the basis of zoogeographical
zones, much lesson that of arid and non-arid _zones. The tesult is that
we know little about the subject of insect adaptation to arid environment.
It would be interesting to analyse all these examples on the Ibasis of arid
and non-arid zones.
In a general way it has been remarked that "the traveller who visits
one of the arid regions of the earth, such as the Kalahari, Sahara, or the
deserts of nort1~-western India or Southern California, will look there in
vain for brilliant greens". grasshoppers, mantids, etc. Instead he will ob-
serve that these creatures are "with few exceptions clad in colours borr-
owed from the desert itself-ochre, buff, brown and sandy grey, broken
perhaps with patterns of dark brown, black and white". There are many
species of moths. especially those belonging to the family Geometridae,
which have unicolourous appearance. These like the desert species, differ
from their near allies living in other types of climate just as much in
their pale ventral surface as in their buff or sandy backs. This unicoi our-
ous appearance is present in various groups of insects found in the desert.
suqh as moths, bugs, beetles, bees, flies and wasps. Some desert insects
have a silvery or sandy appearance. The phenomenon is found in many
bees, wasps, fossors, and in at least 3 families of flies (Bombyliidae.
Nemestrinidae and Tabanidae). It has been said that no one who has gone
on a collector's trip to the deserts of Algeria, Egypt, or Palestine can
doubt that the very pale colours and shimmering silver or golden pruino-
sity in all groups of Hymenoptera in such localities are cryptic characters.
A similar condition prevails in the gad flies of Mesopotamia and South
Palestine and in certain flies of the families Bombyliidae and Nemestrinidae.
The only exception to this generalization is that any desert creature which
is not coloured like its surroundings is black. Examples of black coloured

in desert insects are afforded by Tenebrionidae, Chafer beetles and several
species of Bombyliidae in the Great Palaearctic Desert. There are also a
number of examples of whitish or pale, melanin-free forms in semi-arid
regions of South Africa. Again, there are several species of beetles and
weevils including many white curculionids and a species of light-coloured
tiger beetle whose dark colour stands in glaring contrast to the white
background of the White Sands National Monument in New Mexico. Of
the white pronuba moth it has been stated: "One of the desert's strongest
partnerships is that of this white ghostly moth and the creamy yucca
blossom which it helps to pollinate". Of the 2 seasonal colour forms of
the butterflies of Africa, that of the dry season is said to be the one better
adapted for concealment. Thus, the characteristic colours found in the.
desert are (i) typical desert colour, e.g., sandy grey, ochre, etc., (ii) black. \
(iii) white. It has been explained that high temperatures increase the rate
of oxidizing processes, which in tum may influence the nature and distri-
bution of pigmentation. The most widely distributed pigments in insects-
brownish to blackish or black tints are due to O-dioxybenzol, melanin and
its derivatives.
As regards the biological significance of colour adaptations in desert
insects, Buxton sought to investigate the existing belief that the colour
of desert animals was meant to protect them from their enemies. He was
at once faced with the paradox of the frequency of black, which makes the
creatures more conspicuous against the background of sand and absorbs
more radiant heat resulting in greater evaporation of water, thereby proving
a severe handicap under circumstances in which water is always a limiting
factor. After discussing the subject at length he maintained that "protec-
tive colouration cannot be accepted as a theory to explain the very remark-
able colouration of desert animals"; but he predicted that "a theory of
advertisement might be launched, and could be almost as well supported
as the more popular theory of protection". Within 2 decades of this, the
appearance of Cott's treatise giving innumerable examples of, and unass:
ail able arguments for, both protective and advertising colouration marked
notable progress. "It has now been established beyond question that many
aposematic insects and other animals are highly distasteful, while others,
and especially cryptic forms, are greatly relished by different predators".
About Buxton's objections Cott says: "It is objection of this sort which
Buxton implies when he suggests that the colouration of desert animals
cannot be protective because certain desert forms and various beetles are
conspicuously dressed in black. Such a dress he believes must be regarded
as unsuitable unless we discard the theory of protective colouration. Yet
in certain cases, at any rate, his exceptions support rather than discredit
INSECTS' ADAPTATIONS TO ARID CONDITIONS 107
the theory. for they refer to animaJs whose habits preclude the necessity
for cryptic resemblance." Cockerell, discussing Buxton's objections. says
with regard to the black beetles: "They usually have an offensive odour
and their conspicuousness may be considered to facilitate recognition and
avoidance by possible predators. We have found that the species of the
tenebrionid-genus Eleodes, a characteristic of our south-west, when placed
in a bottle with a wad of cotton over them, will kill other insects placed
in the bottle by their fumes. At the same time some desert insects not
thus protected are intensely black". Thus, it is clear that many of the
examples of dull-coloured and black insects can rightly be expected to be
cryptic and aposematic adaptations, respectively, although it qlay require
close investigation to explain each individual case.
There is yelt another.. possible explanation. Cryptic, aposematic, and
mimetic adapta'tions are all reactions to biotic environment. The utility of
these colours as special adaptations against hot and arid physical environ-
ments requires inve~tigation. Hesse pointed to "the fact that very many
species of black tenebrionids and many carabids and cicindelids are also
among the fastest runners. Tllere is also reaSQll to believe that their
locomotor energy and extreme activity, during the day at least, may be
deriveq from their black colour, which apart from absorbIng heat energy
also favours the absorption of blue, violet, or ultraviolet rays, which
stimulate metabolic activity. The much debated and paradoxical black
colour of the majority of beetles inhabIting hot deserts may thus prove
to be a physiological response to the actinic rays in sunlight, the physio-
logical effects of which are advantageously transformed into locomotor
activity". Similarly, about the white colour he writes: "There are numer-
ous whitish or pale-coloured, melanin-free forms which, though apparently
mimicking the pale colours in their environments, nevertheless respond
to heat and light in that they are able to reflect the heat rays
which tend to raise their body temperatures". Wheeler, too, has
drawn attention to mutillids which have either pure white or even dense
carmine yellow or black pilosity which is serviceable in reflecting rays.
Thus the probable uses of desert colouration are :
(i) cryptic adaptation for protection against enemies,
(ii) aposematic adaptation fot', advertising their distasteful or harmful
nature.
(iii) adaptation for reflecting back the rays of the sun, and
(iv) adaptation for absorbing actinic rays from the sun and trans-
forming that energy into locomotor activity.
The basic plan of insect morphology is in fact an adaptation to aerial
existence as a change from aquatic life, i.e. an adaptation to a change
from moist to comparatively dry environment. However, we are concerned
at present with further specializations in the same direction that have

enabled some insects to face especially inhospitable conditions of arid
and semi-arid regions.
Heavy Well-developed Integument
About the semi-arid -regions of South Africa it has been said: "In
order to prevent the excessive evaporation of water and body fluids which
are physiologically necessary for the maintenance of life, an adaptive
response ip. the form of a very hard almost impervious chitinous exo-
skeleton has been developed in numerous insects, especially in Coleoptera
such as Tenebrionidae, Curculionidae, Carabidae and Cicindelidae. In the
case of such Coleoptera as inhabit arid regions this exoskeleton has pro-
gressed so .far that all the sclerites have become fused, carapace-like".
Similar adaptations have been pointed out by observers in other regions.
In some species inhabiting the hot and dry region of the Mericopa and
Colorado Deserts the large mentum makes it possible to close the buccal
aperture and the interlocking of the last ventral segments and the lower
margin of the epipleura at. the elytral apex, especially in the Eurymetopini,
practically sealing up the body against the drying effect of the desert.
Pilosity
"The significance of such exoskeletal structures as a dense coat of
pubescence, dense reflecting and resplendent hairs, or opalescent scaling
which adorn nllmerous Bombyliid flies, Asilids, and species of bees in
semi-arid regions, is apart from teleological explanations still obscure.
The ornamental value of such exoskeletal structures which usually excite
man's aesthetic sense of colour and harmony is probably insignificant, and
such structures are probably more of the nature of structural responses
which have been developed for reflexion of light and the prevention of
heat absorption". Similar phenomena and views have been recorded by
others for Phyllophaga hirticula and mutillid wasps.
Winglessness or Fused Elytra
Strong winds and violent vortices are among the most important
characteristics of arid regions, and the most striking modification to avoid
their ill-effects is the loss of power of flight. Acquisition of wings and con-
sequent power of flight was the most important modification in the
evolution of insects from aquatic to terrestrial and then to aerial existence.
It is, therefore, interesting to find that in the winglessness of insects speci-
alized for existence under arid environments there came a stage when
the possession of wings became disadvantageous, and a retrogressive step
had to be resorted to. In Algeria and Tunisia about half the Orthoptera
are wingless, and in many desert Tenebrionidae and Carabidae the wings
are fused. The May beetles (Phyllophaga lanceolata) adapted for prairie
existence have wingless females, but those confined to still drier regions
(P. farcta and P. cribrosa) are wingless in both sexes. In the South African
beetles Brachycerus and Psammodes this adaptation, besides reducing the
power of flight, is bound to reduce loss of water from excessive evaporation.
The loss of power of flight is due solely to the wind and not to any other
influence; for the phenomenon is by no means confined to desert species,
and a similar loss occurs in many other environments such as .mountain
tops and isolated islands which are eXPQsed to violent winds but do not
resemble deserts in any other respect.
Long Legs
Hesse stresses the, point that many tenebrionids (Stenocara, Adesmia,
Trachynotus), many cicindelids (species of Mantichora) and carabids (spe-
cies of Anthia) have developed elongated legs which are admirably adapted
to cope with sand and enable these insects to run very rapidly over such a
shifting medium. The utility of these elongated legs in keeping the body
away from the hot sand does not seem to have been studied in these beetles,
but it has been studied in other insects.
Oval or Compressed Body

The species of Zophosis (Tenebrionidae) and to a certain extent many
species of Graphipterus (Carabidae) have developed a smooth, slippery,
hard, oval and compact body which enables these beetles to wriggle along
over the sand with astounding rapidity. The shape of the body is similar
to that of many aquatic beetles. Another sort of adaptation is found among
some grass feeding insects of the order Homoptera, whose bodies exhibit
a strong tendency towards lateral compression when in the prairie. On the
other hand, the extreme dorsoventral compression in the tenebrionid
genera Eurychora, Steira, Lycanthropa and Geophanus and in hemipterous
genera Eupododus, Scantius, Brachyrhynchus and Neuroctenlls enables
them to eke out an existence in the limited and restricted environment
under a flattened stone.
Organs for Digging into Sand

In the case of beetles, such as the tenebrionid genus Gonopus, the
front tibiae have become flattened and blade-like, and are used for digging
into the sand. In the fossorial wasps, such as sphegids and musarids, there
is another type of structural response in the form of a brush on the tarsal
joint which enables these insects to scoop out their nests in a sandy envi-
ronment. In many unrelated genera of desert ants there is a structure called
psammophore which consists, of a row of long curved hairs on the clypeus

and gula or lower surface of the head in workers and females; it is used
as a crate or basket fOll transporting sand to the surface.
Organ for cleaning

There is a remarkable development in many species of ants living iIi
sandy and dusty places which has been called a 'circumoval crate'. This
consists of stiff bristles developed upon the structures which surround the
mouth, and its purpose is probably to clean the strigil on the foreleg, which
in turn is used for cleaning the antennae, etc.
Organs for capturing Prey

All predacious animals have specialized organs for capturing prey.
Several species of Myrmelionidae (Neuroptera) and Cicinde1idae (Coleo-
ptera) possess formidable pincer-like jaws.
Adaptations in internal Anatomy

The respiratory and excretory systems appear to have undergone
special modifications during the evolution from aquatic to terrestrial and
then to aerial life among the Arthropoda leading to Insecta. It is, therefore,
no wonder that modifications in these systems should continue in the
same direction, since their possessors have to face more and more arid
environments. In a description of a unique hydro-filter device for water
conservation in the excretory system of certain beetles it has been pointed
out that the malpighian tubules are characteristically confined to the
mainly terrestrial arthropods (i.e., most insects, myriopods and arachnids).
On the other hand, in those groups of arthropods, which have been
primarily aquatic, i.e., Crustacea and Xiphosura, the renal excretion is
supposed to be performed by nephridial glands. The main physiological
difference between these two types of renal structures is that, whereas the
latter throws the liquid excretion outside the body, the former discharges
it into the gut whence the water component of the excretion is reabsorbed
and retained in the body. Thus, it is quite likely that even the original
development of the malpighian tubules was related to the need for water
conservation during the change from aquatic to terrestrial life. The malpi-
ghian tubules seem to have undergone further modification for still more
efficient conservation of water in the case of certain insects, e.g. meloid
beet{es, several species of which are xeric, by a process of reassociation
of malpighian tubules with the hindgut. This has been shown to constitute
in coccinellid beetles a hydro-filter device for water conservation, which
acts as follows: There is a luminal connection between the haemocoele and
the hindgut in the region of reassociation. The water from the excreta
INSECTS' ADAPTATIONS TO ARID COl'!DlTIQNS 111
stored in the colon is mechanicalIy pressed out througl;l the minute open-
ings at the junction of colon and rectum into a fascial envelope which en-
closes the reassociated segments of the malpighian tubules. These segments
extract the toxic substances from the pressed out water which, after puri-
fication as in sewage works, enters the body cavity through the anterior
opening of the fascial envelope.
Reference to the significance of rectal glands will also reveal a trend,
of specialization in the same direction as indicated above, i.e. more efficient
economy of water. Development of the tracheal system and the m1chanism
of spiracles are directed to the same end, i.e., existence of life a\yay from
aquatic and moist environments; but not much attention appears to have
been paid to further specializations in this system for existence under arid
and semi-arid conditions.
PHYSIOLOGICAL ADAPTATIONS
Besides structural modifications or behaviouristic adaptations there
is a form of physiological acclimatization of certain insects t9 the hostile
conditions of arid regions.
Capacify to withstand Heat

Many insects are able to live on the surface of the soil at midday even
during the hottest season. Temperatures up to 78 'C have been recorded
on the surface of sand dunes in the Sahara. This serves to show the degree
of heat to which insects are exposed on the bare desert. A small shining
black tenebrionid, Zophoris punctata was common at all times of the day
during May, June and July around Jerusalem, moving about on bare soil
when it was as hot as 60° -62'C; with its body close to the surface it was
probably exposed to more trying conditions than a large insect would be.
Field observations and laboratory experiments on comparative capacity for
heat tolerance in insects that normally live on sand dunes and in those
that are not so acclimatized have shown that while the female mutillids,
which are typical dune insects, were able to withstand the highest
temperatures, insects that do not normally live on the dunes seemed
to lack the required endurance and reactions. In general, insects
that live on the dunes during the heat of the day have their minimum
and maximum effective temperatures located at higher points on the scale
than others. The carabid beetle Geopinlls, which is active on the dune at
night, has the lowest minimum effective temperature. On the other hand,
Bembix and Microbembix are typically diurnal and have high minimum
and low maximum effective temperature. For example, a change of 2'C
brought Chlorion (Sphecidae) from dormancy to normal activity, and
they also passed from normal activity to heat rigour very quickly at the
upper limit of the scale.
Adaptations to Existence under Particular Conditioll$ of Humidity

Some insects have adapted their physiology to life under semi-arid
.conditions; among these are some that have become so specialized that
humid conditions prove injurious to them. For example, the chinch bug
(Blissus leucopterus), an important prairie insect, is held in check in ~et
years by the fungus Sporotrichum globuliferum. The false wireworms
(Eleodes spp.) which are typical prairie forms, occur somewhat rarely on
the high prairies of Riley County in USA, but their number increases
westwards as the rainfall ceases. The Hessian fly (Phytophaga destructor),
on the other hand, thrives best in wet years.
Capacity to Jive without Water for Long Periods

A remarkable instance of ability to exist without moisture is furnish-
ed by the straitiomyid fly Hermetia chrysophila. Its larvae are able to
endure for at least 15 months without food or water, and eat decayed cactus
joints whenever moisture is present. Many insect species depend solely
on metabolic water.
ADAPTATIONS IN DEVELOPMENTAL PROCESSES AND STAGES
Without going into details of the evolution of the developmental pro-

cesses in insects, one may not be very wrong in assuming that the post-
embryonic development of insects as a whole is generally well-suited to
life under dry conditions. The introduction of larval stages whose mode
of life and requirements are essentially different from those of the adults
definitely reduces competition between these two stages of the same species;
often the occurrence of the stages is adjusted to different seasons of the
year to avoid the conditions adverse to each. Between these 2 essentially
feeding and reproducing stages is interposed a pupal stage, which is.
generally much more resistant to changes in weather conditions than other
stages and is also free from any food requirement. This is an ideal transi-
tion stage in post-embryonic insect development, and its termination with
the emergence of adults from pupae is in certain cases at least delicately
adjusted to the onset of weather suitable for the adult. Thus, insect life
history, in general, is basically well designed to cope with fairly wide
fluctuations in weather conditions, and we need be concerned here only
with some further modifications that appear to have taken place specifi-
cally in response to extremes of temperature or humidity characteristic of
arid regions.
Hibernation and Aestivation
Several authors have discussed this aspect at some length. Many insect
species of arid regions make good use of their capacity to remain in a
state of suspended animation for long periods. Most insects of the Great
Palaearctic Desert pass the 9 dry months of the year in a state of dormancy
and are active for a very short period which coincides with the presence of
annual vegetation. Similarly, high temperatures and hot winds necessitate
a period of dormancy in insect life of the prairie.
Longicorn larvae are known to' live long. The pebbly speci~s of Osmia
bees from Karoo and Namaqualand can 'sometimes be kept for ,2 or 3 years
before the bees emerge. In the Karoo, Bushmanland in Southwest Africa
the "regions which have been drought-stricken for years becoJ:l1e magically
transformed int'o a paradise, blooming with flowers and teeming with all
kinds of insects after favourable rains". In the case of the phadka grass-
hopper in central India, even if there is rainfall at the proper time, all eggs
may not hatch out, and some may remain dormant for the second or third
summer.' The potential utility of this sort of partial awakening from dia-
pause, is likely to be that, if the environmental conditions in anyone season
do not allow the nymphs to reach the adult stage and lay a fresh crop ,of
eggs, there may still remain eggs to hatch out next season to ensure con-
tinuance of the species.
Several examples illustrate the fact that some insects remain in an
arrested state of development or activity till the following spring, despite
favourable weather in autumn. It has been shown for the summer-winter
aggregations of Coccinella septempunctata on the peak of Elma Dagh
near Ankara that it is not in the interest of the species to resume activity
during autumn because first there are not sufficient aphids for massive
development of a new generation of lady beetles, and secondly the period
of favourable weather in autumn is too short to permit development.
Again, in Israel, there are maggots of Eurytoma amygdali and cocoon-
larvae of Cimbex humeralis in which diapause is not broken during the
favourable weather conditions of autumn. The former, if awakened, would
find no soft almonds for oviposition, the latter no soft almond leaves for
oviposition and as larval food. Thus, diapause has a great survival value
for it raises resistance of insects to unfavourable environment, fits many
of them into their local environmental rhythm, and almost always leads to
a synchronization of the optimal conditions of food and climate with the
active period of the insect concerned. It is the paramount factor in the
maintenance of the species within an environment which is not continu-
ously favourable.
Homodynamic and Heterodynamic Life Cycles

That there is generally an adjustment between seasonal variations in
the environment and the heterodynamic life cycles of insects is quite clear.
The typical prairie species Phyllophaga lanceo/ala and P. submucida pu-
pate in spring rather than in autumn as happens with most Phyllophaga.
This change is attributed to feeding habits, as the adults appear in mid-
summer (July and August) while other species appear in April, May and
June. On the other hand, the homodynamic life cycle has also been inter-
preted as a special adaptive response to conditions prevailing in a dry or
capricious environment because insects with such life cycles develop
successive generations as long as the conditions of temperature and humi-
dity remain favourable.
BEHAVIOUR ADAPTATIONS
For success in the struggle for existence. structural and behavioural
adaptations have to be compatible with each other. Both are purposive in
nature and, the purpose being common, the two have to follow convergent
paths of evolution. Behaviour adaptations of insects specialized for life
under arid conditions can be best described under the following headings.
Behaviour Adaptations for ensuring Security against

Inhospitable Environments
Variability in environment. The most characteristic feature of arid re-
gions is the wide fluctuation in the nature of environment both in time
and in 'space. This fluctuation becomes enhanced if the arid region happens
to be in the interior of a continental area. The recorded absolute range of
temperature fluctuation has been as high as 88°C at Kasalinsk in Turke-
stan and 5TC at Ghardaia in Algeria in one year. and 28 a -29T in summer
and 20o-22'C in winter in one day at In Salah and Wargale in Algerian
Sahara. These fluctuations at times are extremely rapid. In the sand desert
in Transcaspia the shade temperature was recorded as 3 'C, 20°C and,
28'C at 6a.m., 9 a.m., and 1 p.m., respectively. A rise of morning tempera-
ture in 1 hour by up to 4.2'C in August, 3°C in March, and 6°C in
December was recorded at Wadi Digla. Egypt. Fluctuations in relative
humidity are still wider and more rapid. The hygrograph records for August
at Wadi Digla, Egypt, show that humidity at times can drop from 95 %
to ~5% within a few hours. A rise of 75% in relative humidity in 7 hours'
and a fall of 19°C in temperature on the same evening were recorded on
a sand dune in Minnesota.
As regards fluctuation in space it has been reported that in Palestine,
where the surface temperature of the bare carth reaches 55' -62°C at mid-
day in summer, the shade temperature remains about 32'-38°<;. Great
differences in surface temperature within a few years are determined by

slight differences in situation. At Tucson, Arizona, USA. a range 42.5 °C
to 11 °C (i.e., 3I.5°C) was recorded in the shade, 71SC to 15°C (i.e.
56.5°C) just below the surface, 62.1 'C to 22'C (i.e., 40.1'C) at 2 cm
below, 35.6'C to 28.1 'c (i.e., 7.5'C) at 15cm below, and 24,S'C to
24.5°C (i.e., nil) at 100 cm below. The annual temperature range was re-
ported to be 42.2'C in shade, but 34.0'C, 22.6'C, 14.6'C, 9.0°C at 15 cm,
30 cm, 60 cm and 100 cm, respectively, below the surface. In Cairo: "Com-
pared with the range of temperature in a meteorological screen, tqe range
of the black bulb is 38°C greater; the white bulb 15'C' greater; on the
shaded side of the Wadi r less; under a big rock 5' less; in a bird's nest
in a hole in a roc~ 7.6° less'; in a bird's nest on a bush 2' mor:e; on the
surface of the sanG 19.4' more; at 5 centimetres in the sand 5' more and
at 10 centimetres deep in the sand 2' less". 'On a sand dune in Minnesota
it was noted that "there may be a drop of 10 degrees centigrade within an
inch above or below the sand surface when the temperature is high". Also
there may be a drop of as great as 20°C within 6 inches above .the surface.
The relative humidity at 3 inches below the surface was found to be 98 %
when that of the air was only 15%.
Site Preference
The biological significance of such wide variations in the kind of
environment that generally exists in arid regions is that insects adjust them-
selves to the time and place that suit them, and can exercise a choice
which perhaps results in their obtaining more favourable conditions than
they would have in a constant environment, unless that environment is
constant within their narrow zone of activity. Constancy either above or
below this narrow zone would be prohibitive. This scope for what may
be called site preference is actually the secret of the existence of quite a
varied fauna in areas which at first sight appear to be too hostile for most
of their inhabitants. Most desert creatures avoid the extremes of desert
climates by choosing suitable micro-climates for their seasonal or diurnal
resting phases. The following paragraphs will show that a large number
of behaviouristic adaptations of arid-zone insects are directed to this end.
Underground existence. Since soil and sand afford more suitable en-
vironment and better protection against harsh conditions above ground, a
permanent or temporary underground existence is very common among
insects in the arid zones. Ants, which are the most numerous xerophil6us
insects both specifically and individually, are invariably underground
dwellers. Associated with ants under stones are the tenebrionid genera
Arf?asidus, Smiliotlls, Geophanus, Acestus, and Adelostoma; and a number
of Thysanura are found under stones even in the driest parts. The const-
ruction of nests or provision of larders for the developing larvae under-

ground or in sand is the usual method of Psammocharidae, Masaridae,
many Sphegidae and even some kinds of beetles, and these insects are con-
sequently very well represented in sandy environment, where they may be
seen scooping out holes in the sand. The larvae of very many Curculionids,
Tenebrionids, Carabids, Scarabaeids, and even of Diptera are invariably
found in the soil. Many nocturnal Hymenoptera, Coleoptera and Orthop-
tera in the adult stage burrow into the soil or hide under stones, etc., during
the day, and many diurnal forms do so when the temperatures above
ground become intolerable. A number of ant species in the Great Pa]a-
earctic Desert and in central Australia have successfully overcome cIimati'c
difficulties, and many of them come up from their subterranean nests in the
morning and late evening, although some are active even during the mid-
day heat. Insects such as some Carabidae and Laps (Tenebrionidae) live
in burrows constructed by other animals. On sand dunes Geopinus remain
noticeable everywhere before. daybreak, but when the sun coines up they
all cease their other activities and begin to dig down into the sand.
An example of diurnal insects seeking refuge under stones or digging
into the soil is that of the large Tenebrionidae (Adesmia, etc.). They are
conspicuous in Algeria and Israel at all times of the day in March and April,
but seek shelter under stones and bushes in May and June. In the case
of the diurnal bembicid wasps, which are typical dune insects, "the pene-
tration of the hot surface layer is accomplished by a juggling of time and
space during which the wasps alternately dig furiously at the surface for
a short period of time and fly about six to twelve inches above the surface
of the sand. As the burrow deepens these flights become less frequent until
the wasps are well within the uniformly lower temperature of the deeper
sand". Thus, although the bembicids with their narrow temperature zones
of activity may not seem to be well adapted to the sand dunes, the duties
seem to be a suitable place for them to live in.
Some insects take refuge under stones during their seasonal rest. which
is invariably timed so as to avoid the inclemencies of weather. In the Great
Palaearctic Desert in spring and early summer one finds under the stones
such insects as Woodlice, lepismids, earwigs. cockroaches, crickets, beetles
bugs, and ants. Agglomerations of Coccinella septempunctata have
1:)een recorded beneath a few large flat stones on the top of Elma Dag.(an
elevation near Ankara), where a few hundred to a few thousand assembled
in a state of semi-torpor and were observed to remain so from 10 August
1939 to 11 April 1940.
An interesting specialization as an accompaniment of underground ex-
isten.ce is found in the pupal stages of Bombyliidae. the larvae of which
are predacious on eggs of locusts and are capabl~ of wrigglin~ their way
INSECTS' ADAPTATIONS· TO ARID CONDlI:rm'lS 117
out through the sand, coming to rest near the surface where emergence of
adults is facilitated.
Negative phototropism, negative thermotropism and nocturnal habit.
Negative phototropism (avoidance of strong daylight), negative thermotro-
pism (avoidance of high temperature), and nocturnal habit, are all res-
ponses that generally serve the same biological end i.e., keeping the insect
in a safe habitat.
Movement to favourable environments. In the normal course of the day
insects leave the sand surface when its temperature nears 50°C. Ants move
their eggs, larvae and pupae'to suitable situajions. In early spring a great
many colonies oU'semidesert ants inhabit superficial nests on hill slopes.
chiefly those facing the, south. Later, they migrate to north-facing slopes
with more favouraJ>le conditions, and may be found there throughout the
summer.
The army ant (Eciton spp.) cqlonies successfully adapt t~emselves to
adverse dry-season conditions. They generally manage to remain in the
vicinity of favourable zones, such as ravines, withdrawing into moist. and
dark recesses and underground places when the surface terrain is dry, and
going up into the hollows of standing trees during rains. In Riley County.
USA, the soil on the southern and south western slopes of hills dries
quickly, and numerous soil-inhabiting species which are commonly found
under stones or animal manure disappear rapidly, going deeper into the
soil on these exposed slopes than in the same situations on the northern
slopes. In Palestine, Heliothrips haemorrhoidalis feed on citrus leaves in
spring and early summer, but towards the middle of June they not only
migrate to the fruits but also congregate in hidden places on the fruits such
as the point where two fruits touch.
Confinement within small favourable space. Certain small blue butter-
flies (Lycaenidae) which inhabit the Great Palaearctic Desert possess the
power of continued flight within one small bush, from the shelter of which
they seldom issue; examples are the butterflies of the genus Tarucus which
remain confined within Zizyphus, and Chilades galba which remain in an
onion plant.
Construction of egg-pods, larval cases, pupal cocoons and nests. The
most important biological significance of all these structures constructed
by various insects is security against adverse environments. In dry regions
the larvae of numerous species of Tineidae and Psychidae have adopted a
novel 'method of protecting their bodies against enemies, insolation, and
high temperatures by constructing various types of larval cases. Examples
are the peculiarly flattened larval cases composed of silk with an admixture
of sand constructed by the caterpillars of the tineid genus Criticonoma,
and the bags composed of silk and dead leaves or sticks constructed by
caterpillars of psychids such as Acallthopsyche and Euneta and tineids
such as Melasina. As to their protective value. pupal cocoons are too
common to attract the attention Qf ecologists. The data on the develop-
ment of pupae of Earias fabia, which has an ordinary pupal cocoon, show
practically no effect of humidity changes. The pupal cases of many moths
on the other hand are composed of very tough membranes reinforced by
various kinds of materia1s; the cocoons of: many lasiocampid moths ale
often very hard and impervious, and afford effective protection aga,inst
weather in arid regions. The pitcher-like nests of Anthidium and the
pebbly nests of Osmia afford effective protection to these bees, with the,
result that various species of Anthidium are reported to have exploited
practically every type of niche in the barren regions of South Africa. Where
no other suitable niche is available, a species of Anthidium in the sand
dunes of the coastal belt of Namaqualand, and a eumenid wasp in the very
arid sandy coastal belt of Luderitzbucht and Namib, have learnt to utilize
the empty shells of snails.
Behaviour Adaptations for securing Sustenance
The problem of sustenance in arid areas is as difficult as that of secu-
rity against physical hostilities. If the environment is almost annihilating
to insect and other animal life, it is in no way partial to plant life on
. which the sustenance of all animal life depends. The result is that, except
during very short and irregular periods, shortage of food is endemic. Insects
specialized for arid zone: existence have found different solutions to this
perpetual menace. A considerable section of the characteristic insect fauna
of desert areas has become carnivorous, and specialization for predatory
existence has led to the evolution of (i) lochetic species, whose habit is to
wait till their prey comeS within striking distance and then suddenly seize
it with their specially efllarged raptorial forelegs, or (ii) swift hunters. or
(iii) a habit of collecting victims during periods of abundance and storing
them for their progeny. Others have developed highly specialized phyto-
phagous habits along 3 main channels of evolution. Some species have so
thoroughly adjusted their life history to the sporadic and irregular appea-
rance of their food plants that the active feeding stage invariably coincides
with the availability of tbe food plant in the proper stage; some have deve-
loped the habit of collecting their food and storing it against periods of
scarcity; and some have developed a polyphagous habit which enables
them to utilize a much larger spectrum of plant life either at the same time
or in a more less regular succession of food plants in the area they inhabit.
A carnivorous habit is also an effective method of supplementing the

spectrum of plant life on which any particular species has to depend. An
increase by one in the list of victim animal species means a much greater
increase in the primary source of food, unless the victim happens to be a
monophagous species. It has been stated that ratio of predaceous to plant-
eating species of arthropods is very high in desert regions. Certain peculiari-
ties of these predaceous forms are as follows:
Adaptation for carnivorous life. LOCHETIC BEHAVIOUR. The
literal meaning of 'lochetic' is lying in wait. entrapping. This behai,riour has
been exploited by a number of desert insects. In the larval stages several
species of Myrmelionidae (Neuroptera) and also of leptid fiies, which are
peculiarly adapteq' to lIfe in'dry sand, make conical pit in the sand and lie
hidden at the bottom of it. When a victim stumbles in unaware', it is cap-
tured before it is able to get out, the loose sand making escape the more
difficult. Another interesting case is afforded by cicindelids such as Manti-
chora spp., the elongated grub of which lies in wait in a cylindrical burrow
with its body curved in a peculjar way. It can discriminat y vibrations
caused by other insects and by animals.
Active hunting. Some species of ants, e.g., the genera Dorymyrmex and
Myrmecocystus in the deserts of America and the genus Cataglyphis of
the Great Palaearctic Desert, are quite active hunters. They are fast run-
ners and hunt fat and wide. Of the 4 main families of Coleoptera, which
inhabit the desert, Cicindelidae, Carabidae, Meloidae and Tenebrionidae.
the first 3 include active predators.
Specialization for phytophagous life. ADJUSTMENT BETWEEN IN-
SECT . AND PLANT LIFE CYCLES. This point has already been dis-
cussed while assessing the significance of hibernation and aestivation, which
constitute the principal mechanisms for an effective adjustment between
the eating stages of the insects' life cycle and eatable stages of the plants.
As the cyclic rhythm of reproduction in desert plants is to a large extent
dependent upon climatic factors and favourable temperatures, the seasonal
cycle of insects, which is directly or indirectly dependent upon them, coin-
cides with the maximum reproductive, activity of such plants. This is
largely also true for arid areas receiving' a little rain in the winter months.
In the arid region of Koko Head, Oahu, there is a scarcity of vegetation
for the greater part of the year; but a considerable growth of various
grasses and other weeds occurs after a few winter showers, and the insect
populations then quickly increase.
Polyphagous habit. How the polyphagous habit has been successfully
utilized for obtaining a regular succession of food plants was beautifully
illustrated by Papilio machaon, the swallow-tail butterfly in Mesopotamia.
In April and May it fed on the flowers of Ruta tuberculata, and as this
plant began to go to seed, Ammi majus came into flower, and became for
the moment the plant on which eggs were most commonly laid. This plant
was followed by Ducrosia anethijolia, which was not common and pro-
bably not an important food plant. While A. majus and D. anethifolia were
running to seed. Ammi visnaga flowered and became an important food
plant; it was in full flower in June after the first three were quite dry. A
fifth plant, Foeniculum vulgare, was also found to be eaten by the larvae
of P. machaon.
Food storage. The habit of collecting food in times of plenty and storing
it for periods of scarcity is common in harvester ants and honey ants.
Most of the harvesting ants belong to the genera Pheidole, Veromessor,
Messor, Oxyopomyrmex, Goniomma, Holcomyrmex and Pogonomyrmex.
Honey ants store the honeydew of aphids and coccids, flower nectar and
plant exudations in the crops of certain specialized worker ants. This phe-
nomenon is found in several genera, e.g., Pheidole, Melophorus, Myrme-
cocyslus, Camponotus and Prenoiepis, some of which are found in both
American and Australian deserts.
Fungus gardening. This habit appears to be generally confined to the ant
inhabitants of humid regions, but an exception is reported to be Maellerius
versicolor which does fungus gardening in the deserts of Arizona and
Western Texas.
Behaviour Adaptations for avoiding Enemies

The biological significance of various kinds of behaviouristic adapta-
tions for evading the predator as well as deceiving the prey has been dis-
cussed by various authors. The practice of concealment both in offence
and in defence, warning displays, orientation, movement and sound meant
to warn the predator or intimidate the prey, use of poison in offence and
defence, defensive secretions, nauseous taste and odour, adaptive beha-
viour as a complement of cryptic resemblance, and various types of
mimicry have all been discussed.
Reflex bleeding. Many species of the genus Anthia squirt out a fine jet
of liquid which is irritating and smarting to the eyes. The bombardier
beetles of the genera Brachinus and Crepidogaster emit a thick fog-like
cloud on being touched or disturbed. These reflexes have been described
from arid regions of South Africa and are believed to be of defensive
value. Reflex bleeding is also quite common in coccinellid beetles, several
specieS of which exist in arid regions of the world.
Threatening response. A desert mantis, Eremiaphila braueria of Balu-
chistan, when molested, suddenly unfolded its wings and stood up on its
hind legs like a performing bear, thus exposing the brightly coloured
ventral surface of both front hind wings. When the observer moved around
INSECTS' ADAPTATIONS TO ARID CONDIIIONS 121
the insect in a circle. it turned round, always facing and maintajning the
same erect posture. A similar attitude, accompanied by trembling and
shivering movements, is adopted by the carabid Anthia and the cicindelid
M antichora.
Dodging response. Certain Hemiptera and coleoptera play a kind of
hide-and-seek, the insect continuously dodging by slipping round and
round' the branch so as always to keep the branch between itself and 'the
observer.
Death feigning. This phenomenon is described in the case of curculionid
genera Episus, Microcerus and Brachycerus of the arid region 9f South
Africa. It is common in other environments and species also. '
Mimicry. There is a large amount of literature on mimicry. As regards
mimicry among !hSects of' arid regions, attention has been drawn to
resemblances among female mutillid wasps, ants, bombyllid fiies, and
other kinds of flower-visiting Diptera, many species of Bombylius and
anthrophorine bees, species of Systropus, Hymenoptera such as Sphex.
Sceliphron and Belonogaster, carpenter bees (Xylocopa), some robber flies.
syrphid flies, and wasps.
Behaviour Adaptations for ensuring Both Security and Sustenance

Parasites. A vast group of internal parasites have adopted a mode of
life ensuring a continuous supply of food as wen as security against biotic
and abiotic enemies. Alike in number and percentage, there are many
more primary parasites than hyper-parasites. This fact itself is evidence
of the comparative security of internal parasites against biotic enemies;
and their location inside their host affords them comparative protection
against climatic variations. That these advantages of parasitic existence
have been fully made use of by arid-region insects will be clear from
the many examples of typical desert insects such as bombyllid flies and
mutillid wasps which are entirely parasitic during the larval stages. There
are also a number of arid-zone representatives among the parasitic bees
(Crocisa and Coelioxys), Chrysididae, Ichneumonidae, Br~conidae, and
ChaIcidoidea.
Internal feeders in plants. What has been said above about internal para-
sites also applies to phytophagous inse,ct borers in stems, roots and fruits,
to leaf miners, and to gall makers. There are numerous examples of the
first 2 categories, at least in the cultivated areas of arid regions. Certain
cecidomyid flies and chalcidoid wasps make galls in the arid regions of
South Africa.
Symbiosis and commensalism. Attention has been drawn to myrmeco-
phil and termitophily in arid zones of South Africa. Besides several guests
in ant and termite nests, there are in the Karoo several genera, e.g.,
122 . AGRICULTURAL ENTOMOLOGY AND PEST CONTROL
Argasidus, Smiliotus, Geophanlls, Acestus and Adelostoma, that remain

associated with ants under stones.
Behaviour Adaptations for Sex Activity
Thtf two main points here are the tapping activity indulged in by seve-
ral species of Toktokkies (Psammodes) and the marriage flights of ants,
termites, etc.
CHAPTER 8
EFFECT OF TEMPERATURE ON THE DYNAMICS OF

I]'I.ISECT DEVELOPMEl'Sl'
THE problem of temperature effect on organic development is of so much

theoretical interest and economic importance that innumerable investiga-
tions have been conducted on it. Yet it has remained unsolved~ as will
be clear from the remarks of Wigglesworth (1942) : "This curve (of inten-
sity or velocity of development) represents the sum or resultant pf an
immense number, of chemi~al and physical reactions, many of which
must be differently affected by changes of temperature. But in sl?ite of this,
numerous attempts have been made to describe these curves by simple
equations. Such formul~e can be made to fit particular cases and to that
extent they have some descriptive value, but none- of them is of sufficiently
general application to be regarded as embodying any rational principle.
The present chapter attempts to put forward a clearer conception of this
phenomenon than before.
At first sight the topic may appear to be too mathematical for a biolo-
gical reader, but in fact it is not so. As far as possible, efforts have been
made to keep out all difficult mathematical portions so that they may not
disturb a biological reader. The equations which appear in the text stand
there as mere statements of facts; their derivation, etc., are given either
in footnotes or in appendices. This segregation of mathematical and biolo-
gical language may also help a mathematical reader who may like to
check the purely mathematical derivations before concentrating on their
biological interpretations.
Deduction of Equation
In a previous communication (Pradhan, 1945) it was stated that the
value of developmental index (reciprocal of developmental period like egg
period, larval period, pupal period, etc.) of Earias fabia corresponding to
any constant temperature is given accl1rately by the equation :
_ax 2
Y = Yoe (la)*
In that and a later communication (Pradhan, 1946) it was shown how use-
ful this equation, is for tackling problems of fluctuating temperature in the
field. The following paragraphs are meant to show how we logically arrive
at this equation and what physical meaning it carries or, in other words,
which biophysical processes it symbolizes.
The dynamics of development become crystal clear if we consider the
following simple case of hypothetical development. Suppose the single cell

of a zygote begins to divide. If the daughter cells maintain their power of
division unhampered and the development goes on constantly with a speed
of a division a minute, we shall have.
TABLE 8.1
After 1st division or at the end of 1st minute 2 cells or 21 cells
After 2nd division or at the end of 2nd minute 4 cells or 22 cells
After 3rd division or at the end of 3rd minute 8 cells or 23 cells
After 4th division or at the end of 4th minute 16 cells or 24 cells
After 5th division or at the end of 5th minute 32 cells or 2' cells
and so on
It is clear that, although the speed of division, i.e., the number of divisions
per minute, remains constant, the amount of growth (i.e., the addition of
cells) during the successive minutes increases in geometrical progression,
i.e., in accordance with the law of compound interest. If the time is fixed,
say 1 minute, and the speed of division is increased in some way, we shall
have growth as depicted below:
TABLE 8.2
Speed of cell division Amount of development in a fixed period
division per minute 2 cells or 21 cells at the end of 1 minute
2 divisions per minute 4 cells or 22 cells at the end of 1 minute
3 divisions per minute 8 cells or 23 cells at the end of minute
4 divisions per minute 16 cells or 24 cells at the end of minute
4 divisions per minute 32 cells or 2' cells ot the end of minute
and so on.
Thus, although the speed of cell division increases regularly in simple

arithmetical progression, i.e. from I division a minute to 2, 3, 4, divisions
a minute, and so on, the speed of development, i.e., the amount of growth
(addition of cells) per minute increases again in geometrical progression,
i.e., from 2 cells a minute to 4, 8, ] 6, cells a minute, and so on. Based
on the above examples we can write down the generalized equation :
g = 2n .............................. (1a)
A more accurate form of this is
g = en .............................. (2)1
because according to the established mathematical fact we take in account,
y 0 = highest value of developmental index

Y = developmental index at temperature to
e = constant = 2.718282.
X = T-t
T = temperature corresponding to y 0
EFFECT OF TEMPERATURE ON THB DYNAMICS OF INSECT DEVELOPMENT 125
by this modification, development at every moment and not merely at the

time of cell division.
If the total amount of growth (g) required to complete the full develop-
ment under consideration is taken to be constant, it can be shown mathe-
matically (2) as well, as it is easily comprehensible that if the value of (n),
i.e.~ the speed of cell division, is somehow increased, then the period
required for full development, i.e., the developmental period, will corre-
spondingly decrease, and vice versa.
Another point to be considered is that cell division is not ,he primary
activity in organic development; cells do not remain simply dividing and
redividing continuously, but each cell after formation develoI?s for a time
and, so to speak, attains' maturity before it is ready to divide again. Thus
a cell's coming to maturity (when it is ready to divide) also marks the com-
pletion of a series 'of vital processes. This fact is best illustrated by a
unicellular organism in which there is no cell division except at the time
of reproduction, but still development does ~ continue from moment to
moment. Therefore, just as the speed of development of, a multicellular
organism, say, up to maturity, can be expected to depend on the speed of
cell division, similarly the speed of coming to maturity of single cells and
hence the speed of cell divisio!1 (n) should depend on the speed of vital
activities involved in the development and maturity of single cells. Hence,
if we suppose
n = e r .............................. (3)
wherein (r) is the speed of vital activities involved in the development of
single cell and (n) the speed of cell division; then by substituting the
value of (n) in Equation 2 and taking a few mathematical steps 3 we easily
come to
y = e r .............................. (4)
wherein (y) is the developmental index, i.e., reciprocal of developmental
period (1/P).
It is clear from equation 4 that the value of (y) can increase only with
increase in the value of (r), the value of (e) being constant. Therefore, the
experimental fact that the value of (y) increases with the rise of tempera-
ture shows that the speed of vital- activities (r) increases with the rise of
temperature. Further, as shown by the so-called sigmOid curve of develop-
mental indices, the rate of increase in the value of (y) does not remain
constant but is continuously decreasing. These experimental observations
show that a somewhat similar change might also be taking place in the
value of (r) with the rise of temperature. Now, (r) being the speed of
activity, a continuously decreasing rate of increase in its value reminds
one of the principle of retarded motion, and deeper thought on this com-
parison increases the probability of the 2 phenomena being absolutely
similar. Therefore, let us suppose for the time being as a trial that the value
of (r)4 increases with the rise of temperature but with a velocity which
does not remain constant but is getting retarded and with the same rise of
temperature and that the retardation is uniform.
Now, the sort of change in the value of (r)5 as supposed above can be
represented by the equations :
v = u - at............ (5)
and
r = ut - tat 2 +c, .......... . (6)
wherein v = resultant velocity of change in the value of (r)
u = a constant, i.e., value of (v) when t == 0
a = retardation constant, i.e., negative acceleration
t = temperature
c = constant, i.e., value of (r) when t = 0
Further, a few more steps change Equation 6 into
r = - !ax 2 + log k .................. (7)
wherein x represents the same quantity as it does in Equation with which
we started; k is a constant.
Then, substituting the value of (r) from Equation 7 in Equation 4
we geF
e-!ax 2 . . . . . . . . . . . . . . . . . . (8)
y=k
and Equation 8 can be directly written as
-!ax2 .................. (9)
y =
yoe
Equation 9 is the same as Equation 1 except that (a) of Equation 1 is
equal to (!a) of Equation 9.
Thus, by simple presumptions and suppositions we arrive at an equa-
tion which is found to be promisingly applicable to what is supnosed to
be a complicated phenomenon, i.e, development, as will be discussed
further on.
Curve-fitting and testing goodness of fit.

(a) Curve-fitting. As regards curve-fitting it must be made clear at
the outset that for fitting Equation 1 we cannot adopt the usual technique
of fitting the frequency curves which a statistician might like to demand
at first' sight because of a close similarity between Equation 1 and the
equation of normal frequency curve. Neither are we dealing here with any
distribution nor do our data represent a random sample of the total popula-
tion of all the possible data. In fact, our observations have been most
systematically collected for definitely chosen points on the abscissa. i.e ..
the values of developmental periods and hence of developmental indices(y)
EFFECT OF TEMPERATURE ON THE' DYNAMICS OF INSECT DEVELOPMENT 127
have been determined for a few chosen values of temperature, which

actually are not chosen beyond a certain range because of very high pre-
imaginal mortality at higher temperature. Therefore, despite the similarity
rd~rred to above, we have to resort not to the special method of fi~ting
normal frequency curve but to the general method of fitting logarithmic
curve after converting the exponential form into the polynomial form of
equation. Thus, Equation 1 is converted int09
Y==AX2 + BX+C .... , ............. (10) ;
The values of the constants A, B, C can be determined simply by solv-
ing a set of normal equations1o • The calculations involved in this solu-
tion are illustrate9 in Appendix A for ready reference. In Appendix B is
given a modified form commonly known as the Doolittle method for
solving these equations; this modification besides reducing arithmetical
labour provides a check for correctness of calcu!ations at definite stages
and also readily gives figures required to test the goodness of fit of the
equation concerned (vide infra). F~rther, the value at (a) can be found out
from Equation A = -!a, and the value, of T from T = Bfa. The value of
(C) gives the value of log K and hence the value of (yo) by the equation
C = log K - -!a T2. Thus, for example, by fitting the curve to the tem-
perature effect on pupal periods of Earias fabia (vide Appendix A) we get
-!a = 0.001107
T = 97.3
yo = 13.69
Thus, the equation comes to be
- .001107 (97.3 - tOF)2
Y = 13.6ge
Such equations for different sets of data are given in the beginning of each
set in Table 8.4.
(b) Testing goodness of fit. 1. Analysis of variance test. For
testing goodness of fit of Equation 10, and thus of Equation 1, we can
apply the exact statistical test by analysis of variance as illustrated in
Appendix B in case we have individual. observations to start with instead
of average values of developmental periods corresponding to different con-
stant temperatures as have been given by most of the previous investiga-
tors. This test has been applied to nymphal periods of Schistocerca grega-
ria for which Hussain and Ahmad (1936) have fortunately published the
individual observations. It is found (as given in Table 8.3) that the fitness
of Equation 10 is highly significant.
Conclusion-Regression is highly significantl l .
2. Graphical test. Coming to the graphical method of judging the
suitability of curves, let us see what we should expect if Equation 1 corr-
~ctly represents the relationship between developmental index and tem-
TABLE 8.3 ANALYSIS OF VARIANCE SIGNIFICANCE OF DEGREES OF FREE-

DOM USED IN FITTING EQUATION 10 TO THE LOGARITHMS OF
DEVELOPMENTAL INDICES OF Schistocerca NYMPHS AT DIFFE-
RENT TEMPERATURES.
Degree of Description Sums of D.F. Variance Value of F

fitting Squares at 1%
1 Total 1.87978 87
Regression 1.54781 1 1.54781 400.8 6.96
Error 0.33197 86 0.00386
2 Regression 0.29266 0.29266 636.2 6.~6
0.00046
Error 0.03931 85
perature. Equation 1 can be written as

log y = Kl - AX2 .................. (11)
Equation 11 shows that if Equation 1 correctly represents the relationship
under consideration we would get a parabola if log y is plotted against x,
and a straight line if log y is plotted against X2; Fig. 8.1 to 8.6 excellently
satisfy this criterion. These 6 figures illustrtate 6 different sets of data on
developmental periods of different stages of 2 species of insects. Earias
labia and Schistocerca gregoria, belonging to orders Lepidoptera and
Orthoptera, respectively.
Line A represents the curve obtained when y (developmental index) is
plotted as ordinate against the abscissa of (temperature), whereas line B
is the graph obtained when log y is plotted against X 2, i.e., (T-t)2. It is
instructive to note how all the observed points of so-called sigmoid curve
A arrange themselves along almost a clear straight line as demanded by
the hypothesis of this paper. Fig. 8.2 and 8.5 illustrate this fact most pro-
minently, since in these figures graph A exhibits the greatest amount of
curvature. Corresponding points of observation are serially numbered along
both A and B as well as along scales of ordinate and abscissa. It should be .
noted that even points 6 and 9 of curve A in Fig. 8.2 and 8.5, respectively,
have come back to the same straight line in graph B. It may also be inter-
esting to remember at this stage that the advocates of the hyperbolic
relation between temperature and insect development try to find a straight
line in graph A itself. .
Agreement between theory and observation. Having determined the
valu_es of the constants of Equation 1, we have calculated with its help the
developmental periods corresponding to different temperatures and com-
pared them with observed values from 10 different sets of data on various
stages of 3 species of insects, 2 of which are those dealt with in last section
and the third is Microbracon Ielroyi of the order HY!!1enoptera. These
EFFECT OF TEMpERATURE ON THE DYNAMICS OF INSECT OEVELOPMENT 129
....J
-<:--"'--Ahl ---~ J
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,_ e,....
J
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~----A ----. t
• §:~
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I o
<:------- A 501-----~ I
..
I
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I I
I
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ii ~~
.
Co
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EFFECT OF TEMPERATURE ON THE DYNAMIQS OF INSECT DEVELOPMENT 131
calculated and observed values are given in Table 8.4, and it will be found
that the agreement is excellent. The observed values for Fig. 8.1 to 8.6 and
of Table 8.4 were taken from papers published by others-those for E. fabia
and M. lefroy; by Ahmad and Gullamullah (1939), and those of S. gregaria
by Hussain and Ahmad (1936). Therefore, the agreement between theory
and observation is all the more significant, since there can be no question
of the former influencing the latter. Of course, where more than one humi-
dity was tried we have taken the average of different humidities,.
TABLE 8.4. COMPARISON OF CALCULATED AND OBSERVED VAL\lES OF

D:gVELOPMENTAL PERIODS
Temperature Development periods Temperature Development periods

(in d:eyrs) (in da~s)
Calculated Observed Calculated Observed
1. E. fabia - egg period 6. M. lefroyi - larval .period

Y = 37.78 e - .001119 (97.3-tOF)2 Y = 54.1 e - .0011007 (95.02-t'F)2
60.8°F 11.78 11.58 60.8'F 6.70 6.40
68.0'F 6.92 7.10 68.0'F 4.12 4.52
77.0'F 4.19 4.16 77.0'F 2.64 2.56
86.0°F 3.05 3.01 86.0'F 2.02 1.94
95.0°F 2.66 2.68 85.0'F 1.85 1.89
2. E fabia - larval period 7. M. lefroyi - pupal period
Y 11.1 e - .001366 (92.2-tOFF Y = 20.66 e - .00112 (96.6-t'FP
60.8'F 34.64 34.96 60.8'F 19.66 19.44
68.0'F 20.03 19.86 68.8'F 12.10 13.01
77.0°F 12.02 11.92 77.0'F 7.44 7.25
86.0'F 9.49 9.93 86.0°F 5.48 5.23
95·0'F 9.12 8.90 95.0°F .4.85 4.93
3. . E. labia - pupal period 8. S. gregaria - egg period
Y= 13.6ge - .001107 (97.3-t°F)2 Y 9.34e - .002397 (43-t°C)2
60.8°F 31.92 32.30 27.0°C 19.77 20.00
68.0°F 18.69 18.32 30.0°C 16.06 16.00
77.0°F 11.53 11.78 36.0'C 12,05 12.00
86.0°F 8.41 8.39 40.0'C 10.94 11.00
95.0°F 7.75 7.73
4. E. labia - life cycle 9. S. gregaria - nymphal period
Y 13.69 e- .0013 (93.3-t'F)2 Y = 4.783 e - .005747 (38.1-t°C)2
60.8'F 88.00 88.75 24.0'C 64.01 63.00
68.0'F 51.20 50.68 25.0°C 56.21 56.00
77.0'F 31.44 31.06 27.0'C 42.54 46.50
86.0'F 23.83 24.43 30.0'C 30.53 30.90
95.0°F 22.30 22.10 33.0°C 24.34 25.50
132 AGRIcuLtuRAL ENTOMOLOGY AND PEST CONTRoL
TABLE 8.4 (Contd.)

S. M. lefroyi - egg period 36.0'C 21.50 21.20
Y 183.7 e - .00872 (l03-tOF)2 37.0'C 2].}0 20.90
60.8 2 F 2.68 2.68 4O.0'C 21.39 20.70
68.0"F 1.64 1.63 44.0'C 25.60 27.00
77.0'F l.06 1.01 10. S. gregaria - life cycle
8G.O'F 0.71 0.71 y = 1.079ge - .004045 (40.5-t'CF
9S.0"F 0.58 0.58 27.0'C 97.00 97.40
30.0'C 72.30 71.10
36.0'C 50.20 50.80
40.0'C 46.30 46.00
Biophysical Processei involved in Development

The processes involved in organic development should be clear from
the presumptions on which Equation I is based, and the fact that the
values based on this equation are in excellent agreement with the values
observed by various previous workers renders it highly probable that the
presumptions are, if not quite correct interpretations of the natural pheno-
menon, at least good guides for solving this complicated phenomenon. It
will be noted that the following 3 fundamental hypotheses were made in
deducing Equation 1.
(i) Organic development takes place in accordance with the law of
compound i~terest, the speed of vital activity acting as rate of
interest.
(ii) The amount of development from birth to natural death as well
as the amount of development required to complete the well-
defined stages (egg, larva, pupa, etc.) are constant quantities for
the species, making allowance for individual variations.
(iii) The amount of vital activity performed in unit time increases
with rise of temperature, and with a velocity which does not
remain constant but is continuously being retarded with the
same rise of temperature. The rate of retardation, however,
has been supposed to be uniform.
In the first presumption there is nothing new; a number of previous
e~uations, e.g., principle of Q,1o of van't Hoff, p. of Arrhenius, the equa-
tion of catenary curve of Jamsch, etc., are all based on the same hypo-
thesis, although the hypothesis itself was probably based only on an
analogy of autocatalytic chemical reaction. The illustration of a deve-
loping .embryo given in Section II, however, shows that there is no need
of borrowing an analogy from the chemical field. Development does
actually take place in geometrical progression, i.e., in accordance with
compound interest law. at least in earlier stages. Blackman (1919) dis-
cussed this fact at length in regard to plant growth. In later stages of
EFFECT OF TEMPERATURE ON THE DYNAMICS OF INSECT DEVELOPMENT 133
development, although the geometrical progression becomes masked be-

cause of differentiation of cells which gradually lose their meristematic pro-
perties, it is plausible that the essential nature of development is not
altered. Development may not, and actually does not, mean only cell
division, but it is easily imaginable that it implies activities analogous to
the development and division of cells. The applicability of Equation 1 to
observed facts concerning advanced stages of development gives experi-
mental support to this assumption. ,
In the second assumption also there is nothing new. Even/ the sim-
plest of the relations, viz., the straight-line relationship betwe,en .tempe-
rature and developmental indices, presumes the total development to be
a constant quantity. Moreover, the use of a quantity such as develop-
mental index as' average velocity of development presupposes total deve-
lopment to be constant. Anyway, there appears to be no difference
of opinion on this point.
The third presumption is entirely new, and Equation 1 is absolutely
different from all other equations so far used to represent 'the relation-
ship under consideration. This presumption ~s actually a composite
of 2 basic factors. According to this hypothesis the speed of vital acti-
vity or, let us say for convenience of terminology, the amount of vital
activity performed in unit time, increases with, say, a particular velocity
due to rise of temperature, and that velocity itself remains being con-
tinuously retarded with same rise of temperature. Now, the same rise
of temperature cannot be expected to create a positive velocity, and a
negative acceleration (i.e. retardation) unless 2 opposi.te processes are
involved, both accelerated by rise of temperature, but having opposite
effects, one retarding the other. Therefore, if this third presumption is
correct, as it is found to be, then it can be explained only in one way,
i.e .• the amount of vital activity (r) perfonned in unit time increases with
rise of temperature with a particular velocity (u), but simultaneously
there is a sort of resistance against vital activity, and this resistance also
goes on increasing, and consequently retarding the velocity with which
(r) increases. Thus, because of this additional and opposite process of
resistance, the change in the value of (r) is like a retarded motion.
To sum up, the average velocity of development i.e., value of deve-
lopmental index, or reciprocal of developmental period (y or lIP), de-
pends on the value of (g), the absolute velocity of development at any
moment; then (g) itself depends on the value of (r), i.e.. the velocity of
vital activity; and ultimately, as shown by Equation, the value of (y)
directly depends on the value of (r) which itself at any particular tem-
perature depends on the value of (u). which is the velocity of change in
(r) with 1 degree rise of temperature; further, an additional factor of
resistance to vital activities comes into play creating retardation (a) in the
value of (u), and therefore the resultant velocity (v) of change in (r) at
any particular temperature depends on the value of (a). Ultimately, the
value of (y or lIP) of a particular species at any particular temperature
depends on the values of (u) and (a). Thus, if (g) is taken to represent
the velocity of development I, the first acceleration, namely, (r) acts like
the rate of compound interest, i.e., it increases the value of (g) just as
the rate of interest increases the value of capital put on compound inter-
est. Subsequent accelerations work in the usual way as acceleration
works in the case of moving bodies.
It is instructive to appreciate that the com~ination of 2 different
principles from the physico-chemical realm is not unexpected in the
complex phenomenon of temperature effect on rate of organic develop-
ment and growth. Organic development consists firstly of vital acti-
vities and secondly of accumulation of the products of those activities,
e.g., addition of cells, etc. The phenomenon of accumulation of the
products takes place as expected in accordance with the law of com-
pound interest with the first acceleration (r) acting as the rate of interest
- a phenomenon which has its analogue in autocatalytic chemical
reaction. That this does and should do so is evident from everyday
experience. The products of vital activities does not remain passive but
itself begins to show activity similar to those of which it is a result.
Thus, cells produced as a result of vital activities of previous cells do
not remain passive but produce further cells. This is the same thing
as an offspring not remaining passive but producing its own offspring.
It is similar also to the interest of a capital bearing its own interest.
i.e., the principle of compound interest.
An activity is always a result of force. Therefore; so far as the
increase or decrease in the speed of activity is concerned. it is but
natural that it should take place in accordance with the usual principles
governing force and motion. Temperature in the case of vital reactions
is equivalent to! the impressed force in the case of moving bodies. A
narticular temperature is equivalent to a particular impressed force
which should cause a particular acceleration because. according to the
established principles of f"rce and motion. a definite force produces a
fixed acceleration, other conditions remaining unaltered :
Force = rate of change of momentum
- rate of change of mass x velocity
= mass x rate of change of velocitv (mass being inyariable)
= ma~s x acceleration
(Note - This is copied from a ohysics book). Exoerimentally also we
find that corresponding to a fixed constant temoerature there is a fixed
value of (y); hence a fixed value of (g) and ultimat~ly a fixed value of
(r), i.e., a particular temperature produces a particular value of (r),
which represents acceleration of developmental velocity. But as the
temperature is raised it is equivalent to the increase in the amount of
impressed force which must change the value of acceleration. Thus
each degree of rise in temperature means increasing the force by a fixed
unit, which in turn must increase the value of acceleration by a fixed
unit whi~h we have represented by (u). Further, raising the tempera-
ture degree by degree means increasing the impressed force unit,' by unit,
which should mean accelerating the acceleration unit by unit, i.e.! uni-
formly; but an additional factor of resistance to vital activities comes into
play and actually: goes on retarding the secondary acceleration (u). Thus,
the effect of temperature on the rate of organic development may be
summed up by saying 'that the speed of vital activities is increased with
temperature as distance is increased with time in the case of retarded
motion, and this speed of vital activity increases the speed of develop-
ment as the rate of true compound interest increases the capital. I am
not aware of any other phenomenon in nature in which these 2 different
principles are combined as described above. There is no dearth of exam-
ples, wherein these principles act separately. Autocatalytic chemical
reaction has already been cited as following the principle of compound
ihterest. The principle of retarded motion is best illustrated by a bullet
which is shot upwards and continues to go upwards until the retardation
due to gravitational attraction becomes equal to the forward velocity of
the bullet, when it pauses a moment and then comes down.
Having understood aU the biophysical processes and the mathematical
relationships between them, it is worthwhile to summarize again how
temperature affects development. Since v=u-at (eq. 5), it is clear that
the value of (v) goes on decreasing as (t) i.e., the temperature is in-
creased; as long as (u) is greater than (a.t.) the value of (v) remains
positive, i.e., the value of (r) and hence the value of (g) and (y) remain
increasing; but when the temperature increases to such a value that
u=a.t., then the value of (v) becomes zero, i.e., there is no change in
the value of (r) and hence in the value of (g) and (y); further, when
the value of (t) rises still higher then (u) becomes less than (a.t.) and
the value of (v) becomes minus, i.e., the values of (r) and hence of (g)
and (y) begin to decrease with any further rise of temperature. Thus
the value of developmental index should go on rising, i.e., the develop-
mental period should go on decreasing up to a temperature when
u=a.t., i.e., when t =' u/a = T, and thence onwards the effect of tem-
perature should be opposite, i.e., the development should be quickest when
t=T. This is exactly what we find experimentally.
136 AGRICULTURAL ENTOMOLOGY AND PES}' CONTROL
These assumptions not only provide easy and accurate method for
calculating values of developmental periods corresponding to any constant
temperature or any range of fluctuating temperature. (Pradhan, 1945, 1946)
but, what is more important, they give an insight into the possible processes
involved in organic development and, for the first time, explain clearly why
the rise of temperature up to a certain degree accelerates development but
beyond that begins to have retardative effect.
Comparison with Previous Views

For full information on the comparative merits of the various attempts
made to establish relationship between temperature and organic develop-
ment, reference may be made to the papers of BOdenheimer (1926, 1938),
Zwolfer (1934), Shelford (1929), Wigglesworth (1942), and of Janisch (1932)
who has made (in a number of papers from 1925 onwards) the latest funda-
mental contribution to the subject. Botanists have also tried to solve the
problem, as will be seen from the reports of Leitch (1916) and Barten-
Wright (1933). In regard to the basic soundness and exactitude of agree-
ment with observed data, Janisch (1932) has made a systematic comparison
of the equation of his own catenary curve with different equations put for-
ward previous to him. Here it should suffice to point out in what ways the
present Equation I and the related concepts show a definite advance over
the knowledge accumulated up to the contribution of Janisch. He writes:
"Many efforts have been made to express these relations as a mathematical
formula. They are separable into two groups. The first of them is based on
velocity of reaction to chemical processes and represents an exponential
function of various types (van't Hoff's rule. Arrhenius' formula, the ex-
ponential law of Janisch). In the other group the product of temperature
and time expresses a constant. The formula taken as this relation is a
hyperbola (thermal summation. Belehradek's formula. Wardle's formula
and others). Naturally a formula must have a theoretical basis and must
agree with the data observed. It is then certain that an exponential formula
gives the best theoretical basis. since the chemical reactions as well as the
biological phenomena at least depend upon the law of mass action from
which the exponential formula (and therefore the exoonential law also)
follows automatically. However, the simple exponential curve does not
agree with all observed data or with the biological situation. By the resul-
tant of two crossing exponential curves described as catenary curve is
smootned to the obs~rved points and agrees without exception with the
biological situation."
Janisch has rightly formed 2 basic criteria for ~electing a valid formula.
Theyare:
(D sound theoretical basis, and
(ii) a satisfactory agreement with observed data.

Judged by these criteria the second group of attempts putting forward non-
exponential equations are naturally disposed of as unsatisfactory because
they neither have any theoretical basis nor give good agreement with the
observed facts. Of the exponential equations which are supposed to have
their theoretical basis in the law of mass action, Janisch has selected (on
the basis of criterion b) what he calls catenary curve - actually 2 in
number:
(1) Symmetrical catenary curve
m
t = ---- (a T + a -T )
.2
(2) Asymmetrical catenary curve
m'
t = ----
2 (a1 T '+ a
• 2
-'I )
wherein t==time, m=observed shortest time of development, a==constant,

and T = temperature.
Janisch also calls these curves "complex exponential as opposed to
simple one which has 'the formula t=m.a T" and on which other expon-
ential curves put forward prior to Janisch were based but proved to be
unsatisfactory when judged on the basis of criterion b. Now, it is to be
'noted that Janisch has modified the equation of simple exponential curve
into that of his own catenary curve only to satisfy the criterion b, i.e., in
order to get satisfactory agreement ~ith observed data: but by introducing
the modification without fairly sound theoretical basis he has again made
his own curve unsatisfactory from the viewpoint of criterion a. Coming
to Equation I put forward now, it will be found that, although this
equation also happens to be a modification of simple exponential equa-
tion, the modifications are based on more definite theoretical considera-
tions which render our conception of the whole developmental phenomenon
much clearer than before. Further, this equation does not have the defects
of the catenary curve; for example, the value of such fundamental con-
stants as the shortest period of development (m) and the temperature
corresponding to the same have to be experimentally determined by the
catenary curve. and this is a very difficult task. For example, Janisch
(1932) has taken for his illustration m==3.75 days corresponding to 29.6°C.
In order to find the exact value 29.6 one is bound to perform experiments
at very short intervals of temperature near about 29° and 30'C besides
the usual experiments to reach this neighbourhood. The task is all the more
tedious because (m) is the value of developmental period corresponding not
only to the temperature of Quickest development but also to the ootimum
temperature, and this also takes into consideration least mortality, least
138 AGRICULTIJRAL ENTOMOWGY AND PEST CONTROL
variation, etc. On the other hand, in the case of the present equation the va-
lues of Y and T, which represent quickest development and corresponding
temperature, are not to be determined experimentally but are to be calculat-
ed on the basis of all available data taken together (as shown in Appendix
A). Consequently, there is bound to be much less experimental error in y 0
of the present equation than in (m) of catenary curve. As regards taking the
optimum temperature instead of temperature of quickest development as
the origin even of catenary curve, I think it is unnecessarily confusing 2
different issues, viz., the effect of temperature on the velocity of develop-
ment, and the effect of temperature on mortality, variation, etc.
The question of agreement with the observed data has already been
dealt with in section III. where it has been shown how excellently the
present equation is applicable to the data collected by various previous
workers.
- Lastly, the practical advantages strongly' advocated by Bodenheimer
(1938) in favour of hyperbolic equation also need consideration in this
connection. According to this author, the three distinct advantages of
equilateral hyperbola are :
(1) The extremely simple way of computation. an improtant fact con-
sidering the mathematicophobia of most biologists.
(2) The easy way of getting the basic data for calculation and of im-
proving the curve by additional data.
(3) Last but not the least, only hyperbola furnishes us with the value
for lower developmental threshold, which value has proven to be
of highest importance in ~cological conception and calcula-
tions. This value is not obtained by any other formula". Owing to the
virtues enumemted by Bodenheimer, hyperboHc equation is
certainly today the best known and hence apparently also the most
exact relation between temperature and insect development. It is the-
refore necessary to emphasise that
(a) Because of exact similarity between Equation I and the equa-
tion of normal frequency distribution, for which most of the
necessary calculations have already been done and published for
ready reference. the calculations with Equation I are in fact very
simple and. as shown in a previous communication (Prac1han.
1945), easily handled after a little practice even without a tho-
rough understanding of the biological processes on which this
, equation is based;
(b) Exactly the same basic data are needed for Equation I of this
paper as for hyperbolic equation. i.e .. a number of observations
on developmental periods corresponding to different temoera-
tures;
EFFECT OF TEMPERATURE ON THE I?YNAMICS OF INSECT DEVELOPM~NT 139
(c) As stated else where (Pradhan, 1946). almost, all the useful appli-
cations of the hyperbolic equation mentioned by Bodenheimer
are possible with greater exactitude with the help of Equation
I and without taking the help of threshold of development, which
has no scientific theoretical basis;
(d) Besides retaining the 3 distinct advantages of hyperbolic equa-
tion. the present Equation I has rendered it possible (Pradhan.
1945) to fo~mulate an equation for calculating developmental
periods corresponding to variable temperatures of nature, whe-
reas all the previous equations are applicable, if at,' all, only
to constant temperature artificially maintained in the labQratory.
Long after wri,ting the fQregoing account I received some reprints kindly
sent by Professor Davidson of Waite Agricultural Research Institute.
University of Adelaid~, Australia, and another reprint kindly sent by Dr
C. B. Huffaker of Delaware, USA. The work of Huffaker (1944) does not
warrant any urgent discussion, but that of Davidson (1942, 1943) necessi-
tates the following remarks. Davidson has tried to fit to the temperature-
development data on insect eggs the famous Verhulst-Pearl logistic
curve which he, like many others, had previously used 'to describe the
trend of growth in animal populations'. This is certainly a step in the
expected though not altogether right direction. In fact it was surprising
that the so-well,talked-about logistic curve· did not attract attention when
so many far less likely empirical formulae have been tried by students of
temperature-development relation. The logistic curve, in all its essentials,
has been arrived at independently by chemists, biologists and mathema-
ticians, and all of them have found it useful and full of meaning; and it
is but natural that Davidson, being interested both in the growth of animal
population for which this curve was primarily developed and in the tem-
perature-development relation, should be impressed by the similarity
between the so-called sigmoid curves of both these phenomena. But, it
may be pointed out that the logistic curve is an integration or summation
curve, the differential of which is closely akin to the Gaussian curve of
error, and that the logistic curve describes the actual growth of an indi-
vidual or a population and not the rate of growth. Further, if the
logistic curve describes the actual growth, the rate of growth should
naturally be described by a curve akin to the differential of logistic curve;
and what we deal with in our study of temperature-development relation
is rate of growth and development and not actual growth and development.
Therefore Equation I, which is mathematically identical (although based
on different hypotheses) with the Gaussian curve of error, is a natural
outcome of the applicability of the logistic curve to the growth of an
individual or a popUlation. Davidson's attempt, on the other hand. to fit
140 AGRICULTURAL ENTOMOLOGY AND PEST CONTRo'L
the same logistic curve to the actual growth (of population) and to the
rate of growth or development is defective in principle, however close a fit
might be obtained within a particular range. This defect in principle/ also
exhibits itself in practice as the logistic curve becomes asymptotic to the
top horizontal line and generally does not come down, whereas the
temperature-development curve invariably comes down after the peak
temperature. Davidson has tried to overcome this criticism by doubting
the existence of smooth downward curve beyond the peak temperature and
by saying that because of the harmful effects of higher temperatures the
observed data for temperatures above the peak will be less reliable than
data for temperatures below the peak. This reasoning is certainly weak.
The descending of the curve beyond the peak is too universal to be ignored
or doubted. All the figures given by Davidson (1942) and by Davidson and
Swan (1943) give unmistakable evidence of this descending portion of the
curve. Thompson (1942) in his instructive review states: "We see that
there are always certain temperatures at which the rate is maximum; while
on either side of the optimum the rate falls off after the fashion of a
normal curve of error". In this statement we also see that Thompson did
recognise the similarity bteween temperature-development curve and the
normal curve but did not pursue the subject further.
In the end it may also, be pointed out that the logistic curve as applied
by Davidson also has ..the defect pointed out in the case! of catenary curve,
viz., that "peak temperature must be carefully determined by experiment".
and what this operation practically means has already been explained in
the' case of catenary curve.
Effects of V mabIe and Constant Temperatures

Most workers have made use of constant temperature maintained by
various devices within incubators. But how far are the results obtained
under constant temperatures of the incubators applicable to the field under
fluctuating temperatures. Some workers in the past (e.g. Shelford. 1927:
Peairs. 1927) had noted that variable temperatures bring about some
acceleration in insect development. Headlee (1940) invegti~ated this pro-
blem mote critically by maintaining fluctuating and constant temperatures
in parallel experiments, but he found acceleration in some series and
retardation in others. In trying to explain these apparently discordant
results he said that as the characteristic curve of reaction of insects to
external factors of insect environment happens to be sigmoid, the speed
of metabolism is likely to be reduced and more time required to comolete
the same changes in the constant if the variable ranl!e under examination
goes over into the foot curve or into the top curve. Shelford, too. had
observed the retardative effect of fluctuating temperature in some cases but
EFFECT OF TEMPERATUR£ ON THE DYNAMICS OF lNSECT DEVELOpMENT 141
could not make out its significance, and he listed the observations as ex-
ceptional data.
While trying to intercept the data on rearing various stages of. cotton
bollworm in the field insectary, I tried to investigate the relation bet,ween
temperature and insect development by taking the rearing of each batch of,
say, pupae pupating on a particular day as one complete experiment and
then finding out the average pupal period of that batch and the average
temperature of the period during which they lived in the pupal, stage. In
this way I obtained a good amount of data on pupal periods correspond-
ing to a good number of temperature averages extending over the full
range of seasonal. fluctuation. When I graphed the data wIth temperature
as abscissa and pupal period as ordinate, I could draw a curve resembling
the so-called hyperboli~ curve of development. When this curve was com-
pared with the curve obtained by Ahmad and Ghulamullah (1939) under
constant temperature it was seen that the curve obtained under fluctuating
temperature started from a much lower point (temperature) than that
under constant temperatures and then went on gradually approaching it
and going_ above it at higher temperatures. This comparison tends to indi-
cate that development is quicker under variable temperatures than under
a low constant temperature, but slower at higher temperatures.
Analysis of the problem on theoretical grounds revealed still more
~structively how and why development should be~icker under variable
temperatures below a particular range, and slower above it, when com-
pared with development under constant temperatures.
Taking the curve of developmental indices (1/ developmental periods)
to be sigmoid in shape, i.e. a curve with 2 opposite bends and a point of
inflexion between them, the effect of variable temperatures can be ex-
plained as follows. Suppose the average of fluctuating temperature is
X (Fig. 8.1) in the lower bend of the curve. If the temperature remains
constant at X, the rate of development will remain constant, but in the
case of fluctuating temperature it will be quicker when the temperature
goes above X, i.e., the effect will be accelerative; but when the temperature
goes below X the effect' will be retardative when compared with the constant
temperature at X, and the resultant effect of the fluctuating temperature
will be the sum of these accelerative and retardative effects. Suppose the
total effect of the fluctuating temperature is expressed as
Ex = (+ba x ) + (-br x ) = ba x -brl. ............... (1)
Ex =effect of fluctuating temperature with average at X;
b=range of fluctuation about X; ba x = accelerative effect due to tempe-
rature going above X up to b; br x = retardative effect due to tempera-
ture going below X up to b x .
Then, since the graph. in the region of the lower bend is bent upwards,
the rate of retardation below X is slower than acceleration above X, i.e.,

ba x br x .................. (2)
Therefore ba x - br x =a plus quantity ..... : ...... (3)
The resultant effect of the variable temperature is accelerative, i.e.,
round about X the development should be quicker under variable tempe-
rature than under constant temperature * .
Take another temperature (Z) in the upper bend. Here, too, the total
effect of the variable temperature will be
Ex =( + ba x )= bax- br x .................. (4)
Ex = effect of fluctuating temperature with average at z;
other symbols similar to I but round about Z.
But here in the region of the upper bend the graph is bent downwards;
therefore acceleration above Z is slower than retardation below Z, i.e.,
ba z br z .............................. (5)
Therefore
ba x- br z = a minimum quantity ..................... (6)
The total effect is retardative, i.e., development will be slower under
variable temperature than under constant temperature in the region of
the upper bend. Similarly, the effect of variable temperature should be
accelerative below the point of inflexion of the curve of developmental
indices and retardative:.above it, not that retardation is likely to occur only
when the range of variable temperature goes either in the top curve or into
the foot curve as stated by Headlee (1940). Practically, the effect should be
more pronounced in the region of the bends and less so in the intermediate
zone where the accelerative and retardative effects can nullify each other
to a great extent. Probably this is the first time that a clear conception of
the relative effects of variable and constant temperatures has been given.
The Curve of developmental Indices

In the foregoing consideration I have taken the curve of developmental
indices to be sigmoid, whereas the curves published by Ahmad and Ghu-
lamullah (1939), show a bend only on the upper side. the lower end being
produced straight. In trying to see if there can be a bend on the lower
side also or if the produced straight line is fully justified, I tried to fit
some other equations to the data of the above authors besides the hyper-
bolic equation which justifies the straight line on the lower side. I found
that the following equation fits the observed data better than the hyperbolic
curve and also gives a lower bend:
" These ideas can be more accurately expressed with the help of infinitesimal cal-
culus but this algebraic form is used for the sake of simplicity.
EFFECT OF TEMpERATURE ON THE DYNAMICS OF INSECT DEVELOPMENT 143
* y = Yoe -ax"................. ....... (7)

When the pupal periods were calculated for lower temperatures (not tried
by the above authors) by hyperbolic equation and also by Equation (7)
the values got by this equation and were found to be in quite good agr~ement
with the data observed by us for lower temperatures. whereas the values
given by the hyperbolic equation were much more exaggerated. Por exa.
mple. at 55.55'P the calculated pupal periods of Earias sp. should be 65.53
days according to Equation (7) and 229.4 days according to the .hyperbolic
equation. The observed pupal period for this average temperature in the
field is 51.77 days on an average. the maximum going up to 75.days. It
will be seen that 51.77 days under variable temperature is in , quite good
agreement with 65.53 days for constant lemperature, keeping in mind that
at these low temperatures the effect of the variable temperatures will be
accelerative. So, if temperatures below 16 C had been tried it is very likely
0
that the above authors, too, would have got a 'lower bend, as did many
other previous workers. In fact, Pruthi (1940) quoting the data of the same
authors gives a dotted bend besides the straight line on the lower side.
However, the point of major interest brought out by this effect at equation
fitting is that the applicability of Equation (7) may prove to be of great
significance, for it is the most important equation of biological statistics.
It appears that nobody has trieq this equation although a number of hyper.
'bolie, parabolic, and other exponential equations have been tried, some
proving better like Equation (7) than the hyperbolic equation, especially for
extrapolation purposes. The full significance has been given later when
equation (7) is dealt with.
Calculation of developmental Periods
After having a clear conception of the relative effects of variable and
constant temperatures as discussed before, it becomes possible even to
calculate the actual values of accelerative and retardative effects of variable
temperature on development under constant temperature observed in the
laboratory. If it is found, as in the case of E. labia (and probably in insects
in general, as indicated by the illustrations of a number of workers on the
subject), that Equation (7) is applicable to the curve of developmental
indices, the developmental periods under variable temperatures with known
.. y == highest value of Developmental Index (Reciprocal of Developmental Periods);

y :; value of Development Index corresponding to temperature t; t 0 == temperature
corresponding to Yo; X == (t 0 -t); a == constant; e == constant.
+This curve may be taken to be unnatural (as it was at first taken to be by
Mr. Krishna Iyer) because the curve of Fig. 8.1 has not been shown symmetrical
about t. Therellore it is to be pointed out that the other side of the curve does
exist and has actually been realized in practice in a number of species.
range of fluctuation can be calculated as follows.

It is clear that with the help of Equation (7) the value of developmental
index corresponding to any constant temperature can be easily calculated
after finding out. the values of Yo and the constant 'a' with the help of a
few experimentally determined values of developmental periods corres-
ponding to certain constant temperatures. In the case of variable tempera-
tures the values of y also have to remain fluctuating with temperature fluc-
tuation .Therefore, the average value of y has to be calculated in relation
to particular range of temperature fluctuation. This average value of y,
however, cannot be found out merely by taking the mean of the 2 extreme
values of y corresponding to maximum and minimum temperatures because
the rates of increase and decrease in the value of yare not constant,
especially in the lower and higher zones of temperatures.
x 2 -ax2
e dx........................ (8)
Therefore the value of integral Yo
i.e., the integral between the limits of the values of x corresponding to

maximum Xl and minimum X2 temperatures has to .be calculated giving
the total of all the values of y corresponding to every point in the whol6
range of temperature fluctuation, and then the value of this integral has
to be divided by the range of temperature fluctuation in order to get the
average value of y for the given range. The equation for this calculation
thus comes to be
= - - - - - - - -.................. (9)
Y (t1-t2 ) = average value of developmental index corresponding to tem-

perature fluctuation between t1 (max.) and t2 (min.) temperatures,
corresponding to which the values of X are Xl and X2' respec-
tively.
At first sight the above equation, especially the integral portion, will
appear, to be too complicated and tedious to gain general applicability and
popularity. Therefore, it has to be pointed out that Equation (7) is almost
identical with the equation of Normal Frequency Probability Curve (10),
which is the basic and most important equation of statistics and on which
so much work has already been done that calculations based on it have
become comparatively easy. So, the methods employed in calculating the
EFFECT OF TEMPERATURE ON THp DYNAMICS OF INSECT DEVELOPMENT 145
various frequencies (y) in equation (10) can be usefully adapted to find out
the values of developmental indices (y) in equation (7).
y=y o.e -ax 2 . . . . . . . . . . . . . . . . . . (7) Equation of the curve of developmental
indices
or
1 "(10) Equation of the norma,i frequ-
y= - - e - ency or probability cu~ve
)-2
Thus, from eqiIations (7) and (l0) it will be clear that if the numerical
values of x, y, and Yo are kept identical, then
1 or 0 =
a = 2.02 J~ .2a
Of _!_
0
= x
(tt~
And equation (7) can be written as :

y = 1 e _ax 2 (12a)
.J 1
2a
J-2-
or y - 0.3980 2a.e -ax 2 (12b)
Also
1 .x e _X2 = 0.3989 x e _ax 2 dx (13)
-2~dx v'2.a
y'-2-
We find that the values of the integral have been computed corresponding
to different values of x varying from 0.00 onwards at intervals of 0.01 and
published in what are' commonly known. as Sheppard's Tables. Therefore
just after calculating the value of ~ we can directly see from these
y'-I-
2a
tables the value of integral (13). Further the value of
X2 e- ax2dx ............ (14)
0.3989 y' -2-a-
can be found out by finding the value of integral (13) first for the value of
Xl and then for x2• and finally
y'-I-
2a
deducting the former from the latter. The value of integral (8) can be found
out by multiplying the value of integral (14) by Yo
0.3989
The comparative simplicity of the above calculation can be illustrated
by a definite numerical example. Suppose we want to calculate the average
pupal period of E. labia under a variable temperature of 55.82 ± 9.795, an~
also under the constant temperature of 55.82 ± 0 calculated with the help
of the data on pupal periods published by Ahmad and Ghulamullah
(1939) the values of developmental constants for E. labia are
Yo =13.64
to =95°C
a =0.001303 or
v' 1
2a 19.59
From the above example
tl = 55.82+9.795 = 65.615
t2 = 55.82-9.795 = 46.025
Therefore
Xl = 95-65.615 = 25.385
X2 = 95-46.025 = 48.975
Therefore
Xl = 29.385 .
--19~59- = 1.5 .................. (A)
Y 1
2a
X2 = 48.975
--19.59- = 2.5 .................. (B)
Y-l--
2a
From Sheppard's Tables we find as explained above that the value of
integral (13) = 0.4332 corresponding to (A)
= 0.4938 corresponding to (B)
the value of the average may remain unaltered.
In the end it needs to be mentioned that in the above calculations it
EFFECT OF TEMPERATURE ON TEtE DYNAMICS OF INSECT DEVELOPMENT 147
is implied that the fluctuation in temperature is quite regular whereas it is

not strictly true in nature; but this flaw. if at all is common to all correla-
tions between different phenomena and average temperatures.
APPENDIX A
For fitting equation Y =
AX2 + BX +C to the observations on pupal
periods of E. labia under different constant temperatures we proceed as
follows.
First we prepare the following table jn which
P = Developmental periods calculated from the data published by
Ahmad and Gulamullah (1939) after taking the average of their
averages of pup~l periods corresponding to three humidities for
each value of temperature.
X = Values of temperature in degrees Fahrenheit corresponding to
different values of P.
, 100.
Y = Log Y. where Y = --
P
Substituting the values obtained above in a set of normal equations we get
Ca) A (X4)+ B.r(X") + C (X2)= (X2y) or 206351005.2896A + 2489151.712B
+ 30670.64C = 28931.3807.
(b) A (X") + B 2' (X2) + C (X) = (XY) or 2489151.7120A + 30670.640B
+ 386.80C = 351.846.
(c) A (X2)+ B,E (X)NC=(Y) or 30670.6400A+386.800B+5.00C-4.36772
Therefore the value of integral (14) = 0.4938-0,4332
Therefore the value of integral (8) = 0.0606 (y 0)
------
(0.3989 v' 2 a )
0.0606 x 13.64 x 19.59
= ----=-0--=.39=8C::-- - -
9
= 40.58
40.58
Therefore yet 1-
t) -
2 - 65.616-46.025
= 2.072
Therefore pupil period = 100 =48.27 days under a variable temperature

2.072
of 55.82±0.795.
The pupal period at constant temperature can be found much more easily
with the help of Sheppard's Tables as they also contain the value of
1 e- X2 for different values of x. Thus for pupal period at 55.82 (constant)
----
- 20 2
0
y'2
..... 0
..... 0\ 00
"\C> r- 8M 00
oo "0v "000
"'! 0
'"0 00 "'! $:'
00 r-: N .....
-.....
'"
00
0
"<t
M
0
Vl
Vl
\C>
0\
t- o
V
C\I
,....,
'"0\00
M
"
~
'-'
:;1: N
0 0
N
0
\0 .,.,0 ..".
~ -
t- M \C> 00 0\
'".....'"00 00 M
N
1'1
'" 0
00
"..".on $:'
-
Vl Vl 00
..... r-: ~
C'\
N .,.,
0 t-
N
0\ 0
,....,
"'!
on
'-'
..... W
00
'"
0
0\
"<t
.~
".....
t-
\C>
00
00
N
0\
-
'"
'"
"~
.....
00
V
.....
....
...;
M
t-
t-
'-0
..... 8
0 0 0
-.i w
p...9
tI)
;.,
~
- ....
":'
1'1
'"
N
00
....
00
t-:
0\
C':!
00
'"
'"
r-: \C>
0\
00
M
vi ...
0
....
0
g
.,.,
..... W
'"
0\
§ 8 0
0
8 \C>
0
..
:><
00
N
r...: ..0
~ 8
..0
0
.,.;
N
'<I"
.....
11"1
"....
..... ~
M
.....
00
0
1'1
:g
\0 00 Vl 0 Vl
\0
'"
....""N ;;:; r--
'<I"
..".
.....
M Vl 00 N
....
r-:
.... R
....
-.,.,
0-
00
'-'
W
~ ~ 8o 80
N "<t
N
t-
X .,.; N
,., ..0 .,.;
Vl
t-
..... M
Vl
Vl
0 ...,
t-
'<I"
N ~
..... '"
Vl
'" .,.,
r-
N "<t '"
'" 00
"<t
II?
0 ~
'<I" 0
"
\0
.....
0
'-'
W
\0 8 0
0 8 0
.,.;
-
X 10 -.i C'\ 10
..,r-- ~
'"....
\0
N
~
~
11"1
0\
0\
00
..0
00
0 .,.,
00
..... 8
'"
00
\C>
"
t"- '"oo '" W
EFFECT OF TEMpERATURE ON THE DYNAMICS OF lNSECT DEVELOPMENT 149
x x =95-55.82
- - = - - = - - - - = 2.0
o V-I- 19.59
2a
For x = 2:0 we find in Sheppard's Table
a
0.3989 e _ax2 = 1 e- X2
~- = 0.0540
V 2
Therefore y =y o.e -ax 2 = 0.0540 x 13.64
I \ 0.3989 = 1.846
Therefore accelerativ~ value of

Variable Temperature of 55.82±9.795 = 54.16-48.27 = 5,89 days over
full period at 55.82 'C. constant.
From'these calculations it will be clear that it is every point in the
whole range of temperature fluctuation that determines the developmental
period under variable temperatures and not only the average. FOJ example,
the pupal period should be quite different if the raI1ge of fluctuation changes
although the value of average may remain 'unaltered.
,(d) 12445757.560A + 153353.2B + 1934.0C= 1759.22897
(multiplying (b) by 5).
(e) 11863403.552A + 149614.24B
, + 1934.0C= 1689.434096 ...... (c) x 386.8.
(fl 582355.008A + 3738.96B +0=69.7948749 ...... (d)- (e)
(g) 1031755026.4480A + 12445758.560B + 153353.2C = 144656.903626.(a)
x 5
(h) 940688158.0096A + 11863403.552B + 153353.2C= 133959.767741.(c)
I
- x 30670.64
(i) 91066868.4384A + 582355.008B + 0 = 10696.135885 ...... (g) - (h).
U) 339137355342.68A + 2177402080. 711B = 40645394.406629 ...... (f)
x 582355.008
(k). 34495378416.44A + 2177402080. 711B = 39992424.229327 ...... (i)
x 3788:96
(1) 1358023070.76A + 0 = 652970.177302 ...... U) - (k).
A = 0.0004808241.
Substituting
,
the value of A in (f) we have B = + 0.09355682
J
Substituting the values of A and B in (c) we have C = - 3.414575.
T~B= 0.09355682 = 97.3 or T = 97.3
2A 2x.000480241
log yo = log K = C- AT2 = 1.1364 or yo 13.69.
or !a = 0.001107.
-1 A
2a .4343
The equation thus comes to be : y= 13.69 e -O.OOl107(97.3-CFY.
APPENDIX B
The Doolittl,er procedure for fitting equation y = AX2 + BX + C to
the raw individ'ual observations on developmental periods under different
constant temperature was adapted from the book of Goulden (1939). The
following table was prepared from the observations on nymphal periods
of S. gregaria published by Hussain and Ahmad (1936) in their Tables
XIX and XXI. In the following table P=nymphal period in days,
t = temperature in degrees centigrade and numbers within the table repre-
sent the number of nymphs which reached the adult stage at each tempe-
rature. Thus, the table shows that at 24°C 3 nymphs reached the adult
stage in 62 days, 1 in 63 days and 2 in 64 days, and the total number
of successful observations (N xy ) at 24°C was 6.
From Table B.I we prepare the following one in which P is replaced
by Y when Y = log IOO-log P, i,e., logarithm of developmental index;
X = t.
TABLE B.1
P 24'e 2Soe 27°e 30 e

0
33°e WC 37'e 4O"C 44°C Nxy
19 days 1 2 3
20 days 3 2 5
21 days 10 3 12 25
22 days 1 7 9
23 days 1 1
24 days 2 2
26 days 8 S
27 days 5 6
30 days 2 2
31 days 13 13
45 days 1
46 days 2 7-
47 days 4 4
56 days, 1
62 days 3 3
63 days 1 1
64 days 2 2
Nxy 6 7 15 17 20 7 14 88
EFFECT OF TEMPERATURE ON THE ,DYNAMICS OF l~SECT DEVELOPMENT 151
TABLE B.2
Yx 24 25 27 30 33 36 37 40 44 Nyx
0.7212 1 2 3
0.6990 3 2 5
0.6772 10 3 12 25
0.6576 1 7 1 9
0.6383 1 1
0.6198 2 2
0.5850 8 8
0.5686 5 6
0.5229 2 2
0.5086 13 13
0.3468 1 1
0.3372 2. 2
0.3279 4 4
0.2518 1 1
0.2076 j 3
0.2007 1 1
0.1938 2 2
Nxy 6 7 15 17 20 7 14 '" 88
From Table B.2 we construct Table B.3 in which T )IX is calculated as

given for column of X = 24 as example :
0.2076 x 3 = 0.6228
0.2007 x 1 = 0.2007
0.1938 x 2 = 0.3876
T yx = 1.2111
Thus the value of T yx against 24 of column X comes to be 1.2111; other
l
columns are self-explanatory.
TABLE B.3
XTxz
24 6 1.2111 144 3456 82944 1990656 29.0664 679.5936

25 1 1.2518 25 625 15625 390625 6.2950 157.3750
27 7 2.3328 189 5103 137781 3720087 62.9856 1700.6112
30 15 7.6576 450 13500 405000 12150000 229.7280 689l.84oo
33 17 10.0585 561 18513 610929 20160657 33l.9305 10953.7065
36 20 13A722 720 25920 933120 33592320 484.9992 17459.9712
37 7 4.8084 259 9583 354571 13119127 177.9108 6582.6996
40 14 9.5688 560 22400 896000 35840800 382.7520 15310.0800
44 1 .5686 44 1936 85184 3748096 25.0184 1100.8096
88 49.9298 2952 101036 3521154 124711568 1730.6859 60854.6867
(N) (Y) (X) (X2) (Xt) (XY) (X2Y)
-
~
.....
00
-
:!
I
o
o
.... ~
N
C7\
on
N
§
o
00
00 - I
e
c::
-"o ._c::Q)
U...J
-
t--ooC7\O
-
o
EFFECT OF TEMPERATURE ON THE DYNAMICS OF INSECT DEVELoPMENT 153
From Table B.3 we continue the calculation in Table ~B.4 accor?in_g to the
instructions given therein. In these instructions. expressions like (5.1) mean
figures given in line 5 column 1 of Table B.4; expressions .like (1) mean
all the figures given in different columns of line 1.
A = -0.002247 -0.002247 (A) =0.002247 (11, K)
with sign change,d.
B == +0.17516 +0.17516 (B) == + 0.14741 (A) + 0.0275 (6, K) x (6, e)
with silln changed.
C = -2.7285 -2.7285 = -5.8758 (B) (2, i) +2.5799 (A) + 0i5674 (2, K)
x (2, 2) with sign changed,
Check -240.1344 +516.1107 -227.0481 ='49.9282
Y =;' - O.002247X2 + O.175173X - 2.7288

Further calculations as in Appendix A.
Analysis of Variance Test for Goodness of Fit is given in the main
body of chapter in Table 8.3.
1. g = speed of development, i.e., amount of growth (cell addition) Which,

however, goes on increasing in geometrical progression in succes·
sive units of time.
n = number of cell divisions in unit time.
The equation should actually be written as
g = Ken, but for simplicity we take K = I = a constant.
2. From Equation 2 we get (if P=deve!opmental period and G-total
development)
G = (e II ) P :; ell-P = constant by hypothesis.
_.. n.p. = constant because e = constant
or n x I, i.e., as the value of n increased the value of P decreased
P
3. Substituting the value of n from Equation 3 in Equation 2 we get
g = ee
G = (e er ) p :; constant, if P = developmental period
or 1 r
= ee
p
or 1 r
= log e e
Gp
or 1 log G=e r
p
or k = e r because G or log G = constant by hypothesis
.p
or y = e r if y = ~ where k = constant.
p
4. This hypothesis must not be confused with what Janisch (1932) has
written on pp. 150-158 applying the equation of retarded motion to
the course of development.
5. If a little change in the value of temperature is represented by dt .and
a corresponding change in the value of r by dr than
d 2 r = acceleration == -a by hypothesis
dt 2
or d 2r
-- =
dt2 -adt
or dr
(It = v == -at + u, where u is integration constant. .................... (5)
or dr dt
<It = -atdt _+ udt
or r = -iat2 +ut+c, where c is another integration constant ............ (6)
6. Starting with Equation 6
let u = T or u = aT, where T is a constant both u and a being
a constants.
and c + -laP = log K is another constant (this constant K is different
from K used in footnotes I and 3).
or c = log K-!aT2.
Substituting these values in Equation 6 we get
r=-iat 2 +Tt-taP+log K.
= -!a (F-2Tt + P) + log K.
= - ia (T-t)2 + log K.
= -iax 2 + log K if x == T-1 ............ (7)
7. Substituting the value of r from Equation 7 in Equation 4 we get
-!ax 2 _+ log K.
y=e
8. Let the value of x=o which is possible when t=T then,
-tax 2 •••••••••••• (8)
y=Ke o = K.
Therefore K is the value of y when t = T i.e., when x = 0, i.e., K == yo,
i.e., the value of y when temperature is equal to T.
9. Equation 9 which is the same as Equation 10 can be worked back
to Equation 4 and then substituting the value of r from Equation 6 into
Equation 4 we get.
y = e -iat2 + ut + c
or log y = - i at2 + ut + c.
or y = AX2 .+ BX + C
if Y = log y
A = -~a.
t = X
u = B.
10. The equations referred to are
A (X)i + B (X3) + C (X2) = (X2y)
A (X3) + B (X2) +c (X) = (XY)
A (X2) .+ B (X) + NC = (Y)
N being the number of observations and being sign of summation.
11. We find in thj.s case that variance due to an additional d'egree of
freedom used in fitting a third degree curve is also significant showing
that there may be some further gain in precision if a higher degree
equation issued. This point, however, is in no way against the hypothesis
of this pape_r. "
12. From Equation 7 we get
log y = log Yo --!ax 2
or log y Kl - AX2 ........... .if KI = log Yo and A =-!a
(parabolic relation)
or log y - KI - AX I ............ straight line relation 11
Xl = X2
13. To be more accurate, however, it may be mentioned here that the
total weight of evidence is likely to indicate later on that retardation also
is not uniform but goes on increasing slowly.
14. In this connection it must be pOinted out that the logistic curve has
been appli~d to the actual growth of an individual or a population plotted
against time. Therefore, the differential of the logistic curve which has
been recognized to be akin to the normal curve is ordinarily the curve
obtained by plotting the rate of growth against time, i.e., the age of the
individual or population. In the case of temperature-development curve,
on the other hand, the rate of development is plotted against temperature
and not against time. Therefore, the fact that the same type of curve is
obtained when the rate of development 'is plotted either against time or
against temperature is very instructive, indicating that probably time and
temperature are similar in their action on rate of development.
These findings are certainly of immense practical utility, and they are
likely to prove of much theoretical interest. From practical point of view
they present an elaboration of the proof in favour of a method reported
in an earlier communication of estimating the duration of life cycles and
developmental periods of insect stages under any constant temperature or
any ra!1ge of temperature fluctuation. From the theoretical point of view
it puts forward a much clearer conception than before of the phenomenon
of temperature effect on the rate of organic development. For the first time
a clear-cut explanation is given of why the rise of temperature up to a
certain degree accelerates development but beyond that value begins to
have retardative effect. The three main hypotheses formulated in this
connection are the following.
(1) Organic development takes place in accordance with the law of
cQll1Pound interest. the speed of vital activities acting as rate of
interest;
(2) The amount of development from birth to death and the amoup.t
of development required to complete the well-defined stages like egg.
larva. pupa. etc. are constant quantities for the species, making
allowance for individual variations;
(3) The amount of vital activity performed in unit time increases the
rise of temperature but with a velocity which does not remain con-
stant and remains continuously getting retarded with the same rise
of temperature. The rate of retardation, however, has been supposed
to be uniform. The ideas contained in the first two presumptions do
partly exist in literature. but the third presumption is entirely new.
The equations based on these hypotheses have been found to fit
satisfactorily 10 different sets of observations reported by others on
developmental periods of various stages of 3 species of insects be-
longing to 3 different orders.
Biometer
Temperature is the most important factor which determines the dura-
tion of the various stages in the insect's life cycle, and consequently the
number of generations during any period of time. Other factors, such as
humidity and light, cause only a small difference especially within their
non-limiting range. In the words of Bodenheimer (1938): "The duration
of life cycle depends on many environmental factors of which the chief
one is always temperature. Humidity, quantity and quality of accessible
food, etc. are other important factors, but for the normal life cycle of
most animals their importance is small as compared with that of tempera-
ture. Other factors, however. may need to be used for local corrections
of the pure temperature dependency." Recognizing this major importance
of temperature, phenologists have been trying for centuries to establish
such 'a correct relationships between temperature and development of
organisms as would enable estimations such as calculation of duration of
various insect stages corresponding to different temperatures. The oldest
and simplest assumption is that a linear relation exists between tempera-
ture and average velocity of development or, in other words. the time
required ~o complete the development at a particular temperature multi-
plied by the effective temperatu~e * gives. a constant product known as

the thermal constant for the species. Mathematically this relation is also
known as hyperbolic relation between temperature and developmental
periods. This assumption forms the basis of what is commonly known as
'temperature-sum-rule', 'summing of temperatures', thermal summation',
etc., according to which the daily effective temperature can be added from
day to day, and when the total reaches the value of thermal constant it
will indicate the completion of development. I
The idea of thermal summation is now 2 centuries old; and Reaumur
(1735), according to Shelford (1929), is commonly credited with. being
among the first to 'sum temperatures'. During this period, the basic
assumption of straight-line relationship has been proved over and over
again to be contrary to. facts, and a number of more accurate hypotheses
and methods of calculati9n have been put forward by various workers
but the following 2 practical disadvantages have persisted in all these
attempts:
(i) Somewhat as expected, the more accurate a method is, the more
complicated it is.
(ii) Almost all the later workers putting forward different hypotheses
have given methods only for calculation of average developmental
velocity corresponding to particular constant temperatures, but
they have not pointed out any means of estimating the develop-
mental periods under day-to-day and hour-to-hour temperature
fluctuation in nature, which 'thermal summation' at least aims at.
Therefore, partly because of what Bodenheimer calls 'mathematicophobia
of most biologists' and partly because of the tempting utility of the pro-
cess of thermal summation, the oldest hyperbolic assumption, despite its
constant deviation from observed facts, still strongly holds the practical
field of ecological investigation (Bodenheimer, 1924; Zwolfer, 1934).
Shelford (1929), reviewing the subject of 'animal in relation 'to tempera-
ture', comes to the conclusion: "The idea now centuries old to the effect
that temperature can be summed so as to give approximately the time of
development of a particular organism;, must be abandoned wherever
accurate results are desired. This method is likely to fail more or less
completely in the unusual seasons when accuracy is most desired". But
owing 10 the practical utility of thermal summation, Shelford had to
maintain the same basic idea although he had to introduce some ingeni-
ous but laborious corrections for deviation from straight-line relation-
ship.
" Effective temperature, according to definition, is equal to the actual temperature

minl,ls the lowest temperature c~lIed threshold at which development is perceptible.
In order to overcome the disadvantages of later methods and hypo-

theses pointed out above, I am trying to put forward a simple device for
adopting ordinary thermograph to what may aptly be called Biographs
and Biometers. These instruments not only reduce to the minimum the
difficulty of complicated calculations but also make it easy to combine the
basic idea of thermal summation with any of the more accurate hypotheses
and methods, so that there may remain no further necessity of sticking to
the erroneous assumption of hyperbolic relationship only for the sake of
a number of practical applications (enumerated by Bodenheimer, 1938),
which are mere corrollaries of thermal summation.
TABLE 8.5. SUMMARY OF CALCULATIONS FOR CONSTRUCTING

BIOGRAPH AND BIOMETER
t, i.e. (97.288-x) t, i.e. (97.288+x)

y +x t-30 .077 (t-30) t-30 .077 (t-30)
2 3 4 5 6 7 8
0.2 53.927 43.361 13.361 1.0287 151.215 121.215
0.4 47.775 49.531 19.531 1.5021 145.063 115.063
0.6 43.762 53.526 23.526 1.8117 140.050 110.050
0.8 40.690 56.598 26.598 2.0483 137.978 107.978 Not needed
1.0 38.134 59.154 29.154 2.2444 135.422 105.422
1.2 35.908 61.380 31.380 2.4161 133.196 103.196
1.4 33.987 63.30] 33.301 2.5639 131.275 101.275
1.6 32.085 65.203 35.203 2.7102 129.373 99.373 7.6509
1.8 30.381 66.907 36.907 2.84]2 127.669 97.669 7.5214
2.0 28.774 68.514 38.514 2.9648 126.062 96.062 7.3961
2.2 27.233 70.055 40.055 3.0839 124.521 94.521 7.2778
2.4 25.752 71.536 41.536 3.1989 123.040 93.040 7.1647
2.6 24.305 72.983 42.983 3.3097 121.593 91.593 7.0517
2.8 22.887 74.401 44.401 3.4190 120.175 90.175 6.9443
3.0 21.483 75.805 45.805 3.5293 118.7.71 88.771 6.8360
3.2 20.082 77.206 47.206 3.6350 117.370 87.370 6.7282
3.4 18.664 78.624 48.624 3.7437 115.952 85.952 6.6177
3.6 17.227 80.06] 50.061 3.8548 114.515 84.515 6.5088
3.8 15.747 81.541 51.541 3.9682 113.035 83.635 6.3944
4.0 14.]98 83.090 53.090 4.0879 111.486 81.486 6.2747
4.2 12.557 84.731 54.731 4.2141 109.845 79.845 6.1488
4.4 10.745 86.543 56.543 4.3531 108.033 78.033 6.0089
4.6 8.678 88.610 58.610 4.5134 105.966 75.966 5.8492
4.8 ' 6.084 91.204 61.204 4.7120 103.372 73.372 5.6494
5.0 0.000 97.288 67.288 5.1809 97.288 67.288 5.1809
Note - Y represents developmental index as in Eq. I; x also represents the same

~8 in that equation; t represents the temperature corresponding to Y and
.077 (t-30) represents the distance from the large line of the line repre-
senting t.
EFFECT OF TEMPERA TURF ON THE DYNAMICS OF INSECT DEVELOPMEN'T 159
Q.l
• 4 4
'20
110
if
'00 ""or
O
c:::::- :Ii::;
90
i!'
0
~~ '"':r"
I-
70
..z
+ , 60 g
~
-~ ~
0
="
'0
02 \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ I\
Ftg. 8.7. Biograph of the pupal period of 'Earias fabia.
Biograph
A biograph can be had merely by modifying the chart of a standard
thermograph so that the horizontal lines of the chart, instead of showing
different temperatures, may directly show the average velocity of deve-
lopment, corresponding to various temperatures' calculated according to
one of the more accurate methods. Thus, for constructing the biograph for
the pupal period of Earias labia represented in Fig. 8.7 the temperatures
corresponding to different values of developmental index (Table 8.5) at
intervals of 0-2 were calculated with the help of the following equation
(Pradhan, 1945).
-ax'
y = y oe_ ........................ (1)'"
When the constants of this equation were calculated by taking
Y = 100/P we got for pupal period of E. labia (calculations, based on
observations of Ahmed and Ghulamuli~h, 1939):
• y == value of developmental index, i.e., reciprocal of developmental period

corresponding to some particular temperature, say, t'; this reciprocal is, however,
multiplied by 100 in order to obtain convenient values. Therefore, y = IOO/p in
common calculations where P represents the developmental period, i.e., the time
required to complete the development of a particular stage.
y = highest possible value of developmental index species concerned. Let
the temperature corresponding to Yo be T.
±x = (T-t), i.e. t = T - )1:.
a =. a constant.
e == base of Naperian logarithm = 2.718282 (constant).
- .QOll06 (97.288-t)2 ........................ (la)

Y = 13.6g e '
i.e., the highest possible value of developmental index came to 13.69.
Therefore, for constructing the biograph we had to find out the'tempera-
tures corresponding to- different values of y at suitable intervals up to,
~3.~9: ~or this purpo~e the value 13.69 is not suitable because of its being
mdiVIsible by any sUItable interval say, .2 or .5, convenient width of bio-
graph chart.
However, an arbitrary change in the value of y 0 does not change the
value of other constants of the equation. Thus if we take y to be equal
to, say .5, the change will only mean that the value of y does not remain
equal to 100/P as originally taken but becomes (l00 IP x 5 I 13.69 =
36.52/P; i.e., the reciprocal of the developmental period is not multiplied
by ] 00 but by another convenient number 36.52 which does not mean any
deviation from the principle. Therefore in constructing the biograph v(\
was taken to be 5 and the values of ±, i.e. (97.288 - t) in Table 8.5,
Column 2, up to 5 were calculated with the equation.
y = 5 e -0.001106 (97.288-t)2 .................. ,(Ib)
Then from each value of x the 2 values of temperatures (Table 8.5)
corresponding to each value of y were directly obtained by addition and
substraction.
Further, this biograph was to be adapted to a thermograph which was
meant to record a range of temperature from 30 P to 130'P over a width
0
of 7.7cm. Thus, taking 30 P as the base line, each degree above 30 was
0
equivalent to 7.7/100, i.e. 0.077 cm. Therefore for graphing the lines
representing the temperatures (Table 8.5) first 30 w~s deducted from each
temperature and then the remainder was multiplied by. 0.077. The products
thus obtained gave the distance from the base line of the lines representing
the various temperatures (Table 8.5). ,Then a thermograph chart adapted
to a particular thermograph was made in which, instead of drawing the
horizontal lines at regular intervals representing full degrees, i.e., 42 44°
0,
etc. as is the usual practice lines representing the temperatures shown in

Table 8.5 i.e. 43.361 49.513°, 53.526°, etc. were drawn and labelled as
0
,
representing the values of developmental indices, i.e.. 2, .4, .6 etc., res-

pectively. When this chart is put on the drum of the same thermograph
instrument to which it is adapted, the pen, instead of marking the tem-
perature at different times, shows directly the values of developmental
If the biograph is to be used on a different thermograph instrument then its

total width will have to be proportionately altered so as to fit into the other in-
strument .. This can be done on Iv by photographic magnification or reduction and
does not need any fresh calculation or drawinl!. Of course for such photographic
adaptalions the drawings should not contain time lines
index corresponding to different points in the whole range of temperature

fluctuation. Thus the point 0, which would have shown on a thermograph
chart that the temperature at 10 a.m. on Tuesday was 84.7°F, shows on
this biograph that the average velocity of pupal development of E. labia
was 4.2 at 10 a.m. on Tuesday.
Estimation of developmental Periods

Besides showing directly the value of developmental index' at different
times, the chief value of the biograph described in the above lines ~s the fact
that it provides an easy and a rather mechanical method for estimating
the duration of different stages. It is clear that in the biograph the vertical
scale represents the average velocity of development (developmental
index), and the horizontal scale represents the time; therefore the area
below the temperature (here, velocity) tracing (a) in Fig. 8.8 can be
directly ,taken to give the product of developmental velocity and time,
i.e. tbe amount of development itself. Thus the area enc1o~ed has 2 con-
secutive midnight lines on the sides and between 2 consecutive thick
horizontal lines representing whole number value of developmental index,
e.g., between 1 and 2 D.L or between 2 and 3 D.L, represents the amount
of development completed in 1 day if the value of developmental index
. remains 1 D.I. throughout the 24 hours. Let us fix the amount of deve-
lopment represented by this area (a somewhat rectangular space enclosed
by comparatively thick lines in Fig. 8.8 and 8.7) as a unit which we may
call 'biodiem'. If the value of developmental index remains continuously
1 D.L the full pupal development will take 36.52 days.
Y = 36.52{P or P = 36.52 Y = 36.52 x I = 36.52 days.
2 I
~--DL-~
Fig. 8.8. Principle of biometer.

162 :AGRICULTURAL ENTOMOLOGY AND .PEST CONTROL
This means that the full pupal period is represented by 36.52 such
areas as we have termed biodiem. Further, we see from Fig. 8.7 and 8.8
that each biodiem consists of 60 small cells which may be called 'biocells'.
Thus, with the biograph represented in Fig. 8.7 and 8.8, we can say that
the pupal period of E. fabia is represented by 36.52 biodiems or 36
biodiems and about 31 biocells, or by 2,191 biocells.
Now, in orger to determine the duration of the pupal stage of E. fabia
starting on any particular day, it should be necessary only to count from
that day onwards the number of biodiems and biocells below the tempera-
ture tracing till the total comes to 36 biodiems and 31 biocells; the time
and day when this total is reached will be estimated time of emergence,
making all<)wance for individual variations. But a close examination of
the biograph will show that the value of developmental index rises with
temperature up to 5 D.I. and then falls as gradually as it rose. Therefore,
if the the temperature tracing (a) runs along the line of, say, 3 D.I. above
the line of 5 D.l., it shows the same value of developmental index as if
it were running along the line of, say, 3 D.I. above the line of 5 D.l.,
it shows the same value of developmental index as if it were running
along the line of 3 D.l. below the line of 5 D.L Therefore, as shown in
Fig. 8.8 if the temperature tracing runs above the line df 5 D.I., as shown
by the solid tracing (a), the area representing the amount of development
will actually be below the dotted line (b), and it is clear that the area below
the dotted line can be obtained by subtracting the area enclosed by the
temperature tracing below the line of 5 D.1. from the area enclosed by the
same tracing below the line, of 5 D.I. Therefore, the actual process of
determining the pupal period of E. fabia comes to be :
The following record of Daily Biothermic Value~ should be completed
as a matter of routine every week when the biograph chart is changed:
(i) Starting, say, on the 1st of the month, count the number of biodiems
and biocells below the temperature tracing (a), but from the base line
below up to the line of 5 D.1. above, and from midnight line before
the 1st to midnight line after the 1st cells marked by plus (+) in Fig.
8.8 and keep the total as plus quantity in column 2 of Table 8.6. For
example, in Fig. 8.8 this plus quantity comes to be 2 complete biodiems
and 136 biocells, i.e., 4 Bd, 16 Bc on Tuesday and 2 complete biodiems
and 135 biocells, i.e. 4 Bd, and 15 Bc on Wednesday.
(ii) Then count the number of biodiems and biocells enclosed by the
temperature tracing above the line of 5 D.I. and keep this total as
minus quantity in column 3 of Table 8.6. Thus, in Fig. 8.8 this minus
• 'For convenience of terminology the evecyday total number of Biodiems and

BioceJIs is being designated by the term 'Daily Bioth~rmic Value',
'"Cl
e
::l
"0
u
.....o
sCl o 0
--
o 0 o 0
o
on
N ~ ~ _ ~ ~ ~ ~ ® ~
N M ~ on on ~ N
~~ !N !M
TABLE 8.6 (Contd.)

Date Up-to-date Date Up-to-date Date Up-to-date Date Up-to-date
total total total total
Bd. Be. Bd. Be. Bd. Be. Bd. Be.
1-5-44 4 9 15-6-44 65 12 13-8-44 53 53 27-9-44 100 42
2-5-44 8 7 16-6-44 69 47 14-8-44 57 37 28-9-44 104 9
3-5-44 12 3 17-6-44 74 25 15-8-44 61 11 29-9-44 107 24
4-5-44 1(1 5 18-6-44 78 46 16-8-44 65 15 30-9-44 110 32
5-5-44 20 21 19-6-44 83 11 17-8-44 69 19
6-5-44 24 42 20-6-44 87 28 18-8-44 73 27 1-10-44 ·3 30
7-5-44 29 6 21-6-44 92 00 19-8-44 77 54 2-10-44 7 14
8-5-44 33 25 22-6-44 96 46 20-8-44 82 25 3-10-44 11 15
9-5-44 37 40 23-6-44 101 28 21-8-44 86 56 4-10-44 14 49
10-5-44 42 10 24-6-44 106 15 22-8-44 91 19 5-10-44 18 43·.
11-5-44 46 38 25-6-44 111 00 23-8-44 95 42 6-10-44 22 33
12-5-44 50 58 26-6-44 115 11 24-8-44 100 6 7-10-44 26 19
13-5-44 55 22 27-6-44 119 13 25-8-44 104 36 8-10-44 28 57
14-5-44 59 47 28-6-44 123 40 26-8-44 108 57 9-10-44 31 42
15-5-44 64 10 29-6-44 127 57 27-8-44 112 53 10-10-44 34 13
16-5-44 68 45 30-6-44 132 15 28-8-44 116 39 11-10-44 36 34
17-5-44 73 17 29-8-44 120 9 12-10-44 38 54
18-5-44 77 28 1-7-44 4 44 30-8-44 123 53 13-10-44 41 19
19-5-44 81 40 2-7-44 10 27 31-8-44 127 25 14-10-44 43 51
20-5-44 85 54 3-7-44 14 54 15-10-44 46 16
21-5-44 90 7 4-7-44 19 25 1-9-44 3 28 16-10-44 48 43
22-5-44 94 28 5-7-44 23 32 2-9-44 6 58 17-10-44 51 33
23-5-44 98 36 Data absent from 3-9-44 10 24 18-10-44 54 24
24-5-44 102 53 6th to 24th. henee 4-9-44 14 10 19-10-44 57 15
25-5-44 107 4 onwards up-to-date 5-9-44 17 55 20-10-44 59 25
26-5-44 111 20 total from 25th 6-9-44 21 40 21-10-44 61 39
27-5-44 115 33 25-7-44 3 48 7-9-44 25 29 22-10-44 64 4
28-5-44 119 54 26-7-44 8 10 8-9-44 29 19 23-10-44 66 35
29-5-44 124 42 27-7-44 12 40 9-9-44 33 15 24-10-44 69 5
30-5-44 129 16 28-7-44 16 51 10-9-44 37 18 25-10-44 71 29
31-5-44 133 58 29-7-44 20 39 11-9-44 41 18 26-10-44 74 10
30-7-44 24 54 12-9-44 45 6 27-10-44 76 40.
1-6-44 4 30 31-7-44 29 35 13-9-44 48 55 28-10-44 78 58
2-6-44 8 56 14-9-44 52 51 29-10-44 81 16
3-6-44 13 17 1-8-44 4 19 15-9-44 56 46 30-10-44 83 36
4-6-44 16 39 2-8-44 8 18 16-9-44 60 38 31-10-44 85 50
5-6-44 20 27 3-8-44 12 33 17-9-44 64 38
6-6-44 24 30 4-8-44 16 33 18-9-44 68 39 1-11-44 2 27
7-6-44 28 42 5-8-44 20 25 19-9-44 72 40 2-11-44 4 59
8-6-44 33 7 6-8-44 24 39 20-9-44 76 41 3-11-44 7 36
9-6-44 37 43 7-8-44 28 46 21-9-44 80 22 4-11-44 9 55
10-6-44 42 10 8-8-44 32 46 22-9-44 83 50 5-11-44 11 48
11-6-44 46 50 9-8-44 37 16 23-9-44 87 27 6-11-44 13 32
12-6-44 51 50 10-8-44 41 16 24-9-44 91 3 7-11-44 15 14
13-6-44 56 19 11-8-44 45 22 25-9-44 94 19 8-11-44 17 8
14-6-44 60 47 12-8-44 49 36 26-9-44 97 29 9-11-44 19 8
EFFECT OF TEMPERATURE ON THE DYNAMICS OF INSECT DEVELOPMEN.J: 165
TABLE 8.6 (Contd.)
Date Up-to-date Date Up-to·date Date Up-to-date Date Up-to-date

Bd. Be. Bd. Be. Bd. Be. Bd. Be.

10-11-44 21 6 24-12-44 27 10 Data from 5th to 22-3-45 46 30
11-11-44 23 6 25-12-44 27 54 12th absent; 6 Bd., 23-3-45 49 20
12-11-44 25 36 26-12-44 28 46 31 Be. are inter- 24-3-45 52 21
13-11-44 28 12 27-12-44 29 34 polated for these 25-3-45 55 29
14-11-44 30 3 28-12-44 30 34 dates, and added to 26-3-45 57 56
15-11-44 31 33 29-12-44 31 24 give total for 12th. 27-3-45 60 18
16-11-44 32 56 30-12-44 32 16 12-2-45 9 33 28-3-45 62 53
17-11-44 34 14 ,31-12-44 '32 50 13-2-45 10 42 29-3-45 65 36
18-11-44 35 30 14-2-45 11 44 30-3-45 68 35
19-11-44 36 51 1-1-~5 31 15-2-45 12 42 31-3-45 71 53
20-11-44 38 11 2-1-45 59 16-2-45 13 53
21-11-44 39 36 - 3-1-45 1 29 17-2-45 15' 12 1-4-45 2 7
22-11-44 40 46 4-1-45 1 53 18-2-45 16 15 2-4-45 4 12
23-11-44 42 4 5-1-45 2 15 19-2-45 17 23 3-4-4~ 8 13
24-11-44 43 24 6-1-45 2 33 20-2-45 18 39 4-4-45 8 22
25-11-44 44 35 7-1-45 2 55 21-2-45 19 45 5-4-45 11 7
26-11-44 45 39 8-1-45 3 19 22-2-45 21 6 6-4-45 14 36
27-11-44 46 48 9-1-45 3 44 23-2-45 22 22 7-4-45 17 37
28-11-44 47 51 10-1-45 4 2 24-2-45 23 52 8-4-45 20 11
29;11-44 49 26 1]-]-45 4 16 25-2-45 25 45 9-4-45 23 10
30-11-44 51 3 12-1-45 4 33 26-2-45 27 35 11-4-45 29 29
13-1-45 5 5 27-2-45 29 30 12-4-45 32 50
1-12-44 1 33 14-1-45 5 43 28-2-45 31 36 13-4-45 36 16
2-12-44 3 2 - 15-1-45 6 18 14-4-45 40 1
3-12-44 4 48 16-]-45 6 54 1-3-45 2 7 ]5-4-45 43 53
4-12-44 6 22 17-1-45 7 29 2-3-45 4 31 ]6-4-45 47 46
5-12-44 S I 18-1-45 8 13 3-3-45 6 20 17-4-45 52 15
6-12-44 9 33 19-1-45 9 Il 4-3-45 7 15 18-4-45 56 4
7-12-44 10 50 20-1-45 10 5 5-3-45 7 53 19-4-45 59 46
8-12-44 12 36 21-1-45 10 5S 6-3-45 8 41 2()-4-45 63 22
9-12-44 13 54 22-1-45 11 54 7-3-45 9 48 2]-4-45 66 8
10-12-44 15 10 23-1-45 12 53 8-3-45 11 8 22-4-45 70 15
11-12-44 16 00 24-1-45 13 52 9-3-45 12 40 23-4-45 74 20
12-12-44 16 49 25-1-45 14 50 10+45 14 28 24-4-45 78 17
13-12-44 17 47 26-1-45 15 49 11-3-45 ]6 22 25-4-45 81 21
14-12-44 18 27 27-1-45 16 39 12-3-45 18 18 26-4-45 84 59
15-12-44 19 3 28-1-45 17 37 13-3-45 20 37 27-4-45 88 33
16-12-44 19 43 29-1-45 18 14 14-3-45 23 26 28-4-45 92 13
17-12-44 20 29 30-1-45 18 56 15-3-45 26 14 29-4-45 95 58
18-n-44 21 17 31-1-45 19 36 16-3-45 29 21 30-4-45 99 S3
19-12-44 22 5 17-3-45 32 37
20-12-44 22 51 1-2-45 40 18-3-45 35 41 1-5-45 4 6
21-12-44 24 9 2-2-45 1 25 19-3-45 38 32 2-5-45 8 21
22-12-44 25 21 3-2-45 2 16 20-3-45 41 9 3-5-45 12 12
23-12-44 26 26 4-2-45 3 2 21-3-45 43 36 4-5-45 15 57
TABLE 8.6 (Contd.)

Date Up-to-date Date UJ.l-to-date Date Up-to-date Date Up-to-date -
Bd. Be. Bd. Be. Bd. Be. Bd. ,Be.
5-5-45 19 48 16-5-45 65 29 27-5-45 112 40 6-6-45 26 41
6-5-45 23' 49 17-5-45 70 00 28-5-45 117 29 7-6-45 31 19
7-5-45 28 9 18-5-45 73 55 29-5-45 121 51 8-6-45 35 56
8-5-45 32 23 19-5-45 78 4 30-5-45 126 21 9-6-45 40 7
9-5-45 36 18 20-5-45 82 20 31-5-45 130 40 10-6-45 44 15
10-5-45 40 16 21-5-45 87 12 11-6-45 48 16
11-5-45 44 11 22-5-45 91 37 1-6-45 4 29 12-6-45 52 2.1
12-5-45 48 23 23-5-45 95 56 2-6-45 8 47 13-6-45 56 50.
13-5-45 52 30 24-5-45 99 59 3-6-45 13 14 14-6-45 60 51
14-5-45 56 26 25-5-45 103 52 4-6-45 17 36 15-6-45· 65 12
15-5-45 60 51 26-5-45 108 2S 5-6-45 22 2
quantity comes to be nil biodiems and 15 biocells on Tuesday.

(iii) The algebraic sum of the above plus and minus quantities will
give the amount of development for the day, i.e. the Daily Biothermic
Value entered in column 4 of Table 8.6. Thus in Fig. 8.8 the Daily
Biothermic value comes to be 4 biodiems and 1 biocell on Tuesday.
(iv) From the next day onwards, besides repeating processes 1 to
3, keep in column 5 of Table 8.6, an up-to-date total of :Daily
Biothermic Values from the starting day onwards.
Having the above-mentioned records as shown in Table 7.2, suppose
we want to know the date of emregence of pupae pupating, say, on 17.4.44.
For knowing this we have simply to add 36 biodiems and 31 biocells to
the up-to-date total of Biothermic value just before pupation and we shall
get the up-to-date total of Biothermic value just before emergence (Table
8.7). Thus, for the example taken we have 39 Bd and 36 Bc plus 36 Bd and
31 Bc equal to 76 Bd and 7Bc. Now we have to follow down date-wise and'
see when the up-to-date total reaches 76 Bd and 7 Bc. In Table 8.6, we
find this total to be completing on 26.4.44; if we calculate up to the nearest
hour it will be 6 p.m., i.e., the estimated pupal period of pupae pupating
')n 17.4.44 comes to be 9.75 days, making allowance for individual varia-
tions. The actual emergence of the batch pupating on 17.4.44 did take
place on 26.4.44 and 27.4.44. Thus, in short, up-to-date total just before
pup~tion 36 Bd and 31 Be = up-to-date total just before emergence.
Estimation of Number of Generations

The .principle illustrated with the help of the biograph for pupal periods
is equally applicable to duration of full life-cycle, which being known,
the number of generations in a given period of time can be directly
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EFFECT OF TEMP£RATURE ON THE DYNAMICS OF INSECT DEVELOPMENI 175
calculated.
Biometer
Biometer is only a biograph chart drawn on a transparent plate of
celluloid or glass or printed as positive on a photographic film which can
be superimposed on the thermograph records (Fig. 8.8). By this device
the biographic method of estimation of developmental period can be applied
even to past thermographic records. Moreover, the same thermographic
records can be used to study a number of species by superimposip.g their
respective biometers in turn.
Comparison of esti~ated and observed Values

Table 8.7 gives a cO:q1parison of pupal periods determined experimen-
tally and estimated by the biometric method. The actual values were
determined by rearing pupae of E. tabia in a field' laboratory under con-
ditions as mitural as possible. The ,estimations were made with the help
of a biometer drawn on a celluioid sheet and superimposed on the
records of a thermograph which had been working near the rearing cages
in the field laboratory. Table 8.6 was completed first and then with its
help columns 2 and 3 of Table 8.7 were filled in.
\ It will be seen that the agreement is more than ordinarily expected,
especially in view of the numerous disturbing factors involved in nature.
Moreover, this is only a preliminary attempt to test the idea of biograph
and biometer with the help of data collected for a variety of purposes
during the course of general exploration of the problem of fluctuation of
insect population. Consequently, a number of possible causes of discre-
pancy have been present in the estimations given here. Therefore, it is all
the more gratifying that a fairly satisfactory agreement has been obtained
between theory and observation. Some of the possible causes of error not
avoided in the present estimations are as follows :
(i) The biograph is based on Equation 1, the constants of which were
calculated from the data on developmental periods obtained when the
larvae were throughout fed on bhin'di (okra) and not on cotton as
was done in the experimental rearings reported herein.
(ii) Equation I is based on data, averaged for 3 grades of staturation
deficiency, i.e., 0 S.D., 13 S.D. and 14 S.D., but no other correction
for changes in humidity was made in these estimations.
(iii) The biometer used was drawn with hand on a celluloid sheet and
cannot be taken to be quite free from errors of drawing, etc.
(iv) The thermograph instrument, the records of which were used, had
flot been running very accurately.
176 AGRICULTURAL ENTOMOLOGY AND PESl CONTROL
REFERENCES
Ahmed, T. and Gulamullah 1939. Indian J. Ent. 1 : 17-47.
Barton-Wright, E. C. 1933. Recent Adv(lnnes ill Plant Physio/lJgy , London. 254-59.
Blackman. V. H. 1919. Anll. Bot. 33 : 353-360.
Bodenheimer, F. S. 1924. Bull. Soc. Eur. Eg}pt : 149-157.
Bodenheimer, F. S. 1926. Z angew. Ent. 12 : 91-122.
Bodenheimer, F. S. 1938. Problems of Animal Ecology. Olt.ford Univ. Press, London:
29-33.
Davidson, J. 1942. Aust J. exp Bioi, med Sci. 20 : 233-3C).
Davidson, J. 1943. Med. J. Australia, June 12, 1943, 533.
Davidson, J. and Swan. D. C. 1943. Aust. J expo Bioi. mecl. Sci. 21: 107-10. .
Goulden, C. H. 1939. Methods of Statlsical Analysis. J. WilIey & Sons, N.Y. 223'-230.
Headlee, T. J. 1940. J ecoll. Ent. 33 : 361-364.
Huffaker, 'C. B. 1944. Ann. ent. Soc. Am. 37 : 1-27.
Hussain. M. A. and Ahmad, T. 1936. Indian I. agric. Sci. G : 188-262.
Janisch, E. 1932. Trans. R. em. Soc. London 80 : 137-168.
Leitch, I. 1916. Alln Bot. Londoll 30 : 25-42.
Peairs, L. M. 1927. Bull. West. Virinia Agric expo Stll 208 1-26.
Pradhan, S. 1945. Froe. natll. lnst. Sci India 11 (2) : 73-80.
Pradh,Ul, S. 1946. Prx. /iW,'.\'. INSI. Sci. India }2 (6): )[)1.
Pruthi, H. S. 1940. Proc lndian SCi. Congr. 11. 27 (II): 261-308.
Shelford, V. E. 1927. Bull. IU. nat. Hisl. SUYI'. : 16.
Shelford, V. E. 1929. Laboratory and Field Ecology. Williams & Williams Co., London
608 pp.
Wigglesworth, V. B. 1942. Principles of /1lSect PhysiologY. Chapman and Holl Ltd.,
London 827 pp.
Zwolfer, W. 1934. Z. angrew. Enl. 21 : 333-384.
CHAPTER 9
PRINCIPLES OF INSECT CONTROL
THE 3 main pillars on which a stable policy for insect control has to rest
are (i) prevention, (ii) co-operation, and (iii) economics.
Prevention
In the field of insects, history repeats itself twice a year in a ,<ountry
like India, once in winter and again in dry summer, when most of the
popUlation is wiped out. Mter each catastrophe the residual insect popu-
lation starts a new cycle of multiplication. Thus, there are 2 critical points
of time when preventive and curative measures are likely to prove the
cheapest and most effective; for all control measures, including the primi-
tive mechanical control by catch and kill and the sophisticated ones
like the sterile male technique, are best applied when the population den-
sity of the pest is at its lowest. Later on the control become~ increasingly
difficult and costly, because the insect population multiplies rapidly in
geometrical progression. Thus, for example, in the beginning of the
summer the sugarcane pest Pyrilla lays prominent egg-masses on green
leaves and these can be easily clipped and destroyed. If the field is left
unattended for a few weeks, the population may increase lOO-fold assum-
ing that a female lays 200 eggs. Further, if control is delayed for another
generation (a few weeks), the population will grow not 200 times but
10,000 times. Similar is the case with a number of other pests, such as
the red haky caterpillar, in the beginning of monsoon.
Co-operation
Co-operative action in pest control is essential for success. This is
50 because many insect pests move from field to field and migrate to long
distances. It is in the field of insect control that international co-operation
has also been found necessary, effectiv~, and useful for several decades.
In fact, international co-operation in locust control is one of the earliest
of such activities.
Economics
The soundness of all pest control recommendations has to be tested
on the anvil of economics. Of course, both long-range and short-range
benefits have to be taken into consideration. For example, in the case of
the summer paddy crop, although the pest infestation is at times too low
for control measures to prove remunerative, yet these are likely to reduce
the pest inoculum for the monsoon paddy, thus, considering the economics
of both the crops, pest control in summer paddy may prove quite remune-
rative. The same kind of considerations are relevant tq the control of
nematodes by costly soil fumigants which may keep the soil fairly free
from serious infestation for several years. The classification is as follows;
One would. like to classify unde~ definite headings the various methods
now employed in plant protection, but the task is rendered difficult not
so much by their complexity as by their interdependence. However, one
convenient classification is as follows :
Pest control
Natural
I
Artificial
\ I
Climatic Topo- Biotic Biotic
I I I I I
Cultural Mecha- Physical Chemical Biologi-
graphic (Para· (Diseases) nical cal
sites) etc.
Legislative
Natural Control. The first division into artificial and natural control
is generally not done. However, it is useful to keep it, and also to appre-
ciate fully that the value and magnitude of natural control are of a very
high order. The magnitude of environmental resistance has been discussed
elsewhere, and natural control is only another way of highlighting the same
phenomenon. Many a time the magnitude of natural control works bet-
ween 98% and almost 100%.
Cultural Control. Cultural control consists of introducing such minor
changes in the ordinary farm practices and farming machinery that serves
the double purpose of being useful both from the agricultural point of
view and from that of pest control. To control insects by cultural methods
it is necessary to understand with equal intimacy the life economy of the
insect and the cultural practices necessary to grow the crop. A control
that would be effective against one kind might be useless against a closely
related kind because of differences in habits. The operation to be effec-
tive gas also to be carried out at a proper stage of development. In fact,
it is a real test of entomological genius. Hence, it is very difficult to
formulate effective cultural control operations, but once research has
revealed an effective and practicable operation there is no doubt that it
will have many great advantages Qver Qth~r meth()d~ of control. It will
PRINCIPLES OF INSECT CONTROL 179
be the cheapest of all control measures; in fact, often it costs the farmer
nothing in time, money, or convenience. Again, with crops of high acreage
and low unit value, the cultural operation is only control measure that
can be employed profitably. Its greatest advantage is that it is generally
preventive and is effective before the damage has been actually caused.
Its disadvantage, however, is that it is indirect or intangible and the
farmer is not sure how effective his measure has been.
Mechanical Control. The only point to be emphasized isI that the
general prejudice against mechanical control for its being old a,nd out of
fashion is unscientific. The argument that it is impracticable is also not
quite correct. In ,many cir~umstances mechanical control is likely to be
the most convenient way to nip the pest infestation in the bud. It is
the most feasible metbod in certain places, e.g. kitchen gardens. At any
rate, it should always pay to give a good thought to mechanical control
before embarking upon other more impressive methods.
Biological Control. This has been dealt with in the chapter on Positive
Role of Insects.
Chemical Control., Chemical control of insect pests has assumed great
importance in recent years. The major part of current entomological lite-
rature deals with this subject, and papers on it are also appearing in
~hemical, agricultural, and other journals. The result is that it is impossible
to do full justice to the subject within a few pages.
Of the various methods of insect control, none brings about such
prompt and conspicuous relief as chemical control. This is the main cause
of its popularity; but a psychological point is also involved, as in the
case of human disease for which some medicine is prescribed.
Insect toxicology resembles' pharmacology except that the former is
concerned with lethal rather than sublethal effects of chemicals. A phar-
macologist is interested in the maximum dose which all individuals (man,
animals) can tolerate. Likewise, insect toxicologist is interested in (i) the
maximum dose which plants can tolerate without suffering any harm, and
(ii) the minimum dose which will be sufficent to kill all the pests. The effects
of chemical poisons on all living beings are essentially similar, though
variable in degree. Hence, it is practically impossible to have a chemical
which would be a deadly poison for some forms of life and completely
harmless to others. The difference is generally that of degree rather than
of kind, for both plants and their insect pests which are living organisms.
To make the best use of the difference in degree of the effect we have to lay
special stress on the study of what is known as dosage-mortality relation.
This relation is essentially represented by a sigmoid curve. For killing,
we are interested in the higher portion of the curve; but for saving, we
are interested in the lower portion. These dosage mortality curves are alike
in shape but start from different points on the abscissa. A good insecti-
cide is that chemical for which the sigmoid curve for insect pests starts
earliest, for useful insects suitably later, for the plant still later, and for
domestic animals and man latest. The farther apart these curves are, the
easier it is to apply the chemical for pest control without risk to useful
insects, the plants, domestic animals, and man. There are several corollaries
of this basic principle.
Specificity of poisons. There was a time when it was generally
assumed that if an insect species was resistant to 1 or 2 insecticides it
was resistant to most or all and, conversely, if an insecticide was toxic -to
1 or 2 insects it was assumed to be toxic to most insects. It was with
this idea that Campbell (1926) decided that an insect so easily handled
as the silkworm would provide an excellent indication of relative toxicity
of a series of compounds. But when Campbell's sandwich technique was
applied by Richardson to grasshoppers the relative toxicity of the com-
pounds was different: he found that, whereas acid lead arsenate and
sodium fluosilicate were of about equal toxicity to silkworm, the latter
was more than 19 times as toxic to Melanoplus as the former. It was thus
that the importance of specificity of toxic action was realized. Examples
of such specificity have since multiplied in number. Thus, we find that
BHC is 39.86 times as toxic as DDT to Bagrada eruciferarum, 170.9
times to Aulacophora fovieeollis, and 108.4 times to Cylas formiearius;
and its toxicity relative to' DDT is 0.37 for Mylaeems maculosus, and
0.38 for Drosieha mangilerae. Similarly, Aldrin is more toxic than Endrin
to Aulaeophora loveieollis (R.T. 119.9: 66.7) and less toxic than Endrin
to Cylas formiearius (R.T.58.1 :271.99). DDT is more toxic to Myloeerus
maeulosus than BHC. Aldrin, Dieldrin and Endrin, but practically useless
against Cylas lormiearius. DDT is altogether useless against locust hoppers
and adults. sO' much so that if Paul Miller had used locusts instead of
house flies the toxic properties of DDT would not have been discovered
and the history of pesticide development might have been different and
delayed.
Susceptibility and stage 01 development. Not only have different
species been found to behave differently to the same insecticide but now
it is also well known that the various stages of even the same species
exhibit wide variations in susceptibility to the same toxic compound. There
appears to be no dependable regularity in the relative resistance of the
stages'to different chemicals. Lindgreen and Shepard (1932) reported that
the adults of Tribolium confusum are killed with about qne-half the dosage
of carbon disulphide or one-fourth the dosage of Cloropicrin needed to
kill the eggs, but they are 9 times as resistant as the eggs to ethylene oxide.
1t has been shown by Pradhan and Bindra (1956) that the 2Ad. 3rdt 4th
PRINCIPLES OF INSECf CONTROL 181
and 5th stages of locust hoppers are 1.72 times. 4.19 times. 7.46 times and
15.26 times. respectively. as resistant as 1st stage hoppers. So many workers
have made general observations along these lines that it is possible to
generalize that younger nymphal ins tars of Acrididae are more susceptible
than older ones.
The problem oj phytotoxicity. A plant, like an animal, is a living
organism susceptible to poisoning. Hence. a chemical controller must have
full information about the relative susceptibility of different pl~nt species
to specific insecticides. In fact, when poisons like arsenicals are applied
to plants to kill insects on them, there is often only a slight, margin of
safety in favour ,Of the plant. This obviously necessitates a close co-opera-
tion between entomologist and plant physiologist. but very little work
has been done on this aspect.
Insect resistance to insecticides. The ratiqnal explanation for the
sigmoid r,elationship between log·dosage and percentage mortality is that
the sigmoid curve can be takep to be the result of integration of the
normal curve depicting the ordinary biological variation' in individual
susceptibility to insecticide. The whole phenomenon can be visualized as
follows. The different classes (Fig. 9.1) of an insect population require
different doses for their destruction, and the frequency of these classes
,forms. as expected, a normal curve. In practice, however, when the insect
population is treated with any particular dose (say, dose 6 in Fig. 9.1)
mortality occurs not only in the class for which that dose is necessary
but also in such other classes (from 0 up to 6) for which the required
dose is less. Thus, the mortality observed is not 9.02% on the normal
curve, but 27.43% on the sigmoid curve. This becomes all the clearer
in a class like 15. Although the percentage of insects requiring this dose
is only 0.97 (on normal curve), the percentage mortality observed is 98.21
on sigmoid curve. Hence, in actual practice a sigmoid relationship is
recorded between percentage mortality and log-dosage. This clear under-
standing is very helpful in analyzing the development of insect population
resistant to insecticide. When an insect population in the field is treated
with an insecticide dose which brings, about, say, 95% mortality, a refe-
rence \, to the normal curve will show that a 5 % population requiring a
higher dose of the insecticide survives. If the area treated is large enough
not to allow the survivors to mix with untreated population from the
surrounding areas and lose their separate identity, their progeny will
require a much higher dose of the 'insecticide for a 95 % mortality than
the original population. In other words. comparatively resistant individuals
get screened out when an insect population is treated with an insecticide.
and by repeated such screenings a resistant popUlation replaces the original
susceptible population. Another point which should be appreciated at this
182 AGRICULTuItAL ENTOMOLOGY ANO PEST CONTROL
~9.18
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o , I I 1 1 ' \ I \ I , I \ 1
L :::: I : : I t I :
2 3 4 5 6 7 8 9 10 1112 1314 15'
CLASSES REQUIRING DIFFERENT DOSAGE (ON LOG-SCALE)

FOR THEIR MORTALITY
Fig. 9.1. Reaction between insect population classes and insecticidal doses.
stage is that the resistant individuals which constituted a small fraction

of the original untreated population are not necessarily stronger in qua-
lities other than resistance to insecticide. In fact, these individuals have
a comparatively low survival value and this explains their being in a
minority. Various experimental results have shown enough proof of this
assumption (Bhatia and Pradhan, 1960). Therefore, if the insecticidal
application is stopped for a few generations the population is likely to
regain the ~usceptibi1ity of the original population.
PRINCIPLES OF iNSECT CONTROL 183
Terminology [or Insect Resistance to Insecticides

Following the original suggestions of Hoskin and Gordon (1956) the
terms 'vigour tolerance' and 'resistance' are becoming popular as signifying
different types of phenomena involving fundamentally different mechanisms.
It is considered that, whereas it is useful to make a distinction between
vigour on the one hand and continued breeding from survivors on the
other, both of which might bring about a change in insects' abil,ty to with-
stand a toxicant, the distinction between tolerance and resistance is rather
artificial and unhelpful for a correct understanding of the complex phe-
nomenon of in~ect resistance to insecticides. Hence, the following scheme
of classification of the various types of insect resistance to insecticides is
in favour.
Resistance
I
Genotypic Resistance Phenotypic Resistance

or or
Genetic Resistance Vigour Resistance
II
External Resistance Internal Resistance
(to the entry of poison (to the poison that has
into the system) gained entry)
III
Behaviouristic Biophysical Biochemical Physiological

Resistance Resistance Resistance Resistance
This means that all the cases of resistance are to be classified as indicat-
ed above on the basis of 3 different criteria: (i) Origin of resistance, which
may be due to change in genetic constitution or merely vigour brought
about by such factors as temperature, food, humidity, etc., (ii) gross loca-
tion of the mechanism of resistance~ and (iii) nature of the mechanism
of resistance.
It is held that the terms 'susceptibility', 'tolerance', and 'resistance'
should be taken to indicate differences in degree, not in kind, and to
represent different regions on the same scale just as 'cool', 'warm' and
'hot' indicate different regions on the temperature scale.
The considerations on which the above-mentioned terminology has been
worked out are as follows. The general tendency is to reserve the term
'resistance' for those cases in which the 'added ability to withstand an
insecticide' has been acquired by breeding from survivors of a particular
insecticidal pressure over the generations. If this reservation gets fixed,

it will mean that when one insect population requires a significantly higher
LD so ' or suffers a lower mortality than the other, we cannot say that it
is more resistant than the other. In other words, we shall have to make
use of terms such as 'tolerant'~ 'susceptible', etc. The following conside-
rations will show that this practice of using different terms for different
types of funqainentally the same phenomenon is likely to lead to more
confusion than clarity.
1. The basic mechanism of resistance and what may be called tole-
rance can be exactly the same.
(i) Suppose there are 2 natural populations A and B which have not
been subjected to any insecticidal pressure, and that LD 50 of A is signi-
ficantly higher than that of B. If a portion of B is subjected to insecticidal
pressure till its LD 50 approaches that of A, it will lead tOI a situation
where we have 3 populations A, Band Bi in which the LD 50 values of
A and Bi are equal, and higher than that of B. According to the termi-
nology that will develop we shall have to say t}1at A is more tolerant
than B, and Bi is more resistant than B. These apparently different state-
ments will lead the reader to think that the differences between B and A
on the one hand, and between Band B, on the other. are basically different,
although actually this may not be the case. Further. it will be diffic~lt
to decide if A and B, are equally resistant if the terms tolerance and
resistance are used indifferently.
(ii) The mechanism of tolerance (i.e., resistance of a population not
subjected to insecticidal pressure) need not necessarily be different from
the mechanism of resistance in a population subjected to insecticidal
pressure, especially when no mutagenic changes are envisaged as a result
of insecticidal application.
(iii) Let us take one possible mechanism of resistance in which the
body fat may be storing DDT away from the site of action, i.e. there
may be individuals with a different genetic constitution by virtue of which
they have a larger amount of body fat which can store a larger quantity
of DDT away from the site of action. These individuals might be selected
out by the pressure of DDT application and constitute a resistant popu-
lation (X). In another population (Y) not subjected to this insecticidal
pressure there may be a larger amount of body fat as a result of better
nutrition, and this population may also show a degree of resistance equal
to population X. Yet another population (Z). neither subjected to in-
secticidal pressure nor having been on better nutrition. may show resis-
tance equal to X and Y simply because it is a higher temperature at
which the body fat is capable of storing a greater amount of DDT away
from the site of action. According to the terminology which is coming in
vogue, only population X can be called resistant and population Y and Z

should be regarded as showing vigour tolerance, although the mechanism
Js too similar to permit labelling of the 3 populations with entirely
different terms.
2. The term resistance is more appropriate than tolerance or suscep-
tibility. Resistance is positive, susceptibility is negative, and tolerance
indicates passiveness. Insects, like other living beings, try to resis~ all obst-
ructions to the process of living. Thus, 'tolerance' can only impl~ the limit
to which the living organism can successfully resist this obstruction with-
out feeling much pressure, and 'susceptibility' can imply only the negation
of successfull r~sistance, i~e. no positive phenomenon. Hence, the term
'resistance' should be preferred to other terms and should not be confined
to that type of situation where the LD 50 has increased as a result of
insecticidal pressure.
3. It is better to distinguish different types of resistance ,than to have
entirely different terms for each type. Geneticists will agree that all bio-
logical characters are (a) genotypic or (b) phenotypic. Accordingly, resis-
tance as a biological character should be differentiated into (a) geno-
typic resistance and (b) phenotypic resistance. All cases in which the
value of LD 1'0 increases as a result of selection pressure over the gene-
.rations are of genotypic resistance, and those in which LD 50 significantly
increases because of factors such as increased vigour, feeding or starva-
tion, temperature, humidity, etc. are of phenotypic resistance. In other
words, phenotypic resistance is almost equivalent to 'vigour tolerance'.
Resistance can also be differentiated, as has been done by Pradhan and
Bhatia (1951), on the basis of the site where it is offered, e.g., external
resistance to the entry of the poison into the system, and internal resistance
to the poison that actually enters the system. Each of these 2 types of
resistance may be further differentiated according to the nature of the
resistance mechanism into (i) behaviouristic resistance, (ii) biochemical
resistance, (iii) biophysical resistance, and (iv) physiological resistance. It
may be pointed out that all these dtfferent resistance mechanisms can
result in significant increase both in genotypic resistance by selection
pressure and in phenotypic resistance from changes in food and environ-
ment.
4. There are other dangers in locking up the word 'resistance' for the
situation in which the population has undergone change as a result of
selection pressure. By giving an impression that tolerance, susceptibility
and resistance are fundamentally different phenomena it will be rather
difficult to apply conclusions reached in one field to other fields. I have
already at times come across young workers saying that this was the
mechanism involved in tolerance and it could not apply to resistance,
and so on. Different terms will confuse the administrators and financiers
all the more. For example, the mechanism of resistance can be studied
by 2 approaches: (a) We can bring about genotypic change, say, in LD 50
and develop 2 strains, 1 more resistant than the other. Then we can study
whether the penetration pick"up etc. are less or the detoxification, excre-
tion etc. more in the resistant strain. (b) Similarly, we can bring about
phenotypic change in LD 50 by keeping one set at higher temperature
and other at lower temperature, and then determine if the higher or lower
LD 50 at higher temperature is due to higher excretion or detoxification
at higher temperature or due to lower penetration, pick-up etc. at lower
temperature. Both these approaches may give almost similar answers. but
if the word resistance is confined to 'a' above, the work on 'b' may'
receive a setback at least from the point of view of studying the mecha-
nism of resistance.
The only serious argument in favour of reserving the term resistance
for genotypic resistance is that it is this that has created so much con-
cern in the world about the resistance problem in general. This fact does
justify distinction between genotypic resistance and all types of pheno-
typic resistance which are rather temporary and can be called vigour
resistance; so that the public may not be unnecessarily scared by vigour
resistance and the young researcher may not be misled to treat the 2 as
necessarily involving different mechanisms.
Bioassay of Relative Toxicity of Insecticides to Different Species

The LC 50 values of various insecticides for each insect species are
being recorded so that when the tests are repeated after a few years the
development of resistance, if any, can be accurately judged. The infor-
mation'so far collected is published in short instalments in various journals.
These contributions give the values of relative toxicity of insecticides,
taking the toxicity of DDT as 1. Thtr data can be used for several pur-
poses besides detecting changes in the resistance of the insect species.
Analysis of External and Internal Components of Resistance

It has been demonstrated (Pradhan and Bhatia, 1951) in the case
of fumigation with HCN that insect mortality depends on the insect's
external resistance to the entry of the fumigant as well as its internal
resistance to the fumigant that has gained entry into its system, and
that these 2 components of resistance differ markedly in different species.
In studies on the susceptibility of Corcyra cepha/onica larvae, Tribolium
castaneum adults. and adult females of Drosicha sp. it was found that,
when the dosage mortality curves were prepared on the assumption that
the concentration of HCN to which the insects were exposed is the effec-
100
0~--0 C. Cephalonica
)( J( ItT. Castaneum
• • Drosicha sp.
80
"f
~
0
60
>-
.-=
-
(ij
(5
:;E
40
20
o 0.5 1.0 1.5 2.0 2.5 3,0 3.5

Hen. Conc. mgjLitre
Fig. 9.2. Comparative susceptibility of Carcyra cepha/onica, Tribo/;uin caslaneum

and Droslcha sp. to HCN.
100
80,
..
C
.. 60
Q)
tJ
Q)
Il..
e e C. Cephalonica
>- W It T. Castaneum
"
-..
~
III
0
~
40
Drosicha sp.
20
o 2 4 6 8 10 12
H C N Recovery mgj100gm of Insects
Fig. 9.3. Dosage mortality curves based on recovery of HCN from CoTt;),ra
cepha/onica, Tribolium castaneum and Drosicha sp.
tive dose, the resistance of these 3 species was in the order: C. cephalonica,
T. castaneum, Drosicha sp. (Fig. 9.2). But when the dosage mortality
graphs were prepared by taking the amount of HeN recovered per gram
body weight as an index of internal dose, the order of resistance of these
species was just the reverse (Fig. 9.3). These apparently anomalous findings
were explained by assuming that the resistance shown by an insect in an
actual fumigation operation, i.e., to the concentration of HeN to which
it is exposed (external dose), is what may be called total effective resistance
(Fig. 9.1) and that this is the resultant of (a) surface resistance i.e. external
resistance (Fig. 9.4) and (b) internal resistance (Fig. 9.3). Thus, the effective
resistance and high internal resistance giving a very low effective resistance,
as in the case of C. cephalonica, or vice versa giving the maximum resis-
tance as in the case of Drosicha sp.
EpicuticuIar Structure and Insect Resistance

The presence of wax or cement as the outermost layer of the epicuticle
12
0 - - - C. Cephalonica
10 )( )( T. Castaneum
-
CII
u
Q)
CII
C
...--+. Orosicha sp.
-
o
E
8
C)
o 6
o
.-
---E
C)
4
z
U
J:
-o 2
>-
8
Q)
>
0
-.--------~;--------------~.---.
£. 0 0.5 1.0 1.5 2.0 2.5 3.0 3.5
HCN Concentration mg/Litre
Fig. 9.4. Relationship between the concentration and recovery of HeN from
Corcyra cepha/onica, Tribolium castaneum and Drosiclw sp.
was determined by using Wigglesworth's technique in 8 species of insects,

and bioassay studies were carried out to determine the relative resistance
of 6 of these species to 3 different inert dusts, viz, quartz, magnesite, and
ro~k phosphate. The order of resistance was: Aphis craccivora < Bagrada
cruciferarum < Bruchus anaUs < Euproctis lunata < Corcyra cephaloll!ca <
Trogoderma grallarium. A general correlation was found between the areas
abraded by the inert dust and those where polyphenol got exposed by
chloroform treatment. The abrasion was also generally found in areas
which are likely to get rubbed. There was much less abrasion in individuals
that survived the dust treatment than in those that succumbed to it. There
was also a general correlation between (a) susceptibility to inert dust
treatment and (q-) the degree of abrasion caused, loss of booy weight,
presence or absence of wax and cement layers, and nature of the wax
layer.
Lipoid Solubility and Insect Resistance

The contact insecticide has tp penetrate the insect body. wall before
it can produce its effect. and it is reasonable to surmise that the first
step in this process is its dissolution in the epicuticular wax. Experimental
proof of this became possible when a sample technique was developed
for studying the solubility of insecticide particles in the wax covering
of the insect body. It was shown in the case of Trogoderma granarium
larvae and Euproctis lunata larvae that the relative solubility of DDT
crystals in the cuticular wax of these 2 species was negatively correlated
with their relative resistance to DDT suspensions.
Effect of Particle Size on Toxicity of Suspensions

That particle size has a profound effect on the efficacy of insecticidal
suspension has been recognized by various workers, but the exact manner
in which this effect is brought about is not well understood. It has been
shown that toxicity of DDT suspensions used as spray increases with
decrease in particle size up to a certain stage. but further decrease in size
decreases the toxic effect. Both these, effects have been reported sepa-
rately as contradiction of each other. but we have observed them together
and without any anomaly. It has been suggested as an explanation that
this relationship between toxicity and particle size is governed by 2
factors, viz, (a) the contact factor, on account of which smaller particles
prove more toxic because they (i) offer larger surface area for contact
with the insect body, and (ii) fit better into the contour of the body
surface; and (b) the drain-off factor. because the smaller particles remain
suspended longer in droplets of the carrier medium and are more liable
to be_ drain~(l off from the insect body than larger effective particles which
quickly settle down on the bony. The total effective toxicity is the resultant
of these 2 factors.
Fa~totS assudated with Higher Resistance in Higher Instars

This aspect has been studied in the case of different instars of the
desert locust (Schistocerca gregaria) in which resistance to gamma BHC
markedly increases from instar to instar. Of a dozen factors studied, only
surface area per unit body weight, percentage of wax in the exuviae" and
probable thickness of the exuviae (which is aL~o probably an index' of
the insecticide within the insect system) do actually vary in the expecte'd
direction (Fig. 9.5). The first 2 are negatively correlated with MLC values'
for different instars; the third shows a positive correlation. When allowance
is made for these factors, it is found that inherent resistance is actually
more in the case of younger instars. Moreover, it has been shown in the
case of Trogoderma granarium that the extraordinary resistance of its
larval stage is altogether lost in pupal and adult stages.
Efted of Temperature on Insect Resistance
The effect of temperature on insect mortality due to insecticides has
lOCUST INSTARS v
I II III IV
MLC OF GAMA SHe
(RESISTANCE)
BODY WEIGHT
PER UNll AR'EA
PERCENTAGE OF WAX
THICKNESS OF EXUVIAE
Fig. 9.5, Factors tesp0t)1\ibk for im;rease il\ resl,tal1l:e to \I\,ectidde in differell.\
instars of the des~rt locust. '
PRINCIPLES PF INSECT CONTROL 191
been recognized by various workers, but different effects have been

reported in different cases. In the case of contact-and stomach-poisons
the findings in general have been that there is higher mortality at lower
temperature, but in the case of fumigants the reverse effect has been
reported in most cases. It has been shown that a large number Qf these
conflicting observations can be accounted for by the following generali~
zations:
(i) Insect resistance to poisons changes with temperature, as do its other
vital activities, increasing up a critical,degree and afterwards peclining.
(ii) The amount of poison reaching the site of action in unit timeyaries
with temper1j.ture, generally but not always increasing with its rise.
(iii) The appars:nt effect of temperature on insecticidal action is the
resultant of these, 2 factors, viz., resistance and uptake.
In view of these findings it has been concluded that for studies on
insect resistance temperature has to be differentiated into 3 separate
components: (i) temperature during insecticidal application (treatment
temperature) which increases the uptake of the insecticide (Fig. 9.6);
(ii) temperature after the insecticid~l application (post-treatment tempera-
ture) which increases up to a certain degree the physiological resistance
of the insect to the insecticide (Fig. 9.7); and (iii) temperature before the
ipsecticidal application (pre-treatment temperature) which increases both
100 ".__....._--4• ..__....-~•. _ -...

O~ 90°F
_,_,
;;2
80
60
40
/ 80°F
~
I ~ • , 20 <:>
o
...
N ,....
('I') oo:t I.C) co ...,....,.... (XI
,....
.......
C') 0') I.C)
0') co
,.... en 0')
'<t
'<t
co
I.C) en N
,...:
IN 1M IN ,...; I"'; ,..,.:.
LOG. CONCENTRATION
Fig. 9.6. Relationship between the treatment temperature and the insecticidal
uptake.
100
80
60
40
20
o
N ('t)
,.... o:t In <0 ,.... co 0) 0
.- M 0) In .-
,.... ,.... M 0
0) CD
..... o:t .- 0) o:t N 0
10 0) .- N ~ tD co 0
IN IN IN I~ 1.- 1'- 1- 1.- 0
lOG. CONCENTRATION
Fig. 9.7. Relationship between the post-treatment temperature and the insecti-
cidal uptake.
320
-
w
0
z
280
240
DDT SUSPENSIONS
<
I-
(J) 200
(i5
w
-..
a: 160
0 120
.-
X 80
0
10
0 40
.J
14±1.6 24±1 30 35 40
TEMPERATURE °c
Fig. 9.8. Effect of temperature on the insect resistance to insecticides.
the physiological resistance to, and the uptake of, the insecticide. It has
been further shown that atleast in some cases the change in insect resis-
tance to insecticide with change in temperature follows essentially the
same curve (Fig. 9.8) as that obtained with temperature effects on any
other physiological activity, implying thereby a similarity between the
insect's resistance to insecticides and its other physiological activities. It
is possible that the post-treatment increase in temperature increases the
metabolism of the insecticide inside the insect system; if this metabolism
leads tOi non-toxic or less toxic metabolites the insect's resistance may
increase with increase in temperature as stated earlier, but if the meta-
bolites happen to be more toxic than the original chemical the tempe-
rature effect can be reversed. Experimental evidence in favour of, or against
this possibility is' being sought.
Effect of Humidity 01\ Insect Resistance
Experiments were done on the effect of changes in relative humidity
on the toxicity of DDT and DNOC films to adults of Tribolium caslaneum
and larvae of Plutella maculipennls. With T. caslaneum adults the toxicity
of both DDT and DNOC films at higher relative humidities increased.
but with larvae of P. maculipennis the toxicity of DDT film decreased
and that of DNOC film increased. Neither these experiments nor the reports
in literature provide a basis for any general principle. However, it is
'pOSSIble to collect instances showing that, besides the insect and the in-
secticides, there may be various factors, such as range of humidity, strength
of the insecticide, stage of the insect, etc., which must be considered when
studying the effect of humidity.
Factors affecting Stomach-Poison Effect

Using DDT dust as insecticide and 3rd and 4th instar larvae of
Euproctis lunata as test material, the following fundamental aspects of
the bioassay of stomach poisons have been investigated.
Starvation and resistance. It has been indicated that, contrary to
previous notions, resistance of E. lunata larvae decreases with increase in
the period of starvation. The apparent increase in resistance during star-
vation is obviously due to the fact that prolonged periods of starvation
(24-48) make the larvae somewhat lethargic and eat less food. whether
poisoned or not.
Body weight and resistance. It has been shown that resistance to
insecticides increases with increase in body weight. An in«rease in the
quantity of poisoned sandwich eaten with increase in body weight was
observed, but the amount of poison ingested per- gram body weight of
insect remained within a comparatively narrow range and showed no
definite order of variation. It has been suggested that a more or less
, !
constant amQunt of DDT per gram body weight begins to have an inhibi-
tory effect on further feeding.
Development of Resistance an Index of the Intensiveness of

Control Operations
It is intended to invite attention to the idea that perhaps the deve-
lopment of' resistances can be taken as an index of the intensiveness of
the control operations carried out. In other words, instead of starting
from the initiation of control operations in an area and then trying' to
check where resistance has developed and where nbt, we can probably
proceed the other way also. Assuming biological variation to be universal,'
it may not be wrong to state that except in the case of total eradication,
which is rare, really effective and extensive control operations may be
taken to have been carried out only where resistance has developed.
Depending on the frequency of the resistance acquired by the insect
population, a certain minimum degree (which should be actually very
high) of insecticidal pressures should be necessary for the development
of resistance following control operations.
Suppose that, before any control operation has been carried out, the
population had 5% resistant individuals (R) and 95% susceptible ones
(S), i.e., S: R: : 95: 5, (5% R). If the first control operation kills say, 50%
of the population, then obviously those killed will be mostly the susceptible
individuals and the ratio. in the survivors will change to S: R: : 45 : 5
(10% R). After the 2nd cycle of operations the ratio will be S: R: : 20: 5
(20% R); after the 3rd cycle; S: R: : 7.5: 5 (40% R); and after the
4th cycle S: R: : 1.25 : 5 (80% R).
On the other hand, if the control operation is so effective that as a
result of every operation about 80% of the population is killed, then
after the 1st cycle of control operation the ratio will be S: R: : 15 : 5
(25% R) and after the 2nd cycle S: R: : 0: 5 (100% R).
If the control operation is still more effective and 95% of the popu-
lation is killed in every control operation, then after the very first cycle
of operation the position will be S: R: : 0: 5 i.e., a population with 100% R.
The proof for this broad generalization is indicated in similarly broad
observations, as follows:
1. Development of resistance has become more of a challenge in
public health control operation, and needs hardly any discussion that by
the very nature of circumstances the control operations in the field of pu blie
health have to be much more intensive and extensive than in the field
of plant health.
2. Even in the field of public health the resistance problems have
obviously cropped up more frequently either in economically advanced
PRINCIPLES, OF INSECT CQNT~OL 195
countries or in countries in which the advanced countries have had both

intensive and extensive control operations carried out.
3. Although agricultural pests had shown development of resistance
as early as 1914, this problem is still not very serious in agriculture beca-
use the magnitUde of the pest control problem precludes such high degree
of insecticidal pressure as would readily lead to development of resistance
among agricultural pests.
4. In agriculture the earliest resistance problem was with scale insect
infesting garden plants in conditions most suitable for the de~elopment
of resistance. In this insect the power of locomotion is very limited, and
intensive control :operation' is pOssible in the limited enviroilments of
garden farming, so to say, small· islands here and there.
The rational expectations of really effective control operations are:
(i) Development of resistance, which is in the very nature of the protoplasm.
(ii) Palliatiye effect, which is soon nullified by the building up of the
population by the progeny of the survivors; these, however, form a small
proportion of the total untreated population in which they get diluted.
(iii) Eradication, which is only an ideal to strive for especially in large
continental areas, and is to be tried by knock-out intensity of operation.
Of these 3 possibilities, (i) is happening in the field of public health,
(ii) is a common experience in agriculture, and (iii) is practicable only
under specially suitable conditions.
Thus, it may be pointed out that, if the foregoing line of thought is
correct, then resistance must develop sooner or later after really in-
tensive and extensive control operations.
Hazard to Human Health and upsetting the Balance in Nature

This topic has been dealt with at length elsewhere.
lusecticidal Formulations and Appliances
It is better to leave this topic to be dealt with by chemists, physicists
and engineers. From the entomologists' angle. the requirements are (i) low
cost. (ii) minimum risk of overdosing to avoid phytotoxicity, (iii) maxi-
mum uniformity of distribution, and (iv) optimum efficiency.
Sterile male technique. This method has made a lot of impact on
the morale of economic entomologists. The phenomenal success in era-
dication of the screwworm pest of sheep and cattle in the USA by the
release of males sterilized by irradiation has already become an entomo-
logical classic. This novel and economical technique has had a tremendous
impact on the whole. approach to entomological problems all over the
world. In fact, the effect which this particular use of radioactivity has
produc~d is somewhat analogous to its effect in producing mutations.
and the success of the sterile male technique has, so to say, led to much
mutation in entomological thinking and to a vast variety of new research
ideas.
A number of pests other than screwworm have been tried out, and
only partial success has been achieved. This underlines the neceSSIty of
a very careful selection of the pest species for which this method should
be trie<;l. Several scie:qtists including Knipling. the origmator of this
principle, have listed the essential requirements for success in eradicating
a pest by this technique. The following 3 conditions have to be ensured
even for a preliminary selection of a pest species for such a study.
(i) The population of the pest should be either inherently very low or"
should come down to a very low level as a result of seasonal changes or
by conventional methods of pest control. The population of the screw-
\Vorm with which this technique succeeded so well was in the region
of a few hundred per square mile, whereas agricultural pests of any
significance are generally ,in thousands per acre. This is the most essential
requirement because the number of sterile males released per unit area
has to be several times the population of the normal males present in
that area. Hence, the higher the population level of the pest, the larger
the number of sterile males to be released and also the higher the cost
of the operation. Thus, beyond a particular level of the pest population
this technique ceases to be feasible.
(ii) The area from which the pest is to be eradicated should be so
situated that reinfestation can be excluded or kept to a negligible level.
In the case of the screwworm the strictest possible quarantine measures
are enforced.
(iii) There should be a reasonable chance of economical mass-rearing
of adults in such large numbers as would be needed for release.
Thus, the most promising field for success of the sterile male technique
is likely to be the control of storage pests in the microcosm of st9rage
structure or godown. If it is not suitable for this microcosm it will be
much less so for the agricultural fields. This theoretical decision was
arrived at in spite of the rather disappointing conclusion ·with the flour-
mill moth in England. One can envisage using the sterile male technique
neither in a normal godown nor in a general way, but in the following
manner in which both the storage structure and the general 'storage
practices are modified to suit the requirements of the technique.
(i) Earmark certain structures for long storage.
(ii) Make them practically insect-proof by suitable barriers.
(iii) Disinfest them by spraying or fumigation.
(iv) Store fresh grain or fumigated grain which is practically free from
insect infestation.
(v) Before closing the storage structure, introduce adequate numbers

of sterile males of the locally important pest species.
The quantity of foodstuff which the sterile. non-multiplying adults of
the beetle pests are likely to eat is negligible; several pests. particularly
moths. do not eat in the adult stage.
Chemosterilants. In the original technique involving sterile males the
agent for sterilization is the gamma-ray. Certain chemicals have also
been found to bring about sterility. The likely advantage of chemosteri-
lants over gamma-ray is that they can be applied directly to' the field
population. particularly in combination with some attractant. and this
will eliminate the ,need of ryaring the males in very large numbers. How-
ever. before this method becomes practical the problem of special -hazards
involved in the use of ~hemosterilants will have to be satisfactorily solved.
It should prove better than ordinary insecticidal treatment because the
insects treated with chemosterilants will also control other insects not so
treated. The use of sex-attractants is quite promising. and their specificity
is the strongest point in their favour where this is particularly needed.
Juvenile hormones. These hormones are also attracting a good deal
of attention. particularly because of their novelty. Otherwise these chemi-
cals will only add to the already huge list of insecticides along with all
their limitations.,
, Introduction of lethal genes. So far very little work has been done
in this promising field.
REFERENCES
Bhatia, S. K. and Pradhan. S. 1968. Indian J. Ent. 30 (1): 13-32.
Campbell, F. L. 1926. J. gen Physiol. 9 : 433-443.
Hoskin. W. M. and Gordon, N. T. 1956. Ann. Rev. Enf. 1 : 8-122.
Lindgreen, D. and Shepard, H. H. 1932, J. econ. En!. 25 : 248-253.
Pradhan, S. and Bhatia, S. C. 1951. Bull. ent. Res. 42 (2): 399·419.
Pradhan, S. and Bindra. O. S. 1956. Indian J. Ent. 18 (2): 93-111.
CHAPTER 10
PESTICIDE HAZARDS
THIS is a subjec't on which many people have strong views. Some ate
against the ~se of pesticides; others favour it. Both types of opinions are
doing more' harm than good. A balanced view is an urgent need of the
time. There are various aspects of pesticide hazards, but it is proposed
to deal here with only one aspect, viz, the pesticide hazards to man under
present-day conditions in India.
Basic Points to be kept in View

The following basic points have to be kept in view.
(a) Public concern about hazards of large-scale use of pesticides is
universal. Advanced countries like USA and UK have taken cognizance
of these hazards from the very beginning. In UK the Minister of Agri-
culture and Fisheries appointed a working party under the chairmanship
of Professor Zuckerman in 1950, and since then several committees have
gone into the matter. The final stage is represented by Cook's reports
(1964) in which specific recommendations have been made for continuing
or discontinuing the use of particular insecticides for particular purposes.
In USA the actions taken have been all the more legalistic, and a number
of Acts and Amendments have been passed from time to time since 1938.
The latest position is represented by the report of the President's Science
Advisory Committee (1963). In 1963, WHO and FAO held a joint meeting
on the evaluation of the toxicity of pesticide residues. Thus, it is impe-
rative for India to take serious note of the problems of pesticides and to
create effective organizations for keeping the hazards under check. In
fact, this action is overdue.
(b) At the same time, because of the immense amount of good done
by pesticides, particularly in increasing the efficiency of production and in
preservation of food, the hazards involved must be properly weighed
against this benefit before accepting or rejecting the use of pesticides.
It is difficult to visualize a total stoppage of the use of pesticides in
modem agriculture. Before the discovery of DDT, agricultural scientists
used to dream of a, persistent contact poison because all the inorganic
insecticides then used acted only as stomach poisons and hence were of
limited utility. The present wide-spread concern about the adverse effects
of modem pesticides is a testimony to their popularity.
(c) All modem advances involve a certain amount of risk, and all
human activities are subject to accident~. H~pce, accidents associated with
PESTICIDE HAZARDS 199
the use of pesticides, such as those which occurred in Kerala some

years ago and those subsequently reported from the Punjab, should lead
only to greater awareness and alertness of the public and the govern-
ments; they should not be allowed to create undue scare in the public
mind. Even in advanced countries like Japan there have been seyeral
accidental deaths every year from Parathion, the insecticide involved in
the Kerala accident.
(d) Developing pesticides is a time-taking and costly proc~ss. Each
marketable chemical meant an investment of I to 3 million dollars before
it was made available. For investment of such magnitude the industry
certainly requires proper incentives, which should not be marred by
taking an unball;lnced view' of the situation.
(e) When knowledge is scanty, precautions are all the more necessary.
It is often argued that' restriction on the use of pesticides in the absence of
definite information of the harm done by themJ is not quite scientific. It
may be pointed out that collectiqn of necessary information is bound to
take considerable time, and it is not unlikely that if a harmful chemical
is already in indiscriminate 'use it might well cause enough damage before
sufficienf scientific data regarding its hazards are collected.
Types of Pesticides Hazards

There are various types of hazards of the large-scale use of pesticides
in the fields of agriculture and public health. The following classifications
from different angles give an idea of their nature and magnitude, which
have to be kept in view in recommending precautions against them.
(a) Hazards at different stages. (i) Accidental and intentional poison-
ing. Because most pesticides are poisonous to warm-blooded animals in-
cluding man, chances of accidental poisoning are always there. These
poisons can also be used for suicide or homicide. These hazards, how-
ever, are not peculiar to pesticides; they are common to all kinds of
poison, including sleeping pills.
(ii) Operational hazards during manufacture and distribution of
pesticides.
(iii) Operational hazards during application of pesticides.
(iv) Post-application hazards due to pesticide residues.
(b) Risks to different categories. Effort has to be made to safeguard
the following 5 categories from pesticide hazards.
(i) Pesticide users
(ii) Factory workers
(1ii) Consumers of treated food and fodder
(iv) Third parties, such as children, animals. passers-by, and so on.
ic) Qrqded types of pesticide toxicity. (i) Acute toxicity. Two types
of acute toxicity have to be reckoned with, i.e. oral toxicity from acci-
dental or intentional ingestion of the pesticide or heavily contaminated
food, and dermal toxicity from spilling of the pesticide concentrate over
the body or garments of workers. Although oral toxicity is more acute
and dangerous than dermal toxicity, it is the latter which is more difficult
to avoid.
(ii) Subacute toxicity. This occurs when exposure continues for several
days so successively and excessively that the enzymatic function does not
fuIIy recover and the poisons are not completely eliminated during the
night's rest. Such a situation may arise in the case of farmers who try to
complete their spraying schedule within a short period. ,
(iii) Subchronic toxicity. Workers in pesticide factory, personnel'
engaged in pest control and vector control, and aircraft pilots may become
repeatedly exposed to small doses for a full season of pest control.
(iv) Chronic toxicity. This is largely from pesticide residues in food
and environment. Every nation and its laws are generally prepared ade-
quately to face the problems of acute toxicity, but those of subacute and
chronic toxicity are more difficult to deal with. The public, in general,
has become most concerned about hazards of chronic toxicity due to
increasing quantities of pesticides being used in modem agriculture.
(d) Inherent types of operational hazards. These are mainly due to:
(i) spilling over of pesticide formulations in careless handling.
(ii) inhaling of the pesticide mist or dust, and
(iii) coming in contact with treated crops or surfaces, or handling of
used equipment.
Generally, protective clothing and gasmasks are recommended for
avoiding operationa'l hazards, but these recommendations are generally
impracticable in the excessive heat and humidity of tropical and sub-
tropical countries.
(e) Indirect hazards through food chain. Public safety has to be
ensured not only against direct poisoning by pesticides but also against
indirect poisoning through the food chain. Contamination of fodder or
dairy premises may lead to excretion of the pesticide in milk. Grazing by
cows and goats on treated pastures may lead to contaminated milk and
meat. At times this food chain leads to a progressive concentration of the
pesticide.
Review of Recommendations of Various Committees jn UK and USA

and of FAO and WHO in the Li!!bt of Conditions in India
It was easy for the President's Science Advisory Committee {USA)
and the Cook's Committee (UK) to examine each of the uses of particular
insecticides because in either country there exists adequate official machi-
PESTICiDE HAZARDS 201
nery for formulating and implementing what may be called official

recommendations for pest control. In the USA there is a Federal
Pest Control Review Board and an Interdepartmental Committee on Pest
Control. In the UK this work is carried out by (i) the Agricultural Che-
mical Approval Organization and (ii) the Advisory Committee on Poison-
ous Substances used in Agriculture; Food and Storage and its Sub-
committee; and Veterinary Products Safety Precautions Scheme. In India
there is no organizations like these. Although there are a number' of book-
lets containing recommendations for pest control. yet all these recommen-
dations are from individuals and on individual's personal responsibility.
There is no organization or body to scrutinize all these varied 'recommen-
dations and to /formulate' such agreed recommendations as may guide,
and be obligatory for, those engaged in control operations. Thus,
recommendations for the control of different pests vary wide~y as regards
the active chemical 'to be used, its formulations. and the quantitv to be
applied per unit area. For checking hazards it is necessary that these
recommendations are first reduced to a small manageable' number and
then scrutinized from the poiilt of view of residue and health hazards.
We should also constitute what may be called the Insect Pest and Disease
Control Advisory Committee (IPDCAC). which should make official
recommendations for the control of different pests and diseases and also
'for integrated control schedules for different crops. Its functions
should be somewhat on the lines of the Variety Release Committee. which
considers the varieties recommended bv plant breeders and officiallv
releases some of them for adoption. In the same manner. the IPDCAC
should examine all methods of pest control and officiallv anDrove them
before the extension workers carry them to the cultivator. These
recommendations should be reviewed at periodical intervals in the light
of further knowledge regarding their efficacy and the hazards involved
in their implementation.
In both UK and USA there is arrangement for registration of pesticides.
No such arrangement exists in India. In the USA and UK the manufac-
turers have to supply detailed toxicological data before they are aIlowed
to market the pesticides. These data, with necessary modifications. can
be adopted for Indian conditions; but pesticide formulators and distri-
butors should be required also to suoplv accumulated data on pesticide
residues on various crops for which their use is reocmmended.
Public concern in UK and USA regarding pesticide hazards has been
realistic. based on the results of monitoring of pesticide residues. In India,
in the absence of such information. it is not possible to make specific
recommendations. It is, therefore. suggested that provision should be
made for monitoring pesticide residues mainly to assess the extent of
202 AGRICULTURAL El'lIOMOLOGY AND PEST CONTROL
hazards invo)ved in the use of pesticides in agriculture, forestry, fisheries,

and veterinary and public health including hazards resulting from moth-
proofing of human clothing. Work done at the IARI has shown that this
monitoring work will require careful standardization of each technique
to be used in the chemical analysis of a particular pesticide used
on a particular commodity. Investigations on r~sidues of Pesticides
have been. going on for some years at the JAR!. Studies bn the
persistence of some commonly employed pesticides, such as DDT.
BRC, Lindane, Aldrin, Endrin and Malathion, with respect to different
crops and crop materials have provided basic information for
toxicity studies related to man and animals, modification of application \
schedules, -safe interval between application and harvest, and so on. This
is, however, only a small beginning, and much mor~ remains to be done
with respect to many more chemicals, crops, soils, alld climatic conditions
in various regions of the country.
That the recommendations made in other countries have to be care-
f\1\\)l 1>cru.\ini'Leu in \be \igDt t>l ct>noitit>m> in Inoia Y1, b\:lITte 'i.'lu', uJ tbe
fact that the main risk from pesticide residues, oll the basis of which
Cook's recommendations are made, appears to be for predatory birds.
Indian agriculture is becoming more and more concer1led about the number
of species of birds that have become serious pests of both agricultural
and horticultural crops. The balance sheet of the utility and harmfulness
of different species of birds taking mixed vegetable and animal diet is
yet to be worked out. It has also to be decided hoW' far predatory birds
are useful in controlling the pests against which the pesticides are used.
The recommendations of the American committee are (i) to assess the
level of pesticides in man and his environment, (ii) to formulate meaSUres
which will augment the safety of the current practices, (iii) to undertake
needed research for development of safer and more specific methods of
pest control, (iv) to suggest amendments of public laws governing the
use of pesticides, and (v) to educate the public. for the first purpose
the committee recommended the development of a programme for collect-
ing comprehensive data so that levels of pesticidetl can be determined
in occupational workers and others, and a continuing network to monitor
residue levels in air, water, soil, man, wildlife and fish: For the second
purpose its recommendations are to improve and augment the organi-
zation for reviewing (a) residue tolerances and the experimental studies
on which they are based, and (b) current and proposed control and eradi-
cation programmes to reduce their hazards. For the third purpose it
recommended that emphasis on research should be shifted from broad-
spectrum chemicals to such items as could lead to selectively toxic
chemicals. non-persistent chemicals, selective met}lods of application,
and non-chemical methods as well as more research on toxicity studies

related to man. For the fourth purpose it recommended that public laws
should be reviewed to make them more effective; for example, it should
be required that every pesticide formulation carries its official registration
number on the label. For the fifth purpose it recommended that pro-
grammes of public education on the toxic nature of pesticides and thefr
proper use should be augmented.
In the joint meeting of FAO and WHO (1963) on the evaluation of
toxicity of pesticide residues in food a very useful concept of 'acceptable
daily intake' of different pesticides, was developed. On the basis of all
toxicological data available'; an effort was made to determine the quantity
of each insectidide in terms of mg/kg body weight which man could
daily consume without appreciable risk for his full life time. It was
suggested that this information could serve as a basis on wh'ich different
nations mjght form their own recommendations on the permissible limits
of contamination by each pesticide on each commodity based on average
consumption of each commodity and average weight of individuals.
Nature and Extent of Hazards in Pest Control Operations in India

Vegetables. Pests of vegetable crops are active at all stages of crop
,growth, and application of pesticides and harvesting follow each other.
In general part of vegetable plants goes waste; some parts are consumed
by man, the rest by animals. Thus. hazards of toxic residues of pesticides
are for both man and animals. Therefore, the greatest caution is necessary
in applying toxicants to vegetable crops. At present DDT. BHC, Malathion.
Parathion, Endrin, copper fungicides, and mercurials are recommended
for pests and diseases of vegetable crops. The hazards of these on vegetables
like bhindi, cauliflower, and cabbage are obvious. They also fall on the
ground and persist there for some time; root vegetables grown there may
become contaminated and hazardous to the consumer. Even if less per-
sistent pesticides are used, the hazards will be there because of the
continuous process of harvesting day after day. Thus, in the case of
vegetables, pesticide hazards are m~st serious, and strict vigilance is
absolutely necessary to ensure that vegetables with excessive insecticide
contamination are not marketed.
Paddy. On this crop there are more than 2 dozen pests, of which
about a dozen are considered major pests. Most major pests cause
damage in the nursery or earlier stage. when treatment is most needed.
One important pest, the rice gundhy bug. does maximum damage when
the grains are in milky stage; treatment against it is applied in the ad-
vanced stage of the crop. The commonly recommended pesticides against
paddy pests are DDT, BHC, Endrin and Parathion; for certain pests
Aldrin, Diaz!non and Nicotine sulphate have also been recommended.

When the crop is treated in the young stage the danger of toxic residues
on harvested produce is likely to be negligible, but when the crop receives
the treatment in an advanced stage, as for paddy bug, there can be some
hazard to consumers and particularly to animals fed on the treated straw,.
There is also $ome risk of skin contamination at the time of transplanting
paddy if the nursery has been treated recently and the farmer ,has to
plant the treated seedlings for hours together day after day. Hence" it is
necessary that first all the recommendations about the control of paddy
pests are scrutinized by the proposed IPDCAC, and the few treatments
recommended by it are carefully examined by the Pesticide Hazards \
Committee-(pHC) and tolerance limits fixed for both grains and straw.
Sugarcane. There are many sugarcane pests. of which the several
species of borers, pyrilla. and whitefly are most serious. Termites and
root grubs are also a menace in certain areas. The commonly recommended
insecticides against sugarcane pests are BHC, DDT. Endrin and Parathion
fOT anplication to tbe aerial portion of the crop. and Aldrln, Hep\achlor
and Chlordane for soil treatment against termites. Most of these insecti-
cides are persistent; hence. there can be residue toxicity, firstly in cane
juice, which is often consumed as such, secondly in tops which are fed
to cattle, and thirdly in gur, which is not subjected to severe processing
(sugar undergoes too drastic a processing to retain any residue). There
is also some hazard of skin contamination in workers who carry out
various agricultural operations in the treated field. particularly at harvest
time. It is advisable to carry out monitoring work to estimate the extent
of contamination at all these stages.
Cotton. The commonly recommended insecticides against cotton
pests are Endrin, DDT, BHC, Malathion, Parathion and Diazinon. The
main residue hazards from this crop can be (i) through skin contamination
of labour working in treated fields, particularly at the time of picking,
(ii) feeding of cottonseed and seed cake to cattle, and (iii) in the cotton-
seed oil used for hydrogenation. The hazards throu!!h the last 2 media
ordiinarilV should be quite small because the seed remains protected
inside the boll and by lint after the boll ooens. All the same, because
insecticides are generally Iipophilous and cottonseed contains a good
amount of oil, the final decision can be based only on the results of
exoerimentaI assessment.
Groundnut. The groundnut crop is subiect to infestation with a'
number of pests, and the pesticides commonly recommended are BHC.
DDT, Parathion, lIETP and Endrin. These oesticides are applied at
various stages of the crop. and a good proportion falls on the ground.
The main hazard can be to those who eat groundnuts without roasting.
PESTICWE HAZARDS 205
It is, however, desirable to undertake monitoring work in the case of

lrmts, vegetable fat, and the cake.
. Jute. BHC, DDT, Malathion, Endrin and Dieldrin are the pesti-
cldes used (i) the main hazards are occasional use of tender leaves for
culinary purpose, and (ii) skin contamination of workers who. take out
the jute fibre.
Tea. and coffee. These are treated mainly with copper fungicides.
When pesticides have to be used on tea" the residues must remain within
the limIts laid down by importing authorities. I
Coconut and arecanut., Since the produce used for consumption' is

heavIly covered :with husk, the risk involved is negligible; but there can
be skin contamination when the labour comes in contact with treated
truits and plants while picking and extracting the juice.
Chillies. The crop at various stages is treated against the leaf curl
diseases with toxic and persistent pesticides; hence, toxic residues in the
fruits are expected. The actual extent of hazard will correspond with the
proportion of chiilie\) in the diet. There is also risk from contact with
treated plants at the time of picking.
Fruit trees. Among the temperate fruit trees apple and peach receive
most attention; ,insecticides are, ~pplied when the trees are in the dormant
. condition or before fruit formation; so, normally residue hazards are not
likely to exist on harvested produce. Among the tropical fruit trees
mango and citrus are treated with pesticides like DDT and Endrin, mostly
before or at the flowering stage.
Maize. Quite a number of pests occur on this crop. The commonly
recommended insecticides are ,DDT, BHC, Parathion and Endrin. Maize
is commonly used as fodder; hence studies on the assessment of insecti-
cidal residue found on crop treated in different ways with different pesti-
cides are necessary. If the crop is treated when cobs have formed, con-
tamination of grains is less because they are covered with husk; but
assessment of the extent of contamination of mai:ze plants even at this
stage is called for because they are fed to cattle.
Harvested produce. For protection of the produce from storage pests
are recommended (i) fumigants, such as EDCT, methyl bromide, and
Phostoxin, and (ii) BHC for dusting outside the bags. It has been reported
that cultivators and traders resort also to direct mixing of DDT, BHC,
etc. This is very hazardous and should be· banned. Monitoring work is
needed to evaluate the extent of contamination from dusting of bagged
grains and also the persistence of fumigants in the commodity.
Forestry. The main hazard to wildlife is from pest control operations
carried out by the forest departments; but this does not appear to be on
such a scale as to cause any real concern at this stage.
Veterinary. Pesticidal treatment of milch and meat animals obviously

involves greater hazards than treatment of most crops because animal
fat is likely to absorb and store the pesticide, the concentration of which
can become augmented and render meat and milk more harmful than
through skin contamination with the original dose level.
Review of P~ecautions Generally Recommended,
and Additional Suggestions
(1) Measures to prevent hazards at the stage of manufacture and
formulation. (a) MEASURES GENERALLY RECOMMENDED. There is an In~
dustrial Hygiene Division of the US Public Health Sciences which issues
instructions' to the industry to adopt uniform and co-operative steps to
prevent hazards. The instructions include provision of (i) proper exhaust
ventilation, (ii) suitable working clothes, and (iii) adequate washing facilities.
The primary producers of the chemicals have also extended their
responsibilities by advising and controlling the activities of the firms that
formulate their products.
(b) ADDITIONAL MEASURES SUGGESTED. The following further meas-
ures are essential:
(i) Authority with the government to inspect pesticide manufacturing
plants.
(ii) Prevention of faulty disposal of factory affluents and wastes; pesticide
firms. manufacturers and formulators to obtain a waste disposal permit.
(iii) Prevention of dangerously adulterated and mislabelled pesticides from
reaching the market, and provisions for their seizures.
(iv) Provisions to see that the manufacturing enterprise are not located
near populated areas.
(v) Regulations for disposal of empty pesticide containers.
(vi) Provision of warning facilities in cases of excessive contamination
during manufacture and formulation of toxic materials.
(2) Measures to prevent hazards at the stage of marketing. In the
USA the Insecticides, Fungicides and Rodenticides Act prov_ides that every
product must have attached to it a label showing (i) name and address
of the manufacturer, (ii) name, brand or trade-mark under which the
chemical is sold. (iii) the net content, (iv) ingredient declaration, (v) appro-
priate warning or caution statement when necessary to prevent injury to
man, IJvestock vegetation and useful invertebrate animals, (vi) labels of
the world 'Poison' in red skull and cross bones, and a statement of
antidotes. Provision is made also for testing products as they are encoun-
tered in the regular channels of trade to determine whether they are in
compliance with the law. Those that are found to violate the law may be
siezed by the government and removed from the channel of trade.
ADDITIONAL SUGGESTIONS
(i) The internationally accepted common name must be indicated con-
spicuously on the labels.
(ii). The sale of pesticides should be restricted to farmers who are edu-
cated and trained for the purpose and who use them under the
supervision of plant protection, block development, or other officials
of the state department of agriculture.
(iii) Pesticidal formulations, particularly concentrates, should be marketed
in suitable containers of handy size so that there is least chance of
unconsumed material being left.
(3) Measure,s to ,check hazards to workers' engaged in pesticide use
programme. In the UK the Agriculture (Poisonous Substance) Act is
I
designed to protect agricultural workers by ensuring that they are supplied

with protective clothing when working with the more toxiy pesticides
included in regulations made under the Act. In India t4e following
recommendations are generally made: (i) use of protecqve clothing,
(ii) operators not to work for more than 8 hours a day, (iii) operators to
be checked by a physician periodically.
FURTHER SUGGESTIONS
(i) Protective Clothing should be specified, keeping in view the weather
conditions under which the operators have to work. Operational studies
on this aspect need to be carried out; for example, 'it wiall be necessary
to give periodical recess for pesticide operators, and to allow each operator
to stop work; for a suitable period and take rest in fresh air before resum-
ing it.
(ii) The operators should be trained in safe handling of insecticides.
(iii) Popular publications in local languages on correct methods of
application, personal- hygiene and protection as well as storage and dis-
posal of containers would be useful.
(4) Measures to check post-application hazards. (a) MEASURES
ADOPTED IN OTHER COUNTRIES .. (i) Fixation of tolerance limits for each
pesticide on each commodity. The Pesticide Chemicals Amendment to
the Federal Food, Drug and Cosmetic Act (Public Law 518) (The Miller's
Bill) in the USA provides practical methods of establishing safe tolerance
for residues of pesticide chemicals in food. Under this law the petitioner
who requests establishment of a safe tolerance for insecticides submits
data to the government to support the application. Information is required
on the following:
(a) Name, chemical identity and composition of the pesticide chemical
(b) Amount, frequency, and time of application of the pesticide
chemical.
(c) Full reports of investigations made on the chemical (long-term

feeding tests required).
(d) Resul.ts .of tests on the amount of residues remaining, including
descnptlOn of the analytical method used.
(e) Practical method for removing residues which exceed any proposed
tolerance.
(ii) Fixation of acceptable daily intake of each pesticide. A joint
meeting of the WHO and FAO in 1963 prescribed the values of acceptable
dally intake of a number of pesticides on the basis of all available data.
(iii) Fixation of minimum time lag between application of pesticide
and harvest of the crop. In the UK a working party suggested restrictive'
conditions- of pesticide use, viz, use on specified crops only, maximum
rates of application, and minimum intervals between the last application
and harvest.
(iv) Banning or discouraging the use of certain pesticides. This has
been suggested by Cook's Committee (UK) for certain insecticides.
'(b) SUGGESTIONS FOIt INDIA. (i) India should adopt No. (i), enforced
by law in USA. and fix tolerance limits for each pesticide on each
commodity.
(ii) It should fix the minimum time lag between application of pes-
ticide and harvest. and this should be advisory so that cultivators may
be reasonably sure that their produce will comply with legal regulations
for tolerance limits.
(iii) For the acceptable daily intake no specific action is needed sepa-
rately because these values are considered to be applicable to the human
species in genrea1.
(iv) Baruring the use of an. insecticide may be don.e ocly when. theIe
are enough scientific data for recommending a firm decision and there is
a prima facie for it. For example, there can be hardly any difference of
opinion on the necessity of extreme precautions against direct contamina- .
tion of foodgrains and fresh vegetables and indirect contamination of milk.
(v) It should also give such advice as may be necessary for reducing
hazards of skin contamination of cultivators during weeding, picking,
harvesting and handling treated harvest.
It should be easy to recommend, on the basis of existing knowledge
and com~on sense, precautions against operational hazards from pesticides
right .up to the application of pesticides. It is, however, the hazards from
pesticide residues which have caused the greatest concern all over the
world. For recommending precautions against residue hazards many
carefully collected data are needed, but these are not available for Indian
conditions. Hence, only the most obvious precautions can be recommended
at present. It is necessary to constitute certain standing bodies and to create
certain organizational facilities for keeping constant vigil over pesticide

hazards and for collecting necessary data for formulating rational precau-
tions from time to time. .
1. For chemical treatment of vegetables, harvestjng of which continues
for a good part of the season when pesticidal operations have to be carried
out, only safer insecticides of plant origin should be used. Even with
readily breaking down organo-phosphorus insecticides fatal accidents may
occur sometimes because of harvesting and consumption soon after spray-
ing, and it may be difficult to ascribe these to pesticide poisoning;because
by the time the legality of getting the residues is complied with
the insecticides may break down into non-toxic compounds. It is necessary
to make legal prqvisions to' collect samples from vegetable markets to
have them quickly analysed, and to take necessary action against those
found guilty.
2. Special precautions are necessary to check insecticide contamina-
tion of cere~lls, pulses and tubers (potato, sweet-potato), which are consum-
ed in large quantities and to preserve which it is tempting to use insecti-
cides. Hence, it is necessary to discourage this practice as much as possible
and to make legal provisions to collect samples from the market for quick
analysis and prompt legal action. Thjs is particularly necessary because the
alternative fumigation methods are there to check infestation of stored
products.
3. Efforts should be intensified to increase the production of pyre-
thrum, for it provides the safest insecticide so far known. An argument
against pyrethrum is its high cost and non-avaifability. On the other hand,
producers of pyrethrum complain of insufficient interest in its use. A
conference held in Jammu in 1961 passed the following resolutions.
"It was unanimously felt while the users of pyrethrum are not sure
of the supply of pyrethrum in sufficient quantity and at competitive rates,
the producers of pyrethrum are apprehensive that they may not find proper
demand for their produce. When the question of enhancing the pyrethrum
production is discussed, doubts are often expressed whether pyrethrum
will be able to compete with synthetic products. On the other hand, when
the question of using pyrethrum on largy scale is discussed. the argument
is put forth that pyrethrum is not available and is very costly. It is, there-
fore, fully established that on its intrinsic merits, pyrethrum is preferable'
to most of the synthetics especially in the field of storage of foodgrains.
Both pyrethrum industry and food grain storage are suffering mainly be-
cause of a lack of proper co-ordination between the producers and users
of pyrethrum".
"It was recalled that due to the fact that the producers did not find
proper market, the cultivation of pyrethrum had to be given up in Assam
210 AGRICULTURAL ENTOMOLOGY AND ~ST CONTROL
and in the Nilgiris and even in Kashmir area under pyrethrum cultivation
haQ to undergo considerable reduction. On the other hand, it is mainly
because of the non-availability of pyrethrum in sufficient quantity and
at remunerative price that toxic chemicals like Lindane, Malathion, etc.
are being recommended for use in the storage of foodgrains."
"Discussion on various other plant products showed that what has been
stated above about pyrethrum is also applicable in various degrees to
several drugs of plant origin. It was, therefore, unanimously resolved to
recommend the formation of an AIl-India Co-ordination Council for, the
Production and Utilisation of Pyrethrum and other such plant products.
This Council should be entrusted with the task of maintaining a close
liaison between the producers and users of pyrethrum and other such
products to ensure a proper supply of pyrethrum to the users and proper
market to the producers."
4. Specific legal provisions should be made for setting up tolerance
limits of pesticides, as has been in the USA. The Central Committee for
food standards which, it is understood, has made some attempt for
recommending tolerance limits for certain chemicals is not the appropriate
body because tolerance limits are to be fixed not only for food grains but
also for vegetables, fruits and even fodder. The standing committees and the
organizational facilities needed for this and related purposes are indicated
below.
5. A standing Insect Pest and Disease Control Advisory Committee
(IPDCAC) should be set up somewhat on the lines of the Federal Pest
Control Review Board of the USA. This committee should act on the
lines of the American Board and on the Agricultural Pesticides Approval
Scheme of the UK. It should periodically examine all methods of pest
control recommended by various workers in the country and should offi-
cially approve those which show optimum combination of efficiency and
safety. The committee should consist of research and extension workers
in almost equal numbers. It should periodically invite recommendations
for pest control from various workers in the country, consider them, and
issue a list of approved methods of pest control from time to time. This
should be mainly on the basis of efficiency of the pest control methods
recommended, but the committee should take into consideration the haza-
rds also. It should fix and periodically review the minimum time that
must elapse between the last application of insecticide and the harvest of
diffetent crops to ensure with reasonable certainty that the residues will
remain within tolerance limits.
6. A standing interdepartmental Pesticide Hazards Committee (PHC)
should be constituted on the lines of the British Advisory Committee on
Poisonous Substances used in Agriculture ,and Food Storage. This CQ-
mmittee should carry out mainly the following functions:

(a) To fix and periodically review tolerance limits of various insec-
ticides in various commodities of food, fodder, etc.
(b) To determine and periodically review the hazards from grazing
by animals and skin .contamination in men and animals during agricultural
operations in treated fields.
(c) To determine and periodically review all operational hazards in
the manufacture, distribution, dilution, handling. etc. of insecticides.
(d) To determine and periodically review the contamination of the
environment by insecticidal fumes from factories and refuse let 'out into
rivers, etc. '
(e) To keep a vigil oq the new types of hazards arising fro~ the
introduction of new chemicais or new forms of application.
7. The functions of and the interrelations between. the IPDCAC and
the PHC should be clearly spelt out .The interrelations should be as follows.
The IPDCAC should first finalize its selection of control measures
purely on ,the basis of their efficiency. and thereafter the PHC should
scrutinize these selected methods iIi respect of hazards. Finally. the IPDCAC
should consider the recommendations of the PHC and finalize its own
recommendations for extension workers and pest control personnel on the
basis of optimum combination of efficiency and safety. .
8. There should be a legal provision that only those pest control
measures shall be carried out which have been approved by the IPDCAC.
9. An effective monitoring organization for proper assessment of
pesticide residues should be established.
10. Agricultural operations are generally intermingled. For example.
chemical spraying against pests, diseases and weeds may get intermingled
with operations like weeding and repeated harvesting of crops like cotton.
It is necessary to save the workers from contamination with pesticide (e~
sidues on crops. The extent of such contamination should be studied. and
necessary precautions taken against such hazards.
11. The nature of protective clothing should be specified. and there
should be legal provision to have the items periodically checked. The PRC
sh<;mld examine the various types of protective clothing available in the
country from time to time. For exampie. a medical engineering firm put
forth gasmasks and certain other equipment for the use of the workers in
the pesticide industry and research institutions. These devices should be
allowed to be marketed only after they have been approved by the PHC.
12. As regards other operational hazards. those discussed in previous
section should be adopted.
CHAPTER 11
REVOLUTION IN PEST CONTROL
So far, the n~tional efforts have been concentrated mainly on production-

oriented research, and protection-oriented research has been neglected. The
adverse effects of this lopsided thinking, execution and development ,have
begun to pose serious challenges to the very production efforts on which
the nation has been more or less exclusively concentrating so far.
Negative correlation between Success in Production Efforts and

Intensity of Pest Problems
Somebody may argue that production efforts include protection efforts
as well, but in practice it has not been so. The result is that a negative
~orrelation has developed between success in production efforts and inten-
sity of pest problems in major crops such as wheat, cotton, and rice (Fig.
11.1). The production has shown phenomenal increase in the relatively
pest free sop wheat, but such increases are not observed in the other sop
mentioned and which are prone to be damaged by a number of serious
pests. Pest proble!lls constitute the real bottleneck in the case of rice,_and
non-entQmological explanations, for the slow progress in. increasing rice
production do not stand scien~ific scrutiny. Cotton comes between wheat
and rice as regards the intensity of, pe,st problems as w~II as the success of
our e.fforts to increase its production. A similar picture emerges from a
~omp,arison of yield increases in sorghum and bajra. Although, in the
case of s~)fghum much more research effort has been put in, the increase
in yi,eld, has Qeen much faster in bajra, in, which pest problems are much
,milder than in sorghum. Thus, India~s agriculture seems to have been
brought to a stage where insect pests have begun, to determine the amount
of suc,cess the country is likely to achieve in its efforts to increase p~oduc
tion in any particular, crop. This is the statistical revelation of the country-
wide experiment during the, sixties, without anybody being seriously con-
sc~ous of it. This indicates the extent of damage done by neglect of pest
control research.
Extraordinarily higb Losses due to Insect Pests

Results of some recent precise experiments specially designed and
carried out to assess the losses inflicted by insect pests alone on various
high-yielding varieties grown under high-yielding crop husbandry practices
have shown extraordinarily high percentage losses. Table 11-1 shows the
results of trials at 14 different places under the All-II).dia <;o-ordinated
REVOLUTION IN PEST CONTROL 213
70
60
w 50
en
<{
w
a::
u
z
40
LU
<..?
~ PRODUCTJON
Z
w 30
u
a::
w
a.. YIELD '
20 UI1IJ]
10
o
WHEAT COTTON RICE
CROP LEAST CROPS VULNERABLE
VULNERABLE TO PEST
TO FIELO PESTS INFESTATION
Fig. 11.1. Percentage increase in yield and production of crops with different
degrees of vulnerability to pest infestation.
Rice Improvement Project during 1969. The all-India average increases

in yield_due to pest control alone were 194% in 'TNI' and 108% in 'IR-8'
the maximum reaching 729% in 'IR-8' at Warangal and 1005% in 'TNI'
at Maruteru. These losses immediately explain why no rice revolution has

been possible so far, as has been achieved in the case of wheat. A similar
situation is depicted in Table 11.2 for cotton.
-
TABLE 11.1. PERCENTAGE INCREASE IN RICE YIELD DUE TO PEST CONTROL
Location 'IR-8' 'T (N) l' 'W 1963' 'CR 44-93' Local
Warangal 729 615 82 147 375

(4188) (4690) (4084~ (2727) (3~20)
Tenali 253 407 18 57 39
(4332) (3411) (2834) (3706) (3069)
Maruteru 129 1005 35 53 129
(3130) (1602) (1679) (4065) (2215)
Aduthurai 57 19 6 50 15
(3995) (2959) (3077) (3834) (3039)
Cuttack 36 30 -12 -5 -;-3
(4428) (2956) (2814) (3272) (3527)
Patna 59 29 10
, (2792) (517) (-) (-) (2858)
lorhat 37 72 39 23 43
(3667) (4268) (3162) (3482) (3327)
Hyderabad 43 93 10 14
(2460) (3923) (1092) (2905) (-)
Ranchi 62 198 34
(1808) (3125) (-) (-) (2075)
Coimbatore 9 28 5
(3592) (3248) (2717) (-) (-)
Raipuf 5 9 -14
(3660) (5517) (3252) (-) (-)
Pantnagar 3 16 -8 16 15
(7735) (7309) (4522) (7129) (5620)
Karjat 1 22 13
(2622) (2128) (2300) (-) (-)
Kapurthala -3 5 -3 4 13
(3553) (4582) (3057) (4401) (2906)
Mean 108 194 14 45 76
(3903) (3660) (2883) (3947) (3208)
(Source; Progress report of .AJJ-India Co-ordinated Rice Improvement Project

Vol. 3-Kharif, 1969).
Lastly, attention may be directed to losses from pests during storage.

Wheat is highly susceptible to pests during the period from harvest to
consumption and the problem of its safe storage is becoming more serious
with increasing production. In fact, already wheat production is so high
that there is not enough space for storage. and large quantities of wheat
are in what is known as open storage under poly!hene covers directly on
REVOLUnOJl.! IN PEST CONTROL 215
the road.
Increasing Frequency of Pest Epidemics

The conclusions emerging from experimental fields are being equally
emphatically confirmed by the increasing frequency of recurring epidemics
in the country. The new agricultural strategy is only a few years old, and
during this short period there have been a number of serious. epidemics
in different parts of the country, e.g., the bug hopper and oth¥r pest epi-
demics in paddy crops in different states, the caterpillar pest epid,emic on
gram crop in c~ntral India, the Pyrilla epidemics in UP., and the cater-
pillar epidemics in rabi crops in Delhi and its neighbourhood. These
successivel epidemics are only pointers to the shape of things to come.
Lately, the Hispa ot' paddy, which used to be considered a minor pest,
has begun to cause serious concern. It appears that it has been responsible
for the decrease in the rice yield in Andhra Pradesh and has assumed
serious epidemic status in some'districts of U.P.
Reasons for rapidly worsening Pest Situation

The general explanation for the rapidly worsening pest situation is that
action and reaction are equal and opposite. Hence, whenever there is an
organized effort for increasing production substantially, there is also a
resistance to this effort (Fig. 11.1). To appreciat.e the precise mechanism
for the rapidly worsening pest situation with the introduction of H.Y.V.,
it is necessary to grasp the rationale depicted in figure 11.2.
Pest Conttol RevolutioQ preceded Agricultural Revolution

The first revolution in pest control took place during the forties when
the era of chemical control of pests was ushered in. Paul Muller was
awarded the Nobel Prize in 1948 for his discovery in 1939 of the insec-
ticidal properties of DDT. This had set in motion the revolution in pest
control technology, which preceded by 2 decades the agricultural revolution
of the sixties for which N. Borlaug was awarded the Prize in 1970. During
this period a large number of very potent pesticides came into the market
and enabled the potentialities of modern agronomy and high-yielding
varieties being realized. It was with the help of pesticides that the breeder
could pick up high-yielding genotypes (A and B in Fig. 11.2) which were
earlier being destroyed by pests to which they are very susceptible. Formerly,
the good effects of fertilizers were being mastered by higher pest infesta-
tion caused by the higher doses of fertilizer itself (Fig. 11.2).
SeC9nd Revolution needed

The most serious implication of the foregoing concept is that the
IN lliE ABSENCE OF PESTICIDAL UMBRELLA UNDER THE UMBRELLA OF PESTICIDAL APPUCA nON
d _ "'''HURING L EVE l d. -MANURING lEVEL- - -
Fig. 11.2. Relationship between protection effort and yield potential.
production potentials which have been experimentally observed and envi-

sioned under the umbrella of pesticides can be realized on country-wide
scale only under the umbrella of pest control (Fig. 11.2). At this stage a
snag comes in. Chemical control alone cannot take the country much
further. It is quite feasible to apply the highly effective pesticides at th~
experiment stage. It is also feasible to make use of these pesticides at the
stage of national demonstration. But when it comes to making large-scale
use of pesticides on a nation-wide scale it begins to have its own serious
limitations. The cost of pesticides needed annually would be about Rs 800
crore. This constitutes a huge input when the national income from agri-
cultural sources is only about Rs 8000 crore, and even if the nation some·
how managed this high input the adverse side-effects would go out of
control; these are health hazards, environmental pollution, evolution of
resistant pests, and upsetting the balance of nature.
The extremely deleterious results of such an upset of the balance in
nature are already being repented for in Nicaragua and Israel. It ~as
reported in an F AO Conference that during the early sixties the chemical
control of cotton pests in Nicaragua created, as expected, a very high yield.
The people felt that they had got the panacea. Ch.emical firms received
huge ordev and ,enthusiastically executed them. Eventually, the pests

became resistant, the balance of nature was upset; and by the late sixties
it became practically impossible to grow cotton economically. A much
higher decline in cotton yield is taking place in Israel (Fig. 11.3). We
35 . - . I~RAEl
I
.--- .. NICARAGUA
33
/
/'
31
29
27
25
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/
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17
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Fig. 11.3. Yield per hectare of cotton in Israel and Nicaragua during sixties.
should learn a lesson from such gigantic human follies.

Thus. it is quite clear that another revolution in pest control technology
is needed for taking the present green revolution to its next logical stage.
For bringing this about quite a sizable research investment is needed.
This revolutiop. has to be brought about step by step. First, very adequate
survelliance of the quickly changing pattern of pest .infestation has to be
carried out by a competent team of entomologists with adequate ecolo-
gical background. This has to be a research-cum-codtrol activity aimed
at exploring the various intricate ways in which the agricultural revolu-
tion is bringing about a revolution in pest intensity, and at nipping this'
trouble in the bud.
It is well known that man is a slave of circumstances. So, too. are all
other living organisms. But man, not knowing what he knows not often
interferes with circumstances and environment in such a way that he
becomes a slave of his own creations. This is what happened in Nicaragua
and Israel. and is happening in India. We must keep our fingers on the
pulse of nature and remain alert to its reactions to our interference. There
must be an ecological approach in planning crop sequences, particularly in
regard to pests and their control. Another direction in which revolution
in pest control technology is needed is what is known as Directed Integrat-
ed Control discussed in detail in Chapter 12.
Law of Limiting Factor
For optimum exploitation of an economic plant the law of limiting
factor is most important. Any of the factors influencing plant health can
become a limiting factor under certain specific conditions. Under Indian
conditions in recent years the varietal input along with its production
technology was acting as a limiting factor in wheat production. But for
pest control the input is acting as a limiting factor in rice. cotton and
sorghum.
Pest Control Basically Different from other Inputs
The pest control input is basically different from production inputs;
these go on increasing the yield in succession, and if any stage further
inputs are denied the further yield increase is arrested at that level; but
if the protection input is denied the yield ensured by other inputs registers
a steep fall.
Failure of National Demonstrations
The national demonstrations effectively show to the cultivator the
yield at the end level. i.e., after all the necessary production and protec-
tion inputs have made their respective contributions. These demonstra-
tions have replaced the prevailing gloom with hope. Now the difficulty
seems to be that when the final yield is shown to the cultivator practical1y
and actually in the field, and the package of practices responsible for the
high yield are given on paper, the, cultivator is not able to m~~e a proper
analysis of the contributions made by different production and protection
practices. The result is that pest control practices, with which he has not
been traditionally familiar, are likely to be easily overiookep. Hence, it
has now become very necessary to arrange national demonstrations speci-
ally meant to highlight the need of pest control practices. '
I
Wastage of Costly Inputs

Fig. 11.4 depicts how in the absence of proper matching. of pest control
and production efforts the high yielding but susceptible varieties act as
channels for the wastage of large amounts of other costly inputs like fac-
tory-produced fertilizers, which ultimately go 'into the stomach of pests
instead of human beings. The experimental data in suppbrt of this state-
ment ·are given in Table 11.2.
PEST CONTROL PROBLEMS ACTING AS BOTTLENECK IN PRODUCTION
Fig. 11.4. Pest control problems acting as bottleneck in production.

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REVOLUTION IN PEST CONl'ROL 221
Pest Control Researtb Needs not adequately realized

India's agriculture has been brought up against the China Wall of
insect pests mainly because the pest control needs, particularly the research
needs, are not adequately realized. There is a serious lacuna in pest con-
trol research which is acting as a drain across the road to progressl of
Indian agriculture, but it is covered with a very flimsy sheet of ignorance
which the press should help in removing. Some concrete examples may be
Storage problems. Plenty are the problems of plenty. The,' problems
of storage of wheat produced in annually increasing quantitie~ h~ghlight
how inadequately, the protection needs have been kept in view.
cited. /
TABLE 11.3. GRAIN YIELD (KG/HA) IN 'IR 8' AND 'TNI' AS COMPARED
WITH LOCAL VARIETIES WHEN INSECT PESTS WERE NOT CONTROLLED
Location 'IR 8' 'T (N) I' Local LocaL variety used
- --~-----
Warangal 505 656 720 'HR 35'

(4188) (4690) (3420)
Tenali 1227 672 2204 'MTU 20'
(4332) (3411 ) (3069)
,Maruteru 1371 145 969 'MTU 20'
(3130) (1602) (2215)
Aduthurai 2548 2479 2645 'CO 32'
(3995) (2959) (3039)
Cuttack 3251 3031 3620 'CR 1014'
(4428) (3956) (3527)
Patna 1750 400 2604 'BR 34'
(2792) (517) (2858)
Jorhat 2873 2472 2318 'CH 63'
(3667) (4268) (3327)
Ranchi 1116 1050 1550 'BR 34'
(1808) (3125) (2075)
Pantnagar 7479 6301 4876 'CH.4'
(7735) (7309) (5620)
Kapurthala 3657 4378 \ 2521 'Jhona 20'
(3553) (4582) . (2906)
(Source: Progress report of All-India Co-ordinated Rice Improvement Pro-

ject, Vol. 3-Kharif, 1969)
This table clearly indicates that if adequate matching of pest control

research with production goals is not to be done it is better not to advo-
cate high-yielding varieties like 'IR 8' and 'TNI' except at a few places in
the north where pest problem is not so serious.
Imbalance between Development Needs and Research Efforts

For proper, matching of various efforts to attain a particular goal of
production in a partIcular crop the best way is to carry out an exercise
at formulating practical strategy for increasing its production. This imme-
diately highl~ghts the relative importance of the various aspects on which
research efforts have to be concentrated. The following is the result of an
exercise on cotton in a seminar at the IAR!. C
Break-up of annual expenditure Break-up of annual expenditure in

envisaged in the strategy for in- the co-ordinated research project on'
creasing cotton production cotton
Total expenditure Total budget Rs 16.81 lakh
Expenditure on R.s 4.55 crore Pest control re- Rs 2.42 lakh
crop protection Rs 3.49 crore search budget
% expenditure on Expenditure for
crop protection research on pest
mainly on .control 76.7 control on the
basis of which
major portion of
76.7% develop-
mental budget is
t9 be spent 14.4
This highlights the imbalance between research efforts and develop-

mental needs of pest ~ontrol technology. The pest control recommenda-
tions of so~e of the states varied widely in their efficacy.
It may also be emphasized that the term 'plant protection is rather
unfortunate. The discipline concerned with the control of botanical disease-
causing organisms is very' different from that concerned with zoological
pests. Just as different production sciences cannot be combined at specia-
list level in anyone individual. different protection sciences cannot be
combined in anyone individual.
CHAPTER 12
IMPORTANCE OF PROTECTION RESEARCH IN TROPICS
THE impact of the green revolution in wheat brought about during the
late sixties has caused revolution in human thought and reaction all over
the world. One of these revolutions has been in introspection 'by agri-
cultural scientists themselves. How has the green revolution been brought
about now? How was it el~ding the scientists' efforts for more than three-
f
quarters of a century? How is it that the green revolution hag. so far
been possible only in _wheat? What is delaying rice revolution? A horde
of such questions ar~ agitating the scientists and the public in general.
It is desirable to re-emphasize the following:
1. A revolution in pest control technology was ushered ill during the
forties with the discovery of the insecticidal properties of! DDT. This
paved the way for the green revolution of the sixties (Fig. 11.2).
2. Any high-yielding varieties discoverd can perform well only 'under
the umbrella of selective control with pesticides. Since it is not possible
,to provide this on a country-wide scale, losses from pests are rapidly
mounting with the spread of the high-yielding varieties to the serious
detriment of the whole varietal programme (Fig. 11.2 and 12.8 and
Tables 1-5).
3. It has been possible to bring about the wheat revolution because
wheat is the safest crop in the field, for it is relatively free from risks of
pest infestation (Fig. 11.1).
4. Inadequate attention to pest control research is delaying the rice
rev@lution (Fig. 11.1, 12.1. 12.2. 12.3 and Tables 1 and 7).
S. The priorities and funds allocated to pest control research have
been so inadequate that the pests have begun to act as the most serious
bottleneck in increasing the yield. They were the deciding factor in the
country's success in its production eff<;Jrts during the sixties. There, was a
definite negative correlation between success in increasing production and'
productivity during the sixties and intensity of pest problem in particular
crops (Fig. 11.1).
6. Even on world basis the insect pests are so effective that the yield
of rice, essentially a tropical crop, is negatively correlated with the annual
average temperature of the principal rice-growing countries, and the only
explanation available for this negative correlation is that infestation is
relatively less serious in colder regions (Fig. 12.1). '
1. The foregoing revelations lead to an equation on ecological balance
in crop yields in which, the yield potential is equal to production potential
224 A,GRICULTURAL ENTOMOLOGY AND PEST CONTROL
YIELD .
Q HA lie
4 o·
27
... ....
- r------
26 +-ANNUAL AVERAGE TEMP.
r--
25
30,_
r--
24 ....
-
20 -
23
22
21
\ ~
10 - 20.-- r--
r- r- r------
r--
r-
1\
~
(/) <{
w 0 Z u;
Z
- _. z «
19 Q.. «_. « I-
UJ
z Z
Q..
N -ex: ~ (/)
0 >- « -ex:
..J -ex: 4: -
0 0:: :;;2 0
..J
-ex: (/)
Q..
~ z
~
0:: :::c z- -ex:
:::c
- !:: ~
-
Q.. !Xl I- !Xl Q..
18
o
Fig. 12.1. Rice yields in selected countries during 1967-1968.
minus destruction potential due to pests, etc., and this clearly inOlcates
that in the tropics protection research is more needed than production
research.
8. Results of some independent, statistically planned experiments on
the crops' response to important inputs, such as better variety, better
fertilizer, fungicidal treatment and insecticidal treatment, have clearly
demonstrated that the yield increases from pest control are much higher
than from any other single input (Tables 12.2 and 12.3).
9. It is well recognized that cost /benefit ratio is the highest for pesti-
cides among all inputs.
10. Y~t the agricultural ~cientists of the tropical counterparts in the
colder countries are laying major stress on production re~arch, particularly
IMPORTANCE OF PROTECTION RESEARCH IN TROPICS 225
R~
5500
IR8
5000
>-
I- KHARIF ' .... _'---<>
UI
----- -0- '
"«
>
4500
a:
0
u.
-
«
::x:
4000
(,?
~ 3500 KHARIF
.... ......
..... ""0
0
~
~
>- 3000
z
«
w
~ 2500
120 160 200

0 40 80
NITROGEN (KG/HAl
Fig. 12.2. Summary of N-response in dwarf indicas and tall locals (3 khan! and
3 rabi seasons AICRIP).
plant breeding. but protection research. particularly pest control research.

has been sadly neglected.
11. Hence. stagnation in production, except in wheat. seems to be' the
destiny for years to come unless the analysis presented in this contribu-
tion is seriously taken into consideration and made the basis for aIIocating
research priorities and funds.
INADEQUATE RESEARCH ATTENTION TO PEST CONTROL

DELAYJNG RICE REVOLUTION
The considerations on which pest control is regarded as the main
cause for the delay in rice revolution are as follows:
1. Phenomenal Increase in Rice Yield by Pest Control

Table 12.1 and Fig. 12.3 are spectacular. The recorded increase in rice
yield with pest control has been as high as 1005% in 'TN l' at Maruteru
and 729% in 'IR 8' at Warangal. At the same time, these increases have
varied widely at different centres depending either on intensity of pest
I ADDITIONAL YIELD
FROM PRODUCTION
8
1200
~
z
7 . 0
~
100 0
w
~
6 0
a:,
.ct
-
0.'
X
0
en 80 z
w 5
z a:
z w
>
0
t- O
0
4 60 ...J
W
0 >-
...J
W Z
>- 3 w
V)
40 <{
Z w
ex:
<{ 2 0
a: ~
(!)
~
20
1
0 0
:::> ...J ~
-,
...J
a: III ~ ~ a: ~ ...J a: Z
~ Z
~ ~
(!)
a: .... ...,a: 0 -,
a: z (!) 0.
a:
~
a.
~ ~ U Q. or::( '~
3:
LOCATIONS
Fig. 12.3. Effect of insect protection on grain yield of 'IR S' grown in 13 loca-
tions, khari/ 1968. (After Freeman and Shastri).
problem or on efficacy of control measures. All the same, the all-India

average increase due to pest control worked out to 108% for 'IR 8' and
194% for 'TN 1'. This highlights the extent to which pests can act as a
limiting factor in ~increasing rice production.
Revolution in Wheat but not in Rice
A definite negative correlation between success in productiop efforts
IMPORTANCn OF PROTECTION RESEARCH IN TROPICS 227
TABLE 12.1. INCREASE IN GRAIN YIELDS OF RICE BY MAXIMUM P.R01)lCTION
Grain yield (kg/ha)

Location 'IR8' 'TN I' 'W 1263' 'CR44-33' Local Local
variety
used
Warangal Treated 4188 4690 4084 2727 3420 'HR

I
35'
Untreated 50S 656 ,2245 1102 720
Difference 3683 4034 1939 1625 2700
Tenali
.
% increase
Treate'd
729
4332
615
3411
82
2834
147
3706
375
3069 'MTU 20'

Untreated 1227 672 2404 2412 2204
Difference 3105 2739 430 1294 865
% increase 253 407 18 57 39
Maruteru Treated 3130 1602 1679 4065 2215' 'MTU 20'

j
Untreated 1371 145 1241 2655 969

Difference 1759 1457 438 1410 1248
% increase 129 1005 35 53 129
Aduthurai Treated 3995 2959 3077 3834 3039 'Co 32'

Untreated 2548 2479 2908 2562 2645
Difference 1447 480 169 1272 29
% increase 57 19 6 50 15
Cuttack Treated 4428 3956 2814 3272 3527 'CRI 014'

Untreated 3251 3031 3210 3439 3620
Difference 1177 925 -396 -167 -98
% increase 36 30 -12 -5 -3
Patna Treated 2792 517 2858 'DR 34'

Untreated 1750 400 2604
Difference 1042 117 254
% increase 59 29 10
Jorbat Treated 3667 4268 3162 3482 3327 'Ch 63'

Untreated 2673 2472' 2282 2404 2318
Difference 994 1792 800 1078 1009
% increase 37 72 39 23 43
Hyderabad Treated 2460 3923 1092 2905

Untreated 1714 2035 994 2544
Difference 746 1880 98 361
% increase 43 93 10 14
Ranchi Treated 1808 3125 2075 'DR 34'

Untreated 1116 1050 1550
Pifference 692 2075 525
% increase 62 198 34
TABLE 12.1 (Coneld.)
Grain yield (kg/ha)

Location 2 3 4 5 6 7
Coimbatore Treated 3593 3246 2717

Untreated 3280 2530 2592
Difference 313 716 125
% increase 9 28 5
Raipur Treated 6360 5517 3252

Untreated 6077 5051 3775
. Difference 283 466 -523
% increase 5 9 -14
Pantnagar Treated 7755 7309 4522 7129 5620 'Ch 4'

Untreated 7479 6301 4892 6136 4876
Difference 256 1006 -370 993 744
% increase 3 10 -8 16 15
Karjat Treated 2622 2128 2300

Untreated 2589 1739 2033
. Difference 33 389 267
% increase 1 22 13
Kapurthala Treated 3553 4582 3057 4401 2906 'Jhona 20'

Untreated 3657 4378 3151 4219 2521
Difference -124 204 -94 182 385
% increase -3 5 -3 4 13
Mean Treated 3903 3660 2883 3947 3208

Untreated 2802 2353 2644 3053 2403
Difference 1093 1307 239 894 803
% increase 108 194 14 45 76
(All-India Rice Improvement Projects Progress Report - Khari/, 1969, Vol. 3,

Table 11.1).
and intensity of pest problems (Fig. 11.1) is clearly brought out by a

comparison of the percentage increase in production from 1961-62 to
1968-69, i.e., during the green revolution period. Of the 2 major cereals to
which greatest attention has been paid, only wheat has registered a pheno-
menal increase in ,production and productivity. Cotton comes between
wheat and rice in regard to the intensity of pest problem as well as the
success of our efforts to increase its production. Comparison of sorghum
and bajra sb9WS that the increase in yield has been much faster in bajra,
the pest problems in "which are much milder t'han in sorghum, which faces
serious pest problems, -
TABLE 12.2. INCREASE IN COTTON YIELD BY CONTROLLING INSECTS PESTS

(COMPILED BY R. A. AGARWAL - UNPUBLISHED)
In experimental fields • In demonstrational fields

S. State % increase S. State % increase
No. over control No: ov~r control
1. Madhya Pradesh 33.3 1. Madhya Pradesh (R) 52.9
1969-70 2. Gujarat (I) 38.9
2. Gujarat ('Digvijay') 228.8 3. Punjab (I) 99.8
1969-70 4. Haryana (I) 64.3
3. Sriganganagar 16.1 5. Maharashtra (I) 41.9
1967 - 6. MahaJ;"ashtra (R) 73.3
4. Sri ganga nagar 43.9 7. Andhra Pradesh (I) 17.~
1968 8. Mysore (I) 228.8
5. Punjab 1970 149.7
6. Tamilnadu 1969-70 463.0 Total average 68.6
7. Sirsa ('H-14') 25.8
(I) - Irrigated
1962-63
8. Sirsa ('320P') 42.8 (R) - Rainfed
1962-63 • Data recorded by the Directora te
9. Sirsa (1963-64) 17.7 of Cotton Development, Bombaly
10. Sirsa (1946-65) 55.0
11. IARI Centre,
Coimbatore (1962-63) 120.7
12. IARJ Centre,
Coimbatore (1962-63) 356.5
13. JARI, New Delhi 1969 249.1
14. IARI, New Delhi 1969 177.4
Total average 94.4
TABLE 12.3. A VOIDABLE LOSSES DUE TO SORGHUM PESTS

NEW DELHI CBYBRID-CSH-I ')
Grain yield from plots in which Grain yield from plots Loss %
pests were controlled (kg/ha) in which pests were not
controlled. (kg/ha)
Kharif 1965 5404 1212 77.6

KI,(;ri/ 1966 8258 1346 83.7
After Jotwani. M. G., Chandra, D., Young, W. R., Saxena, P. N. and Sukhani,
T. R. 1971. Indian J. En!. 33 (4): 375383.
TABLE 12.4. YIELD OF MUSTARD SEEDS (KG/HA)
Year Plots where aphid was Plots where aphid Loss %

controlled was not controlled
1st 930.0 112.2 87.9
2nd 530.8 256.7 51.6
lrd 219.9 63.5 71.1
From Pradhan el al. 1960. Indian Oi/seeds J. 4 (3); 125·i41.
faddy Yield bighel' in cooleJ; ~egioDs where Pest Problems are

aenerally Mild
Equally interesting revelation is made in Fig. 12.1. which gives per hec-
tare yield for different rice-growing countries. It is seen that although rice is
mainly a tropical crop its yield is highest in Japan and lowest in the
Philippines where th~ International Rice Research Institute (lRR!) has
been working for some years now. There is a definite negative correlation
between rice yield and annual average temperature of the country. and
the only rational explanation for this is that pest problems are
milder in cooler than in warmer regions. Thus it is again pests that seem
to determine the yield in different countries. The pest control experiments
carried out all over lndia (Table 12.1) lead largely to the same conclusions.
Paddy Yield hipel'in cooler Seasons (rabi crop) when

the Pest Problem ii generally Mild
The experience has been that the yield in rabi crop is higher than in
monsoon crop. Even the rice yield response to nitrogen is much higher in
rabi than in kharif; and this difference is much more marked in 'IR 8'
than in the local variety. It has also been explained on the basis of longer
photoperiod in rabi season than in kharif season when the sun is often
shut off by clouds. While the effect of photoperiod cannot be ruled out.
it is difficult to understand so much increase in a variety like 'IR 8' which
is claiOled to be insensitive to photoperiod. On the other hand. it can be
much better explained on the basis of more severe pest infestation during
khari! depressing the yield at various levels of nitrogen than during rabi
when the infestation is much milder. This also explains the higher diffe-
rence in the highly pest-susceptible variety 'IR 8' than in the local varieties.
Another fact that should be taken into consideration is that when pests
were controlled in kharif crop it was possible to get yields as high as
about 700 kg/ha at Maruteru (Fig. 12.2. 12..3). Tbis indicates tbat tbe
photoperiod was enough for producing 700 kg even though it might have
been less' than in rabi.
Nou,-entomologital ExplanatioDs and suggemollS do Dot

stand Scientific Scrutiny
'(i) Photoperiod difference. This has already been discussed under 3
and 4 above. r
(ii) Water management. It has also been explained that the low yield
of monsoon crops is due to difficulti~s in water management. While water
management is certainly more difficult during monsoon, the Ford Founda-
tion's assessment of prospects for increasing rice productivity In India
(March, 1971) has also concluded that "There is more uncertainty PI the
response of wet-season HYV rice than dry-season rice even under the
controlled conditibns of experimental stations."
(iii) Small farmers. ,The percentage of small farmers in traditional rice
areas is generally much higher than in other areas, and much concern has
been expressed that their response to new innovations may be slow; but
in the Ford Foundation report it was stated that !'There ate no really
important differences in the way farmers on different size faims are res-
ponsive to opportunities to modernise their production methods. (they) are
participating in the early adoption of new varieties where tliey
are active in increasing the use of fertilizers. related practices."
(iv) Productivity and acreage~ In India the productivity of rice is higher
in the north and north-west, whereas thel traditional rice acreage is in
the south and east. Unless productivity is increased in the south, or
acreage is increased in the north, rice revolution cannot come soon. Since
efforts to increase productivity in the south have been rather disappointing,
it is being suggested that the khari! acreage in the north should be allotted
to rice. This policy requires serious thinking and scrutiny. It is particularly
necessary to think over this whole question against the background of
the country's need for pulses. It may be better to ,allot the kharif acreage
of the north to pulses than to force the traditional rice-growers of the
south to replace their good crop of rabi rice with other crops.
ECOLOGICAL BALANOE IN CROP YIELDS
On the basis of the foregoing considerations it has been possible to
fonnulate a kind of equation on the ecological balance in crop yield
(Fig. 12.4). This equation depicts the interplay of the forces of production
and destruction. The production potential of a particular variety depends
on climate and other abiotic components of the environment, and what-
ever is produced as a result of the realization of this potential from stage
to stage of the crop gets reduced because of pests and other forces. These
forces constitute the reduction potential depending upon the biotic com-
ponent of the environment, i.e., the consumption potential of pests and
the ultimate yield of the crop is the resultant of these opposite forces.
PRODUCTION POTENTIAL
tiNDER CliMATIC COMPONE"
j REDUCTION POTENTIAL
UNDER BIOTIC COMPONENT
OF ENVIRONMENT i.e. =EnUAL
f
MINUS TO YIELD (Y)
Of THE ENVIRONMENT. CONSUMPTION POTENTIAL Yw-For. Warmer
(P) BY PESTS etc. regIons &
CRJ Seasons
Pw -For Warmer regions &
seasons .
1 Rw- For Warmer regions
seasons
&j Yc -Fe09ri~~o~r
r
Pc-For'Cooler regions & Rc- For Cooler regions & Seasons.
seasons Seasons
[
Pd=Pw-Pc
Pw - Rw =
.
Rd=Rw-Rc .
Yw In Warmer regions & Seasons
VPd<R~ (\
Pc - Rc = Yc In Cooler regions and seasons
Fig. 12.4. Ecological balance in rice' yield.
Further. both these potentials respond to changes in temperature, humi-

dity. rain, etc., but temperature is the most effective and dominant factor,
particularly for insect pests. On the whole, it can be confidently stated
that the co· efficient of change in insect activity and numbers as a result
of changes in temperature is much greater than in the case of plants.
The result is that in warm regions and seasons the reduction potential
due to pests increases much more than the production potential, and the
resultant yield is reduced. This view fully explains the data on the yield
of rice in different countries. These data cannot be explained on any
other basis. If the higher yield were based on better technology, it would
have gone up in the Philippines where both research and extension
activities of the IRRI are expected to provide the best technology. The
fact on the other hand is that the yield is quite high even in Italy. This
and the lower yield in USA than in J'apan cannot be easily attributed to
technological differences between countries (Fig: 12.1).
PROTECTION RESEARCH MORE NEEDED THAN PRODUCTION RESEARCH

Further, what applies to rice is also fairly applicable to other crops
grown in summer and monsoon in the tropics. The relative importance
of various problems of agricultural research is basically different in tem-
perate' and tropical regions. In temperate regions the most important
point for research concentration is to increase the production potential of
the equation mentioned above. What we in the tropics grow in winter
they grow in summer; and what we grow in summer and monsoon is
genenllly difficult to grow in temperate regions. Hence, in the temperate
IMPORTANCE OF PROTECTION RESEARCH IN TRoPICS 233
regions the emphasis has been on plant breeding and detection of geno-
types in summer and monsoon crops of the tropics for growing them in
the temperate regions. This approach has made it possible to grow even
a tropical crop like rice in as cold a country as Japan.
The situation in the tropics. on the other hand. is such that practically
everything grows here in plenty in one season or the other. The crops
grow. as also the organisms which claim the crops; and as crop produce
grows more vigorous the pests. disease-causing organisms. aqimals and
human beings grow. Hence, the main aim requiring highest rt1search em-
phasis in the tropics is to reduce the reduction potential of the equation.
i.e., to reduce the growth of various organisms which claim the major
portion of the' crop and crop produce. Yet agricultural scientists of the
tropical regions have, mostly followed the lead of those in the temperate
1800
:1716
1700
1600.,
1532
1500
0
..J
w 1400
>=
1300
1200
1100
1000
6 4 3 5 2 1
TREATMENTS
Fig. 12.5. Insecticidal trial against the grain and pod borer, Heliothis armigera
(Source: New Vistas in Pulse Production) ..
regions and have. therefore. provided much higher support to production

research than to protection research. This explains the continued failure
of agricultural scientistsi right from the beginning of the century in making
any spectacular increase in crop yields. except in wheat.
Experimental Proof
The yieid increases due to pest control reported in Tables 12.1 to 12.4
(Figs. 12:3 and 12.5) clearly show that the crop's response to pest cqntrol
is much more than can be expected from any other single input such as
better variety. increased fertilizer dose, or fungicidal treatment. In Table 5
it will be seen that pest control alone brought about more increase in'
yield than fertilizer application in each of the 3 years in the improved
variety 'Rai 6342' and in 2 out of the 3 years in 'Laha 101'. Table 6,
of cotton experiments, shows even more spectacular increases due to pest
control. These data also show that yield increases due to pest control are
much more in improved varieties. Also it is well recognized that cost-
benefit ratio is generally highest for pesticides than for any other input.
Table 12.7 shows that at many places it is harmful to sow high-yielding
varieties if proper control measures are not to be taken.
TABLE 12.5. RELATIVE INCREASES IN YIELD OF RAI BY TWO INPUTS

SEPARATELY AND IN COMBINATION
Yield q/ha
Varieties 'Rai 6342' 'Laha 101'
Treatment
Years
1965-66 66-67 67-68 Ave. 65-66 66-67 67-68 Ave.
Control 7.5 10.3 22.1 13.3 6.9 9.5 20.8 12.4
Plant protection 15.0 15.4 30.3 20.2 12.0 12.9 21.7 15.4
measures
(pest control only)
100 kg N. 14.6 9.7 29.1 17.8 11.8 10.9 27.6 16.8
SO kg
P 2 0 5 50 kg
KjlO/ha
Combination 19.2 18.8 35.1 24.4 17.1 16.6 29.8 21.2
of 2 and 3
After Kinra, K. L. and Rao, S. B. P. 1971. Data presented at a symposium held
at Regional Research Station (IARI), Kanpur, in March 1971.
The following analysis will show that while the first pest control
revolution was needed to visualize the green revolution in wheat another
pest control revolution is needed to realize the green revolution in other

crops.
TABLE 12.6. EFFECTS OF FERTILIZER, INSECTICIDE AND FUNGICIDE ON

YIELDS OF COTTON IARI, NEW DELHI (1970 AND 1971)
% increase in yield over control

Variety Fertilizer Yield Ferti- Fungi- lnsee- Fungicide
dose (kg/ha) lizer cide@ ticide +
control (b) Insecticide
1970
'H-14' N-O, p-o 879 0.0 7.2 43.9 54.2
N-60, p-o 1086 23.5 18.8 66.3 74.7
N-120, P-60 ~021 28.4 ~25.1 61.1 72.1
N-120, P-60 1070 21.7 29.1 73.9 69.3
,
'Lockett N-O, p-o 290 0.0 81.3 28!).3 337.2
4791' N-60, p-o 410 41.4 144.4 4'03.7 443.4
N-120, p-o 452 55.8 104.8 432.7 459.3
N-120 P-60 492 69.6 120.3 438.6 520.6
1971 (c)
'H-14' N-120, P-60 45 33.9 33.0 166.7 166.7

'PS-I0' N-120 P-60 270 14.8 7.4 133.3 125.9
'H-14' N-O, p-o 854 0.0 0.0 0.0 6.0

N-60, p-o 588 0.0 0.0 18.9 35.9
N-120, P-O 583 0.0 0.0 30.4
N-120, P-30 677 0.0 0.0 44.4 45.1
'Lockett N-O, p-o 167 0.0 37.1 420.9 299.4

4791' N-60, p-o 219 31.1 146.1 520.3 426.9
N-120, p-o 224 34.1 0.0 445.5 498.8
N-120, P-30 281 68.1 77.8 480.2 420.3
@ = 8 sprayings with 0.2% copper oxychloride

b = 8 sprayings with inseoticides (M onocrotophos, Andrin and Carbarryl)
c = Bad season for cotton crop.
This table clearly indicates that in case adequate matching of pest

control research with production goals is not to be carried out it is better
not to advocate high-yielding varieties like 'IR 8' and 'TN I' except at
a few places in the north where pest problem is not so serious. Compare
tnese data with those of Table 12.1.
TABLE 12.7. GRAIN YIELD (KG/HA) IN 'IR 8' AND 'TN l' COMPARED WITH
LOCAL VARIETIES WHEN INSECT PESTS WERE NOT CONTROLLED
Location 'IR 8' 'TN l' Local Local variety used
Warangal 505 656 720 'RR 35'

Tenali 1227 672 2204 'MTU 20'
Maruteru 1371 145 969 'MTU 20'
Aduthurai 2548 2479 2645 'CO 32'
Cuttack 3251 3031 3620 'CR 1014'
Patna 1750 400 2604 'BR 34'
Jorhat 2673 2472 2318 'Ch 60'
Ranchi 1116 1050 1550 'BR 34'
Pantnagar 7479 6301 4876 'Ch 4'
Kapurthala 3657 4378 2521 'Jhona 20'
(Source: Table 11.1 of the Progress Report of All-India ~oordinated Rice Improve-
ment Project, Vol. 3 - Kharff, 1969).
The pesticide umbrella provided in the first revolution has to be

converted during the second revolution into an effective pest control
umbrella with proper integration of various methods of pest control in-
stead of depending only on pesticides, which have shown seriou~ limita-
tions under large-scale use.
LIMITATIONS OF LARGE-SCALE PESTICIDE USE
High Cost
The cost of the pesticides needed annually is Rs 800 crore. This is a
huge input. Besides, the adverse side-effects will go out of control; these
are health hazards. environmental pollution, evoilltion of resistant pests,
and upset of the balance of nature.
Health, Hazards
The pesticide limitation which has been agitating the public mind
most is the potential hazard to human health anel environment. In 1956
the IARI started work on pesticide hazards; and in the sixties the ICAR
constituted the Thacker Committee, whose report led to the Insecticide
Act of 1971. This Act is mainly to make rules and regulations for keeping
the pesticide hazards to the minimum. Under this kct an Insecticide
Board has come into existence.

It is the considered opinion of scientists that pesticides as su~h do
not involve any serious hazards. What actually causes hazards is the
misuse of pesticides. A recent reviewer categorically stated that there
exists not one authentic record of death caused by the use of any pesti-
cide applied according to the prescribed instructions.
A rather disturbing information reve.aled during the delibeTations of
the Thacker Committee was that some ad hoc post-mortem fat samples
from Delhi had shown DDT content to be as high as 30 ppm, which
was considered ,the highest in the world. It has been shown that unscru-
pulous traders in foodgrains find it convenient to add a little DDT in
their stock rather than use other recommended methods of keeping their
stock safe. Such misuse of pesticides is bounel to decrease considerably
once the Inspectors of the Insecticide Act begin their operations against
such malpractices.
A few remarks are necessary for the well-meaning intelllgentsia whose
concern -regarding pesticidal activities are based on news emanating from
various developed countries. Countries in the temperate zone have to
concentrate mainly on increasing production potential; hence they can
, afford to be fastidious. But our main emphasis has to be on minimizing
the reduction potential, i.e., on pest control. Some experts have rightly
remarked that if one has to choose between death by starvation and
death by insecticidal contamination, the latter may be preferable. Hence,
India should have its own well-considered policy regarding the use of
pesticides depending on our own conditions and should not be misled by
what in other countries is discussed in books, conferences and newspapers.
Even in USA, Senator Jamic L. Whitten, chairman of the House of
Representatives Appropriations Subcommittee for Agriculture for 15 years,
published (1966) a book entitled 'That we may live', This is meant mainly
to remove the public scare resulting from Rachel Carson's 'Silent Spring'
and "unjustified public fears of those invaluable chemicals called pesti-
cides".
Upset. of Balance in Nature
This is the main limitation of pesticide use. When large-scale pest
control is carried out repeatedly with the help of pesticides alone, the
pests become resistant to the pesticides used, and their parasites and pre-
dators get eliminated. This upset of the balance in nature creates a very
difficult situation, as in Nicaragua and IsraeL
THE SOLUTION
Since the subject of pest controi has been a constant victim of neglect
it is difficult to offer very practical solutions of the various problems.
We have reached a stage when it is possible to visualize clearly that the

area requiring highest priorities for research concentration in Indian
agriculture is the protection of crops, particularly from insect pests.
Further success will depend on the amount of investment and the sagacity
with which it is made use of. However, according to the present theoretical
knowledge the solution of the problems created by the limitations, dis-
cussed above lies in what is known as Directed Integrated Control. T)1e
epithet 'integrated' signifies that instead of depending solely on chemical
control we should suitably incorporate all the methods of pest
control into an integrated schedule. The scientific rationale for such in-
tegration lies in the fact that there is in every population individual
variation in every biological characteristic. Thus, the individuals of any
insect population vary in their susceptibility to insecticides. Hence. when
an insecticide is applied some indviduals escape death because they
I require for their kill a dose higher than the one applied. In this way a
resistant population gets screened out, and it is no longer possible to
control it with an insecticide dose which was effective earlier. Thus, the
dose has to be progressively increased, till eventually the insecticide
becomes ineffective. The mechanism by which such a situation arises also
contains the basic principle of integrated pest control.
It may be very pertinently asked why the pest becomes resistant and
its parasite becomes annihilated. The principle involved here is that an
individual which is resistant to the chemical may not be capable of finding
a host at the reduced host density brought about by chemical control.
Hence. while the pest undergoes only I screening (by the insecticide), the
parasite has to face 2 screenings, first by the insecticide and then by the
reduced host density. Thus, while the pest which is subjected only to
chemical control becomes resistant, the parasite which has to face inte-
grated pest control succumbs. Hence, to control the resistant pest popu-
lation it is necessary to use another operation to which it may not be
necessarily resistant. When one control method is alternated with another
the development of resistant strains is avoided or delayed. Moreover.
when chemical control is alternated with non-chemical control the elimi-
nation of parasites and predators is appreciably reduced and the upset
of the balance in nature is avoided. Further, integrated control reduces
the total amount of insecticide used. and the hazards are correspondingly
less. Hence it is necessary to develop non-chemical control measures as
weI],
The integrated control is quite interesting in its principle and appeal-
ing in its philosophy, but very difficult to evolve. Researches have to be
carried out on every minute detail of the life economy of each insect
pest and its natural enemies. These are bound to be costly and time-taking;
but there is no altemat~ve to deeper scientific probes. -
CHAPTER 13
INTEGRATED PEST CONTROL
THE definition and the principle. as agreed upon during the first session
of the FAO panel of experts on integrated control, are as follow~:
Definition of Integrated Control

Integrated control is defined as a pest management system that in the
context of the a~sociated environment and the population dynamics of
the pest species. utilizes all suitable techniques and methods in as compa-
tible a manner as possible and maintains the pest populations at levels
below those causing economic injury. In its restricted sense, it refers to
the management of single pest species on specific crops or in particular
places. In a more general sense, it applies to the co-ordinated ,management
of all pest population in the agricultural or forest environment. It is not
simply the juxtaposition or superimposition of 2 control techniques
(such as chemical and biological controls) but the integration of all
~uitable management techniques with the natural regulating and limiting
elements of the environment.
Principle of Integrated Control

Integrated control derives its uniqueness of approach from its emph.
asis on the fullest practical utilization of the existing mortality and
suppressive factors in the agro-ecosystem. To achieve this goal crop pro-
tection must be guided by the following 2 principles :
1. Pest control should be developed and applied in the total environ-
ment. The pest popUlations are managed in such a manner that existing
limiting and regulatory factors are exploited to the fullest extent possible
and without disturbance to the regulation of other pests. This principle
defines the underlying philosophical approach to integrated control.
2. Additional mortality or regulatory factors are introduced into
the environment at appropriate times to maintain the pest population at
levels below those causing economic injury. These levels should be deter-
mined in terms both of the foreseeable crop loss and of the economics
of crop production and marketing. This principle defines the goal of the
integrated control system.
Strategy of Integrated Control

The strategy for an integrated approach should include efforts (a) to
reduce the seriousness of the pest problem by evolving crop varieties as
resistant to insect pests as possible, (b) to reduce pest infestation by man-

oeuvring the ecology of the crop including introduction of exotic parasites.
(c) to nip the infestation in the bud by such mechanical campaigns as
collection of egg-masses, affected shoots, etc., and (d) to resort to a
suitable combination of chemical and biological methods of control
including some of the modem sophisticated techniques.
In the past the practice has been to recommend control measures
separately for each pest. Those recommended for some important pests
of paddy, in one of the latest official publications (Pl. Prot. Bull. 11, No.
1-4. 1963), are as follows.
Pest Control measures
Rice bugs 1. Dusting 5% BHC, 15-20 lb/acre

Leptocortsa acula Th.,
L. varicomis F. 2. Spraying 0.25% DDT, 60-80 gal/acre
Green jassids I Spraying 0.25% DDT, 6(}80 gal/acre
ephotettix bipllnctatll~ Fb. 1. Spraying 0.15% DDT, 60-80 gal/acre
N. apicalis Mots. 2. Dusting 5% DDT, 15-20 lb/acre
White jassid - do-
Tettigella
spec/ra Dist.
Black bug - do-
Scotinophora
lllrida Burm.
Mealy bug Spraying 0.5% BHC or 0.08% Malathion
Ripersia oryzae Gr. or 0.05% Parathion, 60-80 gal/acre
Fulgorid bug Dusting 5% BHC, 15-20 lb/acre
Nilaparvala lllgens Stal.
Rice root aphid Dipping the seedling in and spraying the
Trtraneura hirsuta B. soil around the plant with 0.05% Parathion
Paddy stem borer 1. Ploughing up and destroying the stubble
Schoenobius after harvest
incertllias Wlk. 2. Collection and destruction of egg clusters
in nursery plants
3. Dipping seedlings in 0.1 % DDT suspen-
sion before transplanting
4. Spraying 0.025% Parathion or 0.03%
Endrin in the field, 60-80 gal/acre
Swarming caterpillar 1. Dusting 5% BHC or DDT, 15-20 lb/acre
Spodoptera
mflUritia Boisd. 2. Spraying 0.25% DDT
INTEGRATED PEST CONTROL 241
Rice caseworm 1. Dusting 5% BHC or DDT, 15-20 lb/acro

Nymphula depunctalis Guen. 2. Spraying 0.25% DDT, 60-80 gal/acre
Armyworms 1. Dusting 5% BHC or DDT, 15-20 lb/acro
Cirphis unipuncta Haw., 2. Spraying 0.25% DDT, 60-80 gal/acre
C. albistigma M.
Paddy leaf roller 1. Dusting 10% BHC, 15-20 lb/acre
Cnapna!ocrocis
medianalis Guen. 2. Spraying 0.25% DDT, 60-80 gal/acre
Semilooper 1. Dusting 5% BHC, 15-20 lb/acre
Psalis securis Hbn.
Rice borer Spraying 0.03% Endrin, 60-80 gal/acre
SCirpoQhaga innotata Wlk.
Pink borer 1. Spraying 0.03% Endrin, 60-80 gal/acre
Sesamia inferens Wlk. 2. Dusting 5% BHC or DDT, 16-20 lb/acre
Striped rice borer Spraying 0.03% Endrin, 60-80 gal/acre
Chilo suppressalis Wlk.
Paddy skippers Dusting 5% BHC, 15-20 lb/acre
Parnara mathias Fb.
Gall fly .1. Light-trapping at the time of sowing for
Pachydipiosis oryzae a fortnight
Wood-mason 2. Spraying 0.25% DDT or .02% Endrin
60-80 gal/acre
Rice hispa 1. Dusting 5% BHC, 15-20 lb/acre
Hispa armigera Olivo 2. Spraying 0.25% DDT, 60-80 gal/acre
H. aenescens Ball'. ~. Clipping off and de!ttroying leaf tl~ in
nurseries when transplanting seedlings
Blue beetle 1. Dusting 5% BHC, 15-20 lb/acre

Leptispa pygmaea Baly.
Root weevil Mixing\ 5% BHC dust in soil before sowing,

Echinocnemus oryzae M. 30-40 lb/acre
Rice grasshoppers 1. Dusting 5% BHC, 20-30 lb/acre

Hierogiyphus banian Fb.
H. oryzivorus Carl., 2 Spraying 0.02% Aldrin, 60-80 gal/acre
Spotted grasshopper - do-

Au/arches mi/iarls B.
Phadka grasshopper - do-

Hieroglyphus nigrorepletus Bol.
Surface grasshoppers - do-
Oxya ve/ox Fb.,
o. heidentata Wlk.
O. multidenta.ta Wilt.,
Ai/opus sp.
Acrida exaltata L., - do-
A. turritae L.
Paddy eelworm - do-
Ditylenchus anJfusta But.
In the above table as many as 40 control measures are recommended

against 26 pests of paddy. It is left to the farmer or to the local
extension worker to select at the moment and on the spot one of the
methods listed against the pest which happens to be doing more serious
damage to the crop at the time. Although at first this appears to be quite
a rational thing to do, there are some practical disadvantages in this
kind of approach. First, the general tendency is to select the most handy
out of the most effectively advertized methods leaving out the most ra-
tional one for which there happens to be no provision in advance.
The result is that some methods which would have proved more effec-
tive and cheaper are not tried, and pest control is reduced to the applica-
tion of an insecticide, the distributing or manufacturing firm for which
has the most effective propaganda organization )n the region. The second
major disadvantage of the existing practice is that for each pest there.
are several recommendations and the result is that the farmer and the
local extension worker are faced with an unmanageably large number
of pest control recommendations for each crop. Neither can they make
advance provision for each of the long list of pesticides, nor are they able
to make a judicious selection. The result is confusion leading to a sense
of frustration and helplessness, which in turn leads to drift and escapism,
allowing the pests to have their full share and to be regarded as unavoi-
dable evils to be tolerated. The obvious remedy is to attempt a kind of
rational synthesis of the diverse recommendations against the diverse
pests of each crop so as to evolve rational integrated control schedules for
growing pest-free crops of each important commodity. Of course, this
schedule should not be too rigid, and it should have a reasonable amount
of built-in flexibility for necessary modifications from year to year and
from place to place. The synthesis has to be done in the following stages.
1. Pest-wise integrated schedule
2. Crop-wise integrated schedule
3. Re~on-wise intewated schedul~
INIEGaATEp PEST CONTROL 243
4. Large-scale campaigns.
Pest-wise Integrated Schedule

The general attitude in the past has been to select the most suitable
of the control measures for each pest species. To this end, there has been
quite a good deal of comparison of the different methods of ~ontrol. The
attitude of thinking entomologists is to work out the integration of as
many of these methods as necessary for the effective control of anyone
or a group at pest species of a crop. Such a synthesis is often quite
rational. A gbod illustration is the control of internal feeders like the
stalk borers of maize and sorghum. The importance of these pests is
increasing every day with the introduction of hybrids and_ other improved
varieties. The methods of control of such internal feeders are far from
satisfactory: Once the larvae enter the plant tissue, they get beyond the
reach of contact insecticides and stomach poisons, and often the insec-
ticides used, instead of killing the pests, continue to kill their parasites
which otherwise would have destroyed a proportion of the larvae inside
the stems. More than 30 species of parasites have been recorded from
the different stages of the stalk borer of maize and sorghum, and a dozen
of them have been found to attack the immature stages of this pest in
different parts of India. Some of them are reported to be effective in
reducing the popUlations of the borer where climate and other conditions
are favourable. The biology and habits of the important larval parasites
have been studied in the laboratory, and techniques for mass breeding
have been developed for a few, e.g., Glyptomorpha deesae and Tricho-
gramma evanescens minutum.
Thus, studies have been in progress to develop a combined ecological
and toxicological approach to solve this problem. and now the stage has
reached when an integrated control schedule on the following lines can
be confidently suggested. The moths of these pests lay their eggs on the
plant, and the larvae hatching from the eggs crawl on the plant surface
for a specified period, which can be determined by actual observations
on each species. Eventually the larvae enter the plant tissue and get out of
the reach of ordinary insecticidal dusts and spravs. Hence, it is sUlzgested
that spraying against these first-stage larvae should coincide with the
hatching of the eggs. which can be easily determined with the help of a
biometer (described below)_ The snravinl! shol ld be CMried out il1~t when
1
the earlv hatching is expected: variation in the hatching period should

determine the persistence of the insecticidal formulation and it~ strenl!th.
which has to be selected. The strenttth of thi' in~ecticioi'~ h::l~ al<:o to hi'
such a~ to ensure that the first-stal!e larvae will nick 1m the If'th::ll oo"e of
inc;ecticide durin!!: the few minutes when thev rr::lwl over the nlant snrface.
If these spe<;ifica~ion~ ~re taken care of, most of the newly hatched larvae
will be killed before they are able to do appreciable damage. Persistence

of the insecticide will last for the :opecific period needed to kill the first-
stage larvae and no more. Parasites of the different stages of larvae and
pupae can be released at a time when the stage against which they are
effective reaches its peak; this can be determined with the help of the
biometer. These biological control operations against larvae and
pupae can thus be integrated with chemical control directed against
first-stage larvae. Lastly, the peak emergence of the moths can also be
determined with the help of the biometer, and at that stage we can use
the latest technique of controlling the development of the next generation
with the help of the sterile mal y technique. Efficiency of the sterile male
technique is maximum at the lowest population density of the pests.
Hence. it is quite rational to reduce the pest population from its peak
at the egg stage to the lowest level at the adult stage by means of chemical
and biological control. the efficiency of both of which is higher at higher
population densities of the pest, and then to use the sterile male technique
at the low population level of the pest at which it is most effective.
Por the integrated control schedule it is necessary to take cognizance
of some recent developments in entomological research. These are (i) in-
vention of the biometer which can pointedly indicate the peak period of
such phenomena as larval hatching, pupation. emergence of adults, and
egg-laying, (ii) evaluation of a technique of bioassay for determining the
relative toxicity to newly hatched larvae coming in contact with insecticide
films for a minimum period (separately determined for each species) for
which these larvae invariably crawl about on plant surface before boring
in. (iii) the sterile male technique, and (iv) the latest thinking on biological
control. The bioassay results indicate the insecticide and the strength to
be used against particular species, and the biometer indicates exactly
when chemical control, biological control, or sterile male technique should
be applied. Some of these topics have been discussed elsewhere in this book.
Crop-wise Integrated Schedule

The main study required in this integration is to find out the widest
common spectrum in the control of different pests and to work out the
schedule of operations, each expected to control the most important pests
infesting the crop at any moment. In fact, whenever a chemical control
operation is carried out it does bring about varying degrees of control of
different pests present at that time, although the operation might have
been only against the most important or spectacular one. This spectrum
can be enlarged and the control of different pests improved substan-
tially if the insecticide and its time of application are chosen by taking
most of tlie pests into consideration. Even more integration effort is
needed in non-chemical control. 'Por example, in the case of sugarcane
iNtEGRATED PEST CONTROL 245
the collection of egg-masses of both Pyrilla and top borer is suggested

and removal of dead hearts is also recommended in the case of several
borers. It is best to collect the egg-masses of top borer and of Pyrilla, the
shoots affected by top borer, and the dead-hearts caused by the. stem and
root borers in one campaign, particularly in the early season of the crop
when infestation of all these pests starts.
Philosophy of Integrated Control

Integrated pest control is a novel concept, but its theoretical simplicity
masks its nove}ty. Even jn recent years, although there has been quite a
good deal of ;talking and writing and the idea has been catching fast,
comparatively few concrete suggestions have been coming forth. The
result is that some of those entrusted with the direction of practical pest
control feel this concept to be utopian. The existence of such diametrically
opposite views, the immense potentialities of this concept. the practical
feasibility of evolving integrated schedules for protecting Ii, number of
crops, and the inevitability of the approach under Indian conditions, have
created the urgent scientific necessity for carefully and forcefully analysing
the philosophy and feasibility of integrated control.
The concypt of integrated pest control was originally meant to bring
about a compromise between chemical control and biological control.
Before the advent of the chemical age in pest control there was no serious
conflict between these two approaches. But following the discovery of
DDT there was such a rapiq influx of new insecticides that these
poisonous chemicals began not only to bring about indiscriminate anni-
hilation of insect fauna but also to kill the very science of entomology.
Some agricultural scientists began to feel that it was not necessary even
to identify the pest trouble. Such unscientific ideas, however, did not last
long and basic phenomena, such as the development of resistant strains
of insects. upsetting of balance in nature, increasing bazards to human
beings and livestock, began to exert their effects seriously. All the same,
at the level of the individual cultivator it is still quite a difficult job to
drive home a balanced view. For any orchardist or grower of a valuable
crop it is difficult to decide not to apply a pesticide in the hope that
parasites and predators will keep the pest under check. In the apple
orchards of Kashmir even experiments on biological control of San Jose
scale became difficult because orchardists would not oblige by desisting
to spray their orchards. Thus. it is desirable that entomologists should
concentrate on formulating rational integrated control schedules in which
both chemical and biological control find their due place, each exerting
its_ optimum beneficial effects. It is feasible to effect not only a compro-
mise between chemical control and biological control but also a workable
integration of all the methods of control of different pests affecting the same
crop or commodity as well as those affecting different crops in a particular

area. Thus, the concept of integrated co:dtrol is rapidly bringmg us to
the threshold of reliable unified approach to our pest problems as a whole.
Recent Need for Integrated Control
There are so many instances in the world where dependence on che-
mical control alone first created health hazards and upset the balance in
nature, and then began to fail in controlling the pest. It is to forestall
these serious limitations of chemical control that integrated pest control
is now being seriously considered. Another recent consideration necessi-
tating integrated control is that agriculture has been following a pro-
gressive trend towards narrowing the germ-plasm in the field. Agriculture
originated in the practice of monoculture in place of mixed vegetation.
thereby 1eading to the origin of insect pests. Now these monocultures of
mixed germ-plasm are being replaced with those of narrower and narrower
germ-plasm in the form of pure varieties. These are further leading to
adverse effects so far as pest ~ontrol is concerned. These adverse side-
effects of modem agriculture can be minimized to a large extent by
replacing single-approach pest control with diversified pest control, which
should be properly integrated and scheduled.
Genetic-cum-statistical Basis for Integrated Control

The novelty of this method lies in the basic principle on which the
superior efficacy of integrated control has to depend. It is a fairly well
known genetic-cum-statistical principle that in any natural popUlation
there is individual variation in almost all biological characters, and the
frequency distribution of a particular character generally follows a normal
curve, a majority of the individuals occupying the central region below
this curve and small minorities lying at two extreme ends. Thus, in the
diagram (Fig. 13.1 B) illustrating the distribution of the individual
susceptibility and resistance (IS and lR) of an insect 'population to an
insecticide there is a small proportion of highly susceptible individuals
(IS) occupying a small portion of the area on the extreme left and also
a small proportion of highly resistant individuals (IR) on the extreme
right, with majority of the individuals of normal susceptibility and
resistance in the major portion of the middle region. Hence, when the
population is treated with the dosage (D 1) of the insecticide, practically
the whole population will get wiped out except the portion (IR) which,
if all otheJ,: conditions remain normal, will multiply in the next generation;
and if this is repeated for a number of generations a resistant strain will
emerge. In another diagram (Fig. 13.1 A) illustrating another distribution
of individual capacity of the same insect population to escape predator
attack. There will be .a few high susceptible individuals (PS) becoming
t·
w
z
::J
ow
....a:
'* IS
R
INSECTICIDE DOSAGE D, [PREDATOR POPULATION DENSITY D2

A CAPACITY TO ESCAPE PREDATOR AnACl(
B
Fig. 13.1 (A and B). Genetic cum statistical basis for integrated control.
prey to a very low population density of the predator occupying a small

area in the extreme left of the diagram, and also a few requiring quite a
high density of. predator population to succumb occupying the extreme
'right-hand area of the diagram (PR); majority with normal susceptibility
to predators will occupy the major portion of the middle area. If this
insect population is exposed to a predator population density (D 2), the
entire population will succumb except (PR). It has to be appreciated that
individuals belonging to PR group (Fig. 13.1 A) and those belonging to
the group (IR) in diagram B are expected to be randomly distributed in
Fig. 13.1 A. Hence, when the population is treated by the insecticide dose
D I most individuals of group PR are likely to be killed, and when the
population is exposed to the predator population density D 2 practically
all group IR will succumb. Thus, if the same population is exposed
successively to both the treatments, i.e., chemical control and exposure
to predator population, both IR and PR groups will be practically wiped
out and development of a resistant s~rain will be considerably delayed.
If the 2 insecticidal applications are replaced with 1 insecticidal applica-
tion and 1 exposure to predators, the amount of insecticide used will be
considerably less, with the least health hazards and upset of the balance
in nature.
There are some advantages of integrated pest control schedules.
1. They fit well in the national economy, and are desirable for
developed countries, these are but mentable for developing countries. Our
pest control activities at present are mainly based on the applications of
chemical pesticides, large quantities of which have to be imported. Our
plans for plant protection are also based mainly on pesticides. The
expenditure runs into crores of rupees even when only 1 or 2 pesticide

applications are envisaged. On the other hand, experiments on hlgh-
yielding varieties show that many more pesticide applications are required.
This means that as the acreage under high-yieldmg varieties increases, it
will be impossible to meet the demand for pesticides if we continue to
depend wholly on them. Thus, a time has come when integrated pest
control is not only advisable but also inevitable.
2. They offer more efficient and cheaper control. In integrated contr:ol
schedules an effort is made to utilize various methods of control including.
use of pesticides, but at times and in some cases it is feasible to nip the
trouble in the bud even by mechanical control, e.g., destruction of egg-
m;:J.Sses of some pests early in the spring. This mean~ much saving of
pesticides, money and of foreign exchange. Very often, pesticides are
applied when the pest has already done considerable damage. This can
be easily taken care of in an integrated schedule of which the successive
steps need to be taken only if the previou~ steps have not been vecy
effective.
3. They avoid upsetting the balance of nature. Chemical control has
often been reported to upset the balance in nature. at times leading to
the upsurge of new types of pest troubles. The seriousness of mites as
pests of several crops has resulted from the use of DDT in many parts
of the world. It is confidently expected that such adverse side-effects will
be much less with integrated control schedules.
_4. They will avoid or delay considerably the development of pesti-
cide resistant strains of pests. Development of pest strains resistant to
pesticides has been the most disconcerting experience of pest control
scientists and technicians. From all that is known to date it is certain
that this phenomenon is the result of killing out of the susceptible portion
of pest population, leaving the resistant individuals to multiply free from
any competition. However, it is not likely that individuals resistant to
one method of control are equally resistant to another method, and it is
possible that those resistant to a pesticide are susceptible to a biological
control agent. Hence, if biological control follows chemical control, the
undesirable screening effects of the pesticide application can be nullified.
In this way the more the methods of control are diversified and practised
as cOl;nponents of a well-planned integrated schedule, the less will be the
chances for the development of resistant strains of pests.
5. They will considerably minimize residue hazards of pesticides.
It is obvious that in an integrated control schedule the use of pesticides
will be consideably reduced. Hence. the pesticide residue hazards will be
automatically minimized.
6. Pesticide industry will get a boost instead of being harmed. Des-
pite what has been stated in the foregoing paragraphs, the pesticide
iNTEGRATED PEST CONTROL 249
industry will not lose but actually gain as a result of integrated control
coming in vogue. In conclusion, therefore, it may be stated that a time
has come when we should stop debating whether integrated control should
be accepted or not, and instead try to formulate integrated control
schedules with existing knowledge. The main aim should be clearly under-
stood as planned diversification of control methods so as tQ checkmate
the pest, as in warfare.
Feasibility of Integrated Control for Pests of Certain Crops
1. San Jdse scale In Kashmir. In recent years highly effective in-
secticides have come into the market for the control of this· cosmopolitan
pest; but in Kashmir the problem continues to cause serious concern,
and the pest attacks such a large variety of fruit and forest trees that it
is not possible to carry out chemical control on all. The result is that
chemical control proves to be only a temporary palliative; for the in-
festation is very heavy and, therefore, the good effect of chemical control
is soon lost, so that the orchards have to be treated again and again
during the same season. In order to overcome this difficulty it is highly
advisable and feasible to integrate chemical and biological control for
keeping this pest economically under control. It is advisable to treat the
orchards of valuable fruits like apple with suitable insecticides and to
follow it up with intensive biological control. In areas like Kashmir it is
quite feasible to rear parasites like Prospaltella in very large numbers
throughout the winter season in specially heated rooms wherein the para-
site population can be rapidly multiplied 10 to 20 times during each
generation of about 4 weeks, and then to flood the areas infested with
the comparatively small overwintering population of San Jose scale on
uneconomic plants around the fruit orchards. In this way the infestation
from outside the orchards can be effectively checked, and fewer insecti-
cidal applications resulting in reduced number of operations will be
required in the orchard itself. This is a very fertile field for an effective
and fruitful integration of chemical and biological control applied simul-
taneously in time but separately in space.
2. Maize and millet crops. Considering the vast variety of potential
dangers from· insect pests to millet crops, the policy of millet! growers
should be to evolve a cooperative routine schedule of control operations
somewhat on the following lines.
(i) After harvesting, the stalks of millet plants should be used up as
soon as possible and in any case before the end of the winter season.
The stalks harbour various stages of borer pests.
. (ii) For the same reason, stubble should be dug out and destroyed.
It is advisable to plough the fields just before the beginning of summer
to uproot the stubble, which should be collected and used as fuel or
burnt, and to disturb and expose the egg-masses of grasshopper pests

and pupae of caterpillar pests to the sun and to predators. The bunds of
the field and the ground around trees and fences, which specially harbour
these pest stages, should be scrapped, ploughed or hoed. Grasses on the
bunds should be used up or destroyed along with the.. pest stages.
(iii) For ~eed. a pest-resistant variety should be selected.
(iv) Sowing should be at a high seed rate so as to enable uprooting
of pest-infested plants without decreasing the plant population below the
optimum level.
(v) The crop should be inspected from the earliest stages for the
appearance of (a) dead-hearts due to borers of any kind, (b) moths
emerging from pupae which have been under diapause and so have
escaped operation (ii), (c) egg-masses laid by these pests, and (d) other
external feeders. In the same round of operations the dead-hearts. the
egg-masses and the sluggish moths should be collected and destroyed.
(vi) Chemical control operations should be carried out according to
the nature and intensity of the pest. In the case of sucking insects a
strong contact insecticide should be used; bUi if there is also an infes-
tation of leaf-eaters. such as caterpillars and grasshoppers. then a more
persistent insecticide should be used. Grasshoppers and cutworms can
also be controlled by poison baiting.
(vii) The precautionary operation (i) to (v). if carried out properly
on a large enough scale and on a co-operative basis. should prevent the
borer population being built up beyond tolerable limits. If. however, for
some reason these precautionary measures are not practicable and inten-
sive chemical operations are yet feasible, a suitable persistent insecticide
should be applied to poison the newly hatched larvae of the borers during
the few minutes when they are crawling on the surface of the plant before
boring into its tissue. For this purpose insecticide sprays or granules should
be used just before the peak period of hatching determined by means of
a biometer for the species which it is most necessary to control. This
chemical control should be integrated with other methods emphasized
earlier.
3. Sugarcane pests. The main characteristics of some of the serious
pests to be considered in devising measures for their control are as follows.
(i) The top borer and Pyrilla lay eggs in large prominent masses,
which can be easily located from a distance against the green background
of the leaf-surface. This makes these pests quite amenable to mechanical
control, i.e., collection and destruction of egg-masses. This is quite a
feasible proposition under Indian conditions where batches of boys and
girls can easily clean up large areas. Supervision by visual survey is also
easy.
(ii) All 3 types of borers, viz., top-borer, stem-borer, and the root-
INIEGRAlED PEST CONTROL 251
borer. cause characteristic dead-hearts in the early season of the crop.

Collection of such affected shoots along with the associated insects can
be combined with the campaign mentioned under (i) above. The top borer
attack can be easily detected in all stages of the crop.
(iii) Both Pyrilla and borer pests are often heavily parasitized in
their different stages.
(iv) All types of borers lay eggs on plant surface, and their larvae
crawl on the surface for a few minutes b.efore getting into the tissue out
of reach of insecticidal chemicals. Hence, the different operations of the
control schedule have to be precisely timed, as follows:
(a) Sugarcane planters should be made fully conscious of the fact
that sugarcane IS a graminaceous crop and, that wild graminaceous plants
growing in the vicinity and on the bunds can serve as alternate hosts for
sugarcane pests. Hence, keeping the area clean of such plants as well as
stubble of the previous season's crop is a prerequisite for successful sugar-
cane cultivation. This is a precautionary measure which shquld not wait
for the appearance of any pest or even for sowing of the crop.
(b) For seed, a pest-resistant variety should be selected, if available.
(c) The next precautionary measure to be adopted at the time of
sowing is against termites and .root borers, especially in areas where they
'cause troubles year after year. This measure consists of chemical treat-
ment of the furrows in which the sets are to be sown, so that termites and
young larvae of root-borer trying to approach sugarcane plants may get
a lethal dose of insecticide.
(d) If the termite attack is becoming serious despite the above-men-
tioned precaution or because that precaution has not been taken, then
charging the irrigation water with a repellent or a suitable insecticide has
to be resorted to. This operation need not form a regular item of the
control schedule.
(e) Full preparation and all arrangements should be made to orga-
nize campaigns on as large a scale as possible to nip in the bud the
attack of borers and of Pyrilla as s90n as egg-masses and dead-hearts
begin to appear in the young crop. 1:hese should be collected and des-
troyed. This step is the most crucial operation for success against sugar-
cane pests. Depending on the experience of the past years, one or the
other pest species may be expected to make its appearance every year.
Hence this operation should form one of the routine items of sugarcane
cultivation and should be continued as long as necessary.
(f) Egg-masses of both Py'rilla and top borer are at times heavily
parasitized. Therefore, arrangements should be made to conserve the
parasites by keeping the egg-masses in wire gauze cages from which only
the -minute parasites can fly out but the Pyrilla nymphs and borer
larvae are either trapped in the cage or made to fall in a pail of kerosinized
water.
(g) If the foregoing operation has been carried out effectively and on
a sufficiently large scale so that its beneficial effect is not upset by migra-
tion of insect pests from neighbouring areas, then the problem should be
significantly reduced in magnitude in the main part of the crop season. All
the same, the farmeu has to keep vigil and adopt control measures as and
when required. It may be necessary to collect and destroy the egg-masses
of top borer and Pyrilla and the tillers affected by borers in the maiQ
season also, but generally chemical control operations will have ot be'
resorted to. Pyrilla can be controlled by a number of contact insecticides,
but it is advisable to time these operations with the help of biometer so
that they coincide with the peak hatching of borer larvae and before they
bore into the cane. To this end it would be well to use a contact insecti-
cide of suitable duration of persistence; thereafter effort should be made
to integrate this chemical control with biological control. In case mite
infestation also appears, it would be desirable to use a chemical which
controls both insects and mites instead of a specific chemical like sulphur
which kills only the mites.
4. Pests of paddy. There are 3 major characteristics which should
be kept in view in evolving a rational integrated control schedule. The
first 2 are rather disadvantageous. First, as many as 3 crops a year are
taken from the same field, and this unbroken continuity creates ideal
conditions for the development of paddy pests. Secondly, there is contin-
uity in space also because the land used for paddy is in many places not
fit for other crops. In other words, paddy cultivation is continuous both
in time and space, and this is very conducive to pest multiplication. The
third characteristic is very helpful. Often the paddy seed is first sown in
nurseries and later the seedlings are transplated. Since the acreage of this
nursery stage is much smaller than that of the transplanted crop, the control
measures are bound to be less costly and more effective in the nursery
stage. Further, at the transplanting time practically each seedling is indi-
vidually handled, and with a little extra care it should be possible to
eliminate the affected seedlings, clip the prominent egg-masses of certain
pests, and treat the seedlings against certain other pests. Lastly, a number
of p~ddy pests are multi vol tine and consequently the flying adult stage
comes again and again in the same season. Univoltine species such as
grasshoppers are capable of migration. It is, therefore. necessary that
farmers are made fully aware of the fact that control measures carried
out in a few fields will not yield the desired results unless all the farmers
pool their resources and carry out the control operations on co-operative
basis.
(a) Regular items of the schedule
The following items should be included in the regular agricultural
operations for successfully growing the paddy crop.

(i) As a number of rice pests, e.g. Spodoptera, gall fiy, gundhy bug.
Hispa, breed on grasses during the off-season, it should be a definite policy
of rice growers to keep their area free from wild grasses.
(ii) As preparatory operations for rice cultivation, propeJ.:' ploughing
and destruction of stubble are necessary to destroy pupae and caterpillars
of stem-borers. pupae of §warming caterpillars, and eggs of grasshoppers.
In areas where,' grasshopper trouble is likely to be serious the scrapping
of the bunds should also be attended to as a routine.
(iii) For seed, a 'pest-resistant variety should be selected, if available.
(iv) Light-traps should be set up right from the beginning not only
to attract and destroy the moths but also as indicators of the activity of
these pests in the field so that timely control operations can be carried
out. It should, however, be borne in mind that sometimes the area in the
immediate vicinity of light-traps gets more severe infestation because the
moths that evade the trap can cause severe damage round about the light-
trap. To prevent this, such arc!ls should be spccially treated with suitabll!
'persistent contact insecticides.
(v) Right from the early stage, the fields should be inspected regularly
for the appearance of prominent egg-masses of stem borer and swarming
caterpillar, and as soon as they appear they should be collected and
destroyed. It should be noted that destruction of egg·masses is the surest
way of controlling these pests, particularly the stem-borer, the later stages
of which are much more difficult to destroy. Care should be taken to
observe, collect and destroy the characteristic dead-hearts caused by stem
borer, silver shoot caused by the gall fiy, and the tunnels in leaves con-
taining immature stages of rice Hispa. Thus, vigilant farmers working
in co-operation can confidently hope to nip in the bud the infestation of
these -5 important paddy pests.
(vi) The operations mentioned under (v) should also be carried out
when the seedlings are collected together in convenient bundles before
transplanting the seedlings. Also at this stage the seedlings can be thor-
oughly treated with a suitable persistent insecticidal dip for which Para-
thion emulsion for systemic action against stem-borer larvae has been
suggested by some.
(vii) The swarming caterpillar and cutworms can also be checked
from entering the field from outside by means of a trench around the field.-
(b) Control operations to be carried out as and when necessary
(i) The crop and bunds should be treated with a suitable insecticide
(say, 5% BHC dust) whenever any of the surface feeders like gundhy bug
and grasshoppers increase in numbers bevond tolerable limits.
(ii) Cutworms and swarming caterpillar can also be destroyed by
flooding the field. A Httle oil (6 Htres/hectare) or better a suitable insee-
ticidal solution may be mixed with the flood water. Even nymphs of
gundhy bugs can be made to fall in this oily water by moving a bamboo
lightly across the plants.
5. Pests of pulse crops. Considering the different important pests
of pulse crops it should be clear that farmers should keep themselves pre-
pared for the following 4-pronged attack against insect pests.
(i) A thin top layer of the soil should be treated with a strong per-
sistent soil insecticide. particularly in areas subject to cutworm attack year
after year. This can be done just before or after sowing, or the insecticidal
dust can be raked in even at a later stage depending on the time when,
cutworm attack starts in the region. This treatment will not only poison
the cutworms hiding in the soil during the day but also take care of
certain othel1 pests, such as gram pod borer and gujhia weevil, which
are somewhat alike in this respect.
(ii) For seed. a pest-resistant variety should be selected, if available.
(iii) As far as possible, as soon as the pest attack starts, a common
campaign may be organized for collection and destruction of such pests
and infested material as can be easily spotted, e.g. leaves attacked with
leaf miners, or even pea stem borers, twisted and deformed pods attacked
by the red gram pod borer, and cutworms detectable by freshly cut
shoots dragged into the soil.
(iv) If the trouble persists in spite of the first 3 operations or because
they have not been carried out, then the crop should be treated with a
penetrating formulation of a good persistent insecticide which acts both
as a contact insecticide and as a stomach poison. This treatment should
take care of almost all the pests. It kills gram pod borers mainly by con-
tact effect while they are eating the developing gram in the pod from
outside, poisons cutworms when they feed on treated shoots, has a lethal
effect on adults laying eggs in the crops, reaches leaf miners within their.
galleries, and controls pests like aphids. The main snag in this approach
is that since leafy parts of leguminous crops are often used for both
human and animal consumption, extreme care is needed to avoid residue
toxicity hazards.
6. The locust problem. In recent years the main emphasis has heen
on pesticides in dealing with problems of locust control. Our studies at
the IARI have resulted in 2 new approaches for dealing with the locust
problem. One of these can be profitably and immediately integrated with
the pesticide, and there is a prima facie case for integrating the other
one also after some further studies. The first is the use of deterrent-cum-
repellent material present in the kernel of neem (Azadirachta indica)
seed. A 0.1 % suspension of this seed kernel in water when sprayed keeps
the crQP safe from locust for 2 to 3 weeks (Pradhan et al., 1963. Bull.
Reg. Lab., 1: 149"151). Therefore, in a locust invasion the farmers can
INIEGRATED PEST CONTROL 255
ensure the safety of their crops with the neon kernel while the govern-
ment agencies take the usual action of wiping out the locust swarms by
insecticidal chemicals. Further, our theoretical studies have shown that
locust outbreak is delimited both in time and space by biotic agencies,
and we hope that after some further studies it may become possible to
use these biotic agencies to nip the locust outbreak in the bud. Thus,
we can use the integrated approach (a) for dealing with locust population
explosion in the outbreak areas with the help of the enemi,bs of locust,
(b) if the locust outbreak does occur to ensure the safety of the crops
with deterrent. sprays of neem seed suspension, and (c) to wipe out the
locust swarm by insecticidal chemicals.
7. The phadka grasshopper. Chemical control of grasshoppers like
phadka is well known and well established, but the chemicals are generally
applied to the infested crop. Our field ecological studies at the IARI
have shown that (a) the eggs qf this species which have been hibernating
during winter and summer hatch about 10 days after the first good mon-
soon showers, (b) these eggs are found in appreciable numbers not in
cultivated fields but on the sides of bunds and in uncultivated areas near
about the cultivated fields, and (c) the young ones of the grasshopper
after hatching remain confined to the bunds and uncultivated areas for
about 14 days before moving into cultivated fields. Hence, we have made
the recommendation that, instead of waiting for the crop to be infested
and then treating it" insecticidal application shoud be made as soon as
the first monsoon rainfall take~ place not only on cultivated fields but,
also on the bunds and the uncultivated areas in the neighbourhood. In
this way the pest can be controlled with much less expenditure and
before it inflicts any damage. This is a good example of integrating
ecological and chemical approaches for the control of a pest, like phadka.
It is likely that similar integration is possible in other species too.
8. Storage pests of grain. Another example illustrating rather
spectacular success of the integrated approach is the Pusa bin for grain
storage. The efficiency of this structure in keeping stored grain safe from
pest infestation and deterioration is based on the integration of measures
against different pests. The study at the JARI (Pradhan, 1968 Indian 1.
Ent. 30: 94-103) of the requirements, habits and idiosyncrasies of different
pests has shown that (a) certain species, such as the weevil Sitophilus
oryzae, are very susceptible to reduction in moisture content of the grain,
so much so that it has become a common belief that if the grain is kept
reasonably dry its safety from pest infestation can be ensured - a belief
embarrassingly belied by pests such as the khapra beetle which can breed
in_grain with very low moisture content; (b) certain species, e.g. khapra
beetle, are very susceptible to reduction of oxygen tension and there is
also a strong belief that the safety of the grain can be ensured by storing
256 AGRICULTURAL ENIOMQLOG¥ AND PEST CONTROL
it under air-tight conditions - another belief belied by species like the

grain weevil mentioned above which can breed under very low oxygen
tension; (c) a number of complications arise from moisture migration
within the grain mass, especially if the container is made of a good con-
ductor of heat - so, for making a storage structure a poor conductor of
heat should be selected. Owing to these different requirements of different
pests. recommendations for safe storage of grain have been quite confusing
and contradictory.
In devising the Pusa bin all these considerations were taken care of.
lts walls are made of mud or unburnt brick, and there is embedded in
them a film of polythene which is reasonably impervious to water vapour
and gases. When grain like wheat is dried to less than about 10% moisture
content and stored in the Pusa bin, it is unable to absorb atmospheric
moisture during the monsoon, and the oxygen tension inside is reduced
by respiration of grain and any insects present. So neither pests like
khapra nor those like grain weevil can breed in this bin. Internal moisture
migration is reduced to the minimum by the wall being made of mud
or unburnt brick. Thus, Pusa bin represents an excellent example of
successful integrated approach in pest control.
9. The giant African snail. Our concern about this pest has been
fast increasing. Studies at the IARI have provided some excellent leads
to developing an integrated chemical-cum-biological control of this serious
non-insect pest. The difficulty in integration of chemical and biological
controls of insect pests has been that generally the pest as well as its
enemy is an insect and by and large it has been quite a difficult job to
find such selective chemicals that will kill the pest but not its enemy. In
the case of the giant African snail also the predators mostly talked about
and recommended are 2 species of predatory snails, and the chemical
and biological approaches to the control of this pest have been conflicting.
But recently the effectiveness of 2 arthropod enemies of this molluscan
pest has been spectacularly demonstrated. These arthropods, viz. several
species of hermit crab of the genus Ceoenobita and a species of the
millipede Orthomorpha. are not affected by the common molluscicide,
the metaldehyde. Hence, the path seems to have been well cleared for
an effective and useful integration of chemical and biological controls in
the case of this pest. This also opens up a vista for putting greater
emphasis than before on exploring the use of biological control agents
belonging to distant zoological groups. There seem to be enough reasons
to believe that non-insect enemies of insect pests are much more effective
under natural conditions than hitherto supposed.
Use of stomach poisons in integrated control. Before the advent of
DDT. ins.ecticides used to be rather sharply divided into stomach poisom.
contact poisons. and fumigants. This classification was obliterated with
INrEGRATED PEST CONTROL 257
the discovery of a number of modem insecticides, many of which act

both as contact poison and as stomach poison, and some also as fumigant.
At first this combination of these 2 or more types of action was quite
welcome because it improved the effectiveness of the insecticide consi-
derably. Later on the same combination of actions began to prove irksome
because the contact action was found to kill the parasite adults. Hence,
efforts were made to avoid the contact action of DDT with. some sort
of coating on its particles during formulatipn (Ripper et al., 1948) Nature,
161: 484). At this stage it occurred to us that instead of trying to coat
DDT particles it might be useful to try the older stomach poisons like
basic lead arsen~te which have no contact action. Consequently, some
work has been in: progress with promising results for integrating biological
control with chemical control of the coconut leaf caterpillar with lead
arsenate.
Large-scale organized campaigns. There are points in favour of and
against co-operative farming, and it is considered to be one of the most
controversial subjects under Indian conditions. All the same, for pest
control co· operative endeavour is essential. The most spectacular pecu-
liarity of insects is that they have ·an effective and efficient capacity for
free flight. There is no other form of life which is so minute in size and
yet able to undertake active flight for a long distance. Bacterial and other
spores are carried about with the wind but passively and at the mercy
of the wind. Besides the development of wings the secret of this extra-
ordinary capacity for free flight in insects is that they have developed
very effective water-proofing mechanism against dangers of desiccation.
which acts as a limiting factor in the evolution of smaller size in other
free-living terrestrial and aerial' fauna; for the smaller the size of the'
animal, the more the exposed surface area per unit body weight and the
more the dangers of desiccation of active aerial existence.
Insects have evaded this general principle by developing special anti-
desiccation mechanisms. The implication of this functional and structural
superiority of insects over other forms Qf life is that they cannot be easily
controlled unless the control operations are carried out over as large an
area as possible and also as simultaneously as possible. Otherwise the
good effect of control operations is diluted or nu11ified bv the migration
of insect from the surrounding untreated areas. Hence, it is absolutely
necessary that progressive farmers should try their best to carry with
them their less enlightened colleagues in pest control operations. In
any case. special efforts are necessary to omanize lame-scale pe!<t control
campaigns as depicted dia1!rammaticallv in Fi!!. 13.2. ~llch nlanning is
very necessary in case of eradication campai$ms. but it should also form
the bi!<is of large-scale control camnailms for emmrin!! ontimum henefits.
The idea is that a suitable compromise has to be worked out between
Fig. 13.2. Spatial strategy of pest control campaign.
intensiveness and extensiveness of each individual campaign, depending

on the nature and distribution of the pest, its mode of dispersal, crop
season. etc.
The strategy depicted in Fig. 13.2 consists of selecting a spot or spots
(A, B, C, etc.), at which the pest is specially serious, as centres or sub-
centres of pest control operations. Thereafter, the required intensity of
the operation has to be carefully decided as the first step, and its econo-
mics has to be calculated. Then, depending on the available resources a
circle of manageable dimensions should be drawn with the chosen spot
as the centre, and during the first season control operation should be
concentrated within that circle to bring down the pest population to the
desired level (or for eradication, if this is the aim)., As a result of this
intensive operation during the first year it is expected that the pest
problem in the following year will be much milder within the treated
circle. Hence, next year the control operation can be extended to a bigger
circle around that of the first year. In this way the area under controt can be
increased year after year, and it can be ensured that the control achieved
in the inner treated area is not nullified by migration of pests from the
surrounding untreated area. Where more than one spot has been selected
to start the campaign, as shown in Fig. 13.2, the expanding concentric
circles around the different spots (A, B, C, etc.) will in due :course meet
one another, and the areas left untreated between these circles can be
taken care of' in subsequent years. Thus, a much bigger circle of the
treated area will emerge. In this way the effect of the control campaign
can be saved from -being diluted or nullified by migration. The area has
to be treated in circles instead of squares or rectangles to ensure uniform
borer effect around the treated area.
These are some of the spcial requirements of pest control as distinct
from other operations of crop husbandry. A pest control campaign is
best organized by such government of other agencies as can ensure 100%
coverage of the area. This can be accomplished only with insurance
against pest infestation or with what may be called Plan Health Scheme,
somewhat on the lines of the Central Government Health Scheme, so that
some levy can be charged and freedom from pest reasonably assured.
Otherwise, the good effect of piecemeal control is bound to be diluted
or nullified, depending on the areas over which it has been carried out
and on the migration propensities of the pest concerned.
The idea of crop insurance is slowly catching public imagination. It
is not, however, urgently and inevitably needed for pest control. I ad-
vocated it in as early as 1958 in the All-India Entomological Research
Workers Conference.
EPIWGUE
From the evidence that is accumulating it is becoming amply clear
that insects, which inherited the earth some 500 million years before man,
still hold effective proprietorship of the land, and man has been exercising
only a sort of tenancy right. The human tenants till the land and produce
the crop, and the insect proprietors appear on the scene and claim the
major share. To abolish this interspecific zamindari system, humanity has
to fight a major war. However, proper appreciation and understanding
of this global phenomenon is the first step in the correct direction. The
strength, strategy. and secret of success of insects lie not in an ostentatious
display of terrorizing power but in the dodging humility of their minute-
ness. They seem to have hit upon the philosophy that the humble shall
inherit the earth!
INDEX
A Atherigona soccata, 91, 92

Attacus at/as, 9
Acamhomia sp., 90 Aulacophora /miecollis, 180
Acanthopsyche, 118 Aularches miliaris, 241
Acestus, 115. 122 Avena sp. 91
Acrida exaltata, 242 sativa, 90
turritae, 242 Azadirachta indica, 254
Adelostoma, 115, 122
Adesmia, 109, 116 B
Adhesive trap, 56, Bagrada cruciferarum, 180, 189
Aenelomia yaria, 88 Bait trap, 56
Agrlotes sp., 81 Belonogaster, 121
Agrotis ypsilon, 87 Belostoma grande, 9
Aldicarb, 98 Bembix, 111
Aldrin, 204 Bemisia tabaci, 83
Alfalfa blossom midge, 89 ber, 32
Alfalfa weevil, 89 BHC, 203
Amaurosoma sp., 97 Biograph, 159
Ammi majus, 120 Biometer, 156, 175
visnaga, 120 Biotic formula, 4S
Amount of damage, 58 Biotic potential, 42
Analysis of biotic potential, 47 black bug, 240
Anaulocnemus sp., 90 black cutworms, 87
Anoxia segetum, 87 Blastophaga peenes, 21
vil/osa, 87 'Blepharidopterus angulalus, 60
Anthia, 109, 121 Blissus, leucopterus, 112
Antlddium, 118 blue beetle, 241
Apantelesl 26 Bombus, 20
flavipes, 24 Bomby/ius, 121
Aphe/inus mali, 23 Bombyx mori, 33
Aphidius, 26 Brachycerus, 109, 121
smithi, 23 Brachyrhnchus, 109
Aphis, 56 Bracon greeni le/royi, 24
craccivora, 189 Bruchus ana/is, 189
Apiculture, 31 Busseola rusca, 86
Apis dorsata, 20, 27 Butea monosperma, 32
florea, 20, 27
indica, 28
c
Appendix A, 147 Cactoblastis cactorum, 27
Appendix B, 150 Camponotus, 120
Aprotis, 84 Cataglyphis, 119
Arecanut, 205 Chemosteriiants, 197
Argasidus, 115, 122 Chilades galba, 117
Assessment of losses due to insect pests, Chillies, 205
75 Chilo zone/lus, 24, 88, 91, 92
Organisational aspects, 94 Chi orion, 111
Percentage increase in yield, 93 Cimbex humera/is, 113
T<:chnique of estimating losses, 76 Cirphis
262 AGRICULTURAL ENTOMOLOGy· AND PEST CONTROL
albistigm a,241 E
unipuneta, 241 '
Cnaphaloeroeis medianalis, 241 Earias, 69, 70, 129
Coeeinel/a septempunclata, 26, 113, 116 fabia, 42, 118, 128, 131, 159, 162, 187
suppressalis,241 Echinocnemus oryzae, 241
Coconut, 205 Eeiton sp., 117
Coelioxys, 121 Effect of temperature on the dynamics of
Coffee. 205 insect development, 123
Coleoptera, 9 Biograph, 159
Collection of marked insects, 59 Biometer, 156, 175
Colonies of social insects, 59 Biophysical processes involved in
Colos~al damage by insec5,ts development,132-
Comparison of different types of samples, Comparison with previous view, 136
61
Con/arinia medicaginis, 89 Comparison of estimated and observed
sorghico[a, 88, 92 value, 175
Copper fungicides, 203 Curve of developmental indices, 142
Coreid bug, 90 Curve fitting and testing of goodness
Coreyra cephalonica, 22, 186, 187, 188, of fit, 126
189 Deduction of eqaution, 123
Criticonoma, 117 Effect of variables and constant tem-
Crocisa, 121 perature, 140
Crop samples, 57 Estimation of developmental periods,
cydixa p/yehora, 89 161
Cylas formicarius, 180 Estimation of number of generations,
,D 166
Eleodes, 107, 112
Dacty[opius indicus, 27 Emergence cages, 58
spp., 27 Emoasca lybiea, 83
tomentosus, 27 Endosulfan, 98
Dalopins sp., 84 Endrin, 203
DDT,203 Entomology as an applied science, 4
Diabrotica longicornis, 86 Epizestia, 60
Disulfoton, 98 elu/ella, 58
Dity/enehus angus/us, 242
Epilogue, 260
Dominance of insects, 6
Episus, 121
Developmental characteristics, 14
Erebills agripinna, 9
High fecundity and controlled repro-
duction,15 Eremiapizila braueria, 120
Popula tion of individual insects, 9 Eriosoma lanigerum, 23
Protective adaptation and device, 16 Estimation of insect population, 52
Secrets of insects inherent strength, 9 Cumulative effect on insect popula-
Size of individual insects, 8 tion,68
Specificity of feeding, 15 Method of computing pest incidence,
Structural perfections, 10 66
Zenith of evolution, 17 Mode of sampling, 61
Dorymjlnnex,119 Nature of sample, 54
Drosieha, Number and size of samples, 62
mangiferae, 180 Stage to be counted, 52
sp., 186,187, 188 Etiella zinckenella, 91
Drosophila, 2 Euneta, 118
Duerosia ane/hifolia, 120 Eupodo/us, 109
INDEX 263
Euproctis lunata, 189, 193 I
Eurychora, 109
Eurygaster integriceps, 41 Importance of protection research in
Ellrytoma amygdali, 113 tropics, 223
Ecological balance in crop yield, 231
F Inadequate research attention to pest
control, 225
Felt loss, 95
Limitation of large-scale pesticide
Ficus spp., 32
use, 236
Fixed volume or area of earth, 57
health hazard, 234
Foeniculum Jlulgare, 120
high cost, 236
Forestry, 205
the solution, 237
Fruit trees, 205
upset of balance in nature, 237
Fulgorid bug, 240
Paddy yield higher in cooler regions
Fundamental vs, ~pplied entomology, 2
where pest problems are mild, 230
G Phenomenal increase in rice yield, 225
Protection research needed more than
Gall fly, 241 production research, 232
Gall midge, 81 experimental proof, 234
Geophanus, 109, 111,115,122 Revolution in wheat but not in rice,
Giant African snail, 256 266
Glyphiterix cramer, 39 Insects,
Glyptomorpha deesae, 243 Colossal damage by, S
Gnorimoschema opercrelella, 97 Development characteristics, 14
Goniomma, 120 Dominance of, 8
Gossypium barbadense, 83 High fecundity and controlled repro-
Grc.phipterus, 109 duction, 15
green jassids, 240 Protective adaptation and device, 16
groundnut, 204 Secrets of inherent strength. 9
GlIlerma melanapus, 90 Size of, 8
Specificity of feeding, 15
H Structural perfection of, 10
Haplodiplosis equatris, 81 Zeni th of evolution, 17
Heliothis armigera, 84, 91 Insects adaptations to arid conditions,
haemorrhoidalis, 117 105
obsoleta, 54 Adaptations in developmental
Hemiptera, 9 processes, 112
Heptachlor, 204 hibernation and aestivation, 113
Hessian fly, 81 homodynamic and heterodynamic,
Heteroderaavenae, 81 114
rostochinensis, 84 Behaviour adaptations, 114
Hieroglyphus banian, 241 for avoiding enemies" 120
nigrorepletus, 80, 241 for ensuring security, 114
oryzivorus, 241 for securing sustenance, 118
Hispa, 253 site preference, 115
aenescens, 241 Morphological adaptations, 105
armigera, 241 adaptations in internal anatomy,
Holcomyrmex, 120 110
Hordeum sp., 96 hea vy well-developed integument,
Human records and natures preservation, 108
61 long legs, 109
264 AGRICUL TURAL ENTOMOLOGY AND PEST CONTROL
organs for capturing prey, 110 Lepidoptera, 9

organs for cl.!aning, 110 Leptispa pygmaea, 241
organs for digging into sand, 109 Leptobylemyia coarctata, 96
oval or compressed body, 109 Leptocorisa acuta, 240
wingless or fused elytra, 108 varicornis, 240
Physiolo~ical adaptations, 111 Light trap, 55
capacity to liv~"without water, 112 Lindane, 98
capacity to withstand heat, III Lycanthropa, 109
existence under particular conditions Lycopodium, 61
of humidity, 112 M
Insects useful to man, 18
Insects enemies of weeds, 27 Macrodontia cervicornis, 9
Insects yielding products of commer- Macrosiphum pis;' 24
cial value, -27 Maellerius versicolor, 120
Introduction of exotic parasites and Maize, 205
predators, 38 Mala thion, 203
Modem partnership between man and Mantichora, 109, 119, 121
medium-sized honeybee, 28 Mealy bug, 240
Parasites and predators, 22, 36 Medicago sativa~ 88
Pollinators, 19, 35 Medurasia obscurella, 91
Utilization of indigenous parasites Megachile, 20
and predators. 38 Megasoma elephas, 9
Integrated pest control, 239 Melanagromyza phaseo/i, 90
Cropwise integrated schedule, 244 Melanophus, 180
Definition of pest control, 239 Melasina, 118
Feasibility of integrated control, 249 Melophorus, 120
Genetic-cum.3tatistical basis for inte- Mercurials, 203
grated control, 246 Messor, 120
Larg-scale organized campaign, 257 Microbembix, 111
Pestwise integrated schedule, 243 Microbracon Ie/roy;, 128, 131
Philosophy of integrated control, 245 Microcerus, 121
Principle of integrated control, 234 Microctonus indicus, 25
Recent needs for integrated control, Mirperus sp., 90
245 Moganeura, 9
Use of stomach poison, 239 Monocrotophos, 98
Strategy of integrated control, 24 Mylocerus maculosus, 180
Introduction of lethal genes, 198 Myrmecocystus, 119, 120
J N
Jute, 205 Narcotized collections, 55
Juvenile hormones, 198 Nephantis serinopa, 24, 25
Nephotettix apicalis, 240
K bipunctatus, 240
Kusum, 32 Net sweeping, 54
Neuroctenus, 109
L
Nilaparvata lugens, 82, 140
Lac insect, 32 Nomia, 20
Laccifer lacca, 32 Nymphuia depunctalis, 241
Laps, 116
LD60,186
o
Leather jacket, 96 Oactylis giomerata, 89
INDEX 265
Odonata,9 Philaenus spumaris, 89
Oecophylla smaragdina, 36 Phleum pratense, 97
Ophichthy boro, 5 Phyllophaga cribrosa, 109
OpUlitia dilleni, 27 forcala, 109
elatior, 27 hirticu/a, 108
tipp., 27 lanceolata, 109, 114
vulgaris, 27 submuclda, 114
Origin of insect pests, 40 Phyllotreta cruciferae, 25
balance in nature, 42 Phytomyza atricornis, 91
population density, 41 Phytophaga destructor, 81. 1i 2
with origin of agriculture, 48 pink borer, 241
Orthomorpha, 256 Place of entomology,
Orthoptera, 9 in relation to man, 1
Oryza sativa, 82 P/usfa peponis, 25
Oscinella frit, 91 Pogonomyrmex, 120
Ostrinia nubilis, 96 poppy root weevil, 90
Oxya bidentata, 242 Population of individual species, 9
multidelltata, 242 Preno/epis, 120
velox, 242 Principles of-insect control, 177
Oxyopomyrmex, 120 Analysis of external and internal resis-
tance, 186
p
Biological control, 179
Pachydiplosis oryzae, 82, 241 Bioassay of rela ti ve toxicity of insecti-
paddy,~03 cides, 186
paddy eel worm, 241 Chemical control, 179
paddy leaf roller, 241 Cultural control, 178
paddy skipper, 241 Coopera tion, 117
paddy stem-borer, 240 Development of resistance an index
Papaver somniferum, 90 of the intensiveness of control opera-
Parathion, 203 tion, 194
Parnara mathias, 241 Economics, 177
Pest control, 51 Effect of humiditv on insect resistance
l~ ,
Pest of,
maize and millets, 249 Effect of temperature on insect resis-
paddy, 252 tance, 193
pulse crops, 254 Effect of particle size on toxicity of
sugarcane, 250 suspension, 189
Pesticide hazards. 198 Factors associated with higher resis-
Basic points to be kept in view, 198 tance, 190
Nature and extent of hazards in pest Factors affecting stomach poison, 193
control, 203 Hazards to human health and upset-
Review of precautions recommended ting the balance in nature, 195
and additional suggestions, 206 Insecticidal formulations and appli-
Review of recommendations of various ances 195
committees in U. K. and U. S. A., Lipoid solubility and insect resistance,
201 189
Types of pesticide hazards, 199 Mechanical control, 179
Phadka grasshopper, 113 Natural control, 178
Pharnacea serratipes, 9 Pre'Jention, 177
Phaseolus vulgaris, 90 Terminology for insect resistance, 183
Pheidole, 120 Prodenia litura, 25
266 AGRICULTURAL ENTOMOLOGy AND PEST CONTROL
Prodigiousness of destruction, 43 Schislocerca, 128

Prodigiousness of reproduction, 42 gregaria, 128, 129, 131, 132, ISO, 190
Pronuiba yuccasella, 21 Schizaphis graminum, 88
Prospaite/la, 249 Schleichera oleosa, 32
Psalis securis, 241 Schoenobius incertulas, 240
Psammodes, 109, 122 Scirpophaga innotata, 241
Pyrilla, 60, In, 204, 250, 251, 252 Screen trap. 54
Scotinophara lurida, 82, 140
R
sermlooper, 241
Rattus, Sesamia calamitis, 86
no;vegicus, 89 inference, 241
rattus, 89 sight counting, 56
Revolution in pest control, 212 silk worm, 33
Extraordinary high losses due to insect Silolroga cerealella, 22
pests, 212 Smiliotus, 115, 122
Failure of national demonstrations, Solanum sp., 97
218 tuberosum, 84
Imbalance between development n~eds sorghum midge, 88
and research, 222 Sphex, 121
Increasing frequency of pest epidemics, Spodoptera mauritia, 240
215 Sporotrichum globuliferum, 112
Law of limiting factor, 218 spotted grasshopper, 241
Negative correlation between success Steira, 109
in production efforts and intensity Stenobracon deesae, 24
of pest, 212 Stenocara, 109
Pest control research needs not adequ- Stenocaxnlls, 90
ately realized, 221 Sterile male teChnique, 195
Pest control revolution preceded agri- Striped rice borer, 241
cultural revolution, 215 Suction trap, 56
Pest control basically different from Sudden trapping, 54
input, 218 sugarcane, 204
Reason for worsening pest situation, surface grasshopper, 241
215 Sylepta derogata. 25
Second revolution needed, 215 Systropus, 121
Wastage of costly input, 219 swarming catterpillar, 240
rice borer, 241
rice bugs, 240 T
rice caseworm, 241_ Tanymecus dilatocollis, 87
rice grasshopper, 241 Tarucus, 117
rice hispa, 241 tea, 205
rice root aphid, 240 termites, 81, 87
rice weevil, 241 Tetraneura hirsllta, 240
Ripersia oryzae, 240 Tettigella spectra, 240
root wqrm, 86 Thrips imaginus, 56
Ruta tuberculata, 119 Timothy fiies, 91
s Trachynotus, 109
Tribolium confusum,180, 186, 187, 188,
Saccharum officinarum, 88 193
sp., 89 Trichogramma. 22, 23, 26, 38
Scant ius, 109 evenescence minlltllm, 243
Sceliphron. \21 Trichospilus, 26
INDEX 267
pupivora, 24 w
Trioxys indicus, 23
Water trap, 55
Tripu/a sp., 96
White jassids, 240
Triticum aestivum, 81
spp., 81 x
Trogoderma grallarium, 189, 190
Xy/ocopus, 20
Tryporyza incertuias, 87
Tychius jlavus, 89 Z
Zea mays, 84, 86, 87, 96
v Zizyphus, 117
Veromessor. 120 mauritialla, 32
Veterinary, 205 I Zophosis, 109
Vigna unguicuia,a, 89, 90 punctata, 111

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AGRICULTURAL

INDIAN COUNCIL OF AGRICULTURAL RESEARCH

Chief Editor P. L. JAISWAL

Chief Production Officer KRISHAN KUMAR

Chief Artist M. K. BARDHAN

© 1983, by the Indian Council of Agricultural Research, New Delhi

Printed in India by Singhai Printing Press, JabaJpur -482002 and published by

he was to have read at a conference of plant breeders in the last week

About the Author iv

4. Origin of Insect Pests 40

8. Effect of Temperature on the Dynamics of

9. Principles of Insect Control 177

11. Revolution of Pest Control 212

12. Importance of Protection Research in Tropics 223

13. Integrated Pest Control 239

PLACE OF ENTOMOLOGY IN RELATION TO MAN

Fig. 1.1 A typical representative of class Insecta.

term Schatpada, also meaning 6-legged, was there in Amarkosh as eady

about the utility, futility, or even harmfulness of the knowledge thus

Entomology as an Applied Science

destroys mosquito larvae and has been recognized as definitely useful in

Colossal Damage by Insects

and livestock health is incalculable, as also it~ repercussion on the

itself was in its infancy. Similarly. epidemiological studies on insects of

INSECTS came into existence, according to various estimates, 250-500

• Metcalfe, Z. P. 1940. Ent. News 51 : 219-222.

1. The largest insects

Population of Individual Species

other group of animals or in plants; the significance of these has yet to

with 3 linear dimensions. Hence, when the size of an animal in-

may be on the antennae, on mouthparts, or even on tarsi or cerci. Such

requirements of food and habitat that competition between the parent

High Fecundity and Controlled ReproductioD

borers, and top-borers of sugarcane.

Protective Adaptation and Device

lNSECfS USEFUL TO MAN

ALTHOUGH a large number of insects are injurious to crops and are to

has to be carried by some agency to the stigma of female flowers. What is

bees were introduced from outside. The total value of insect-pollinated

bing the stamen, scraping off a quantity of pollen grain, carrying it to

Parasites and Predators

866 individuals have been recorded to emerge frbm 1 host pupa of

Coleoptera. The adults are medium to fairly large-sized insects, round or

Insect Enemies of Weeds

Insects Yielding Products of Commercial Value

numerous -species of flowering plants and in this process bring about

the workers it differs in being bigger in size and having an elongated

colony till it gets exhausted and dies.

The Lac Insect

is highly density-dependent and is reduced with reduction in host popula-

Parasites and Predators

population goes down so that the efficiency of the parasite population

are various precautions possible in the caSe of chemical control. Such

ORIGIN OF INSECT PESTS

THE earliest fossil remains of insects are traceable up to the Devonian

known rate. Hence, by determining the proportion of radioactive carbon

evolutionary march changed their status.

Despite all the above considerations in deciding whether a particular

Let us also see some examples of reproductive potential in other

overlooked. Some experimenters could rear only 8 % of the E. fabia eggs,

tion in the field of electricity: C =:!_ , in which C is the electric

Fig. 4. L A diagrammatic sketch illustrating the decrease in the number of

resistance factors in both cases are complex combinations of a number