REPRODUCTIVE BIOLOGY Of A LEHDOPTEROUS PEST

DUSERTATION SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS

FOR THE AWARD OF THE DEGREE OF

Muittt of $I)tlo!gop!f IN

AGRICULTURE (ENTOMOLOGY)

BY

RUBINA HAFEEZ

INSTITUTE OF AGRICULTURE ALIGARH MUSLIM UNIVERSITY

ALIGARH (INDIA) 1993

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D E P A R T M E N T O F Z O O L O G Y AUGARH MUSLIM UNIVEnSITY

ALIGARH.U.P. INDIA 202002

Phonos ^Universily: 285 \puhlir. ; 260 AC

Sections : 1 ENTOMOLOGY 2 PARASITOLOGY 3 ICHTHYOLOGY & FISHERIES 4 AGRICULTURAL HEMATOLOGY 5 GENETICS

Ref. No .„

Oafo....^.....9l**?.*?:®F/„.12??.

This is to certify that the dissertation for

M.Phil. degree in Agriculture . of the Aligarh Muslim

University/ Aligarh has been completed by Ms. Rubina

Hafeez/ under my supervision. It is original in

nature and I have permitted the candidate to submit

it in partial reuirements for the award of the

degree.

^

lAYON MUF

,.l ( HOMAYON MURAD ) Supervisor

• ~ ^

CONTENTS

PAGE NO.

Acknowledgement

I INTRODUCTION 1

II MATERIAL AND METHODS 5

III REVIEW OF LITERATURE 9

IV RESULTS (Biology)

i) £gg 13

ii) Larval instars t-vi 14

iii) Pupa . 18

iv) Adult 19

V. DISCUSSION 24

VI. RESULTS (Food Consumption and Utilization).... 30

i) Consumption ind^x 32

ii) Growth rate 32

iii) Efficiency of conversion of ingessted food 33

iv) Efficiency of conversion of digested food 33

v) Approximate digestibility 33

VII. DISCUSSION 35

VIII.REFERENCES 38

ACKNOWLEDGEMENT

It gives me immense pleasure and honour to express

my deep sense of gratitude to Prof. Humayun Murad who has

guided me with scholarly patience and aplomb. Prof.

Murad's wide experience and knowledge in Entomology and

his willingness to share it with me was of great help in

completing the present work.

I am greatly indebted to Prof. M.Sharoim Jairajpuri,

Director/ Institute of Agriculture and Prof. Man Mohan

Agarwal/ Chairman/ Department of Zoology for providing

necessary facilities. Special thanks are extended to Mrs.

Naushaba Murad/ Reader/ A.M.U. Women's College/ for being

the constant source of encouragement and moral support

during the course of this work.

I am highly thankful to my senior Dr. Archana

Bansal and colleagues Ms. ' Robina Naseer and Mr. Arshad

All Siddiqi for their critical suggestions and valuable

contributions. It is my pleasure to thank my colleague

Mr. Mohd. Amir who sacrificed his valuable time and

without his constant help the present work would have not

been completed.

I remember with gratitude the help rendered by

Mr. Khwaja Jamal in the field collection/ culture

maintenance and the photography of the experimental

insect.

Most cordial thanks are due to my friends Ms.

Subuhi Khan and Mrs. Dipender Vohra for being generous

with support and encouragement whenever I needed.

On a more personal note I wish to thank my eldest

brother Dr. Anwar Ghani/ Senior Scientist, Ministry of

Agriculture & Fisheries/ New Zealand, whose encouraging

comments and suggestion provided me* with the confidence

and capability to bring this work to completion.

I would like to express my sincere gratitude to

my family particularly to my parents, whose teachings of

hard work and discipline have inspired me throughout the

arena of my study.

Finally I would like to express my thanks to

Mr. Kafeel A. Khan and Mr. Abid All for taking pains in

meticulously typing the manuscript.

Above all I am thankful to Allah almighty for

giving me the opportunity and strength to complete this

work.

( RUBINA HAFEEZ )

INTRODUCTION

India is predominantly an Agricultural land with nearly

70% of its working population engaged in Agriculture. It

accounts for about 40% of the total National income and is

therefore the mainstay of Indian economy. Therefore, the speedy

development of agriculture is vital to the progress of our

country. For securing maximum crop production the best use of

the knowledge of the latest tools and techniques has to be

done. Our country having varied climatic conditions is in

unique position to grow almost every possible crop and occupies

an outstanding position in the world with respect to several

agricultural products.

Insects are the chief rivals of human beings and their

unceasing struggle is continued because of the fact that both

man and certain insects constantly require the same thing at

the same time. In an attempt to secure food/ the insects

inflicit considerable damage to every part of the plant causing

an indirect effect on the economy of the country. Insects

affecting the crops are causing two fold damage to our economy,

one by affecting the quality and reducing the quantity of the

product and the other by requiring the investment of money and

involvement of mechanical and manual labour for their proper

control measures. The insects have an intimate relationship

with the plants. The more luxuriant and varied the flora, the

more abundant and diversified its accompanying insect fauna.

:2:

The seriousness of the attack is decided by the nature

of the injuries inflicted/ the susceptibility of the plant to

the attack, the feeding efficiency and the host plant

relationship. The insects cause damage at various levels,

starting from the field upto the storage. The losses caused by

insect pests are 20-35% out of which 12-15% is contributed by

the stored grain pests alone (through personal communication

from the Plant Protection Department, Ministry of Agriculture).

There is a significant lowering in the losses caused by insects

in recent years because of the advancement in the scientific

knowledge and awareness of the farmers.

Among the diverse forms of insects infesting agricul­

tural crops, those belonging to Lepidoptera are very important.

Though no accurate estimate of the annual losses caused by

Lepidopteran pests in value of the product is done in our

country, but it has to be admitted that the extent of damage

may occasionally be considerable. Very little information is

available as to the exact identity of the pest species and

their relative importance.

For an effective and economical application of

different methods of control so far known, it is essential to

have proper idea of the insect pest (identification and

biology) one has to deal with. The knowledge of the life

history of an insect will be very helpful in adopting such

measures against any pest. The biology or the life cycle of an

insect means the record of all that the insect habitually does

:3:

and all the changes in form and habits that it undergoes from

the beginning of its life until its death/ including the

situations where each life stage and every season is spent and

the length of the time occupied by each stage.

The hairy caterpillars or the "Wooly Bears" are well

known foliage feeders. Many of them feed exposed on the leaves

while others feed within the rolled or folded leaves. The Bihar

hairy caterpillar, Diacrisia (= Spilosoma) obliqua Walker

belongs to the family Arctiidae- The members of this family are

found devouring the foliage of almost any low growing crop, but

a few of them are very specific. D^. obliqua is a polyphagous

pest, widely distributed throughout the country, found very

commonly in Uttar Pradesh, Madhya Pradesh, Bihar and Punjab

(Atwal, 1986). It causes considerable damage to pulses,

cotton, vegetables, sorghum, maize, ragi, oil seeds, small

millets, sugarcane, pady, wheat, guinae grass, jute, sunhemp,

beet-root, potato, and sweet potato (Mathur, 1962; Pant, 1964).

Besides, they also attack other agricultural and fibre crop as

well as medicinal plants (Pant, 1964).

For the assessment of the damage caused, insect plant

relationship, ecological success and the control measures

required for a particular pest, it is essential to have a basic

knowledge of identification abundance, distribution, food

range, damaging stage, damaging potential, mode of damage, life

history and the nutritional ecology-

:4:

Most of the work done in the area of nutritional

ecology was based on the food preference, preparation of

artificial diets or on the total utilization profile. But for

the better understanding of the insect plant relationship and

the success of the pest in the ecosystem, it becomes all the

more important to work in detail the food consumption/

utilization, digestibility and conversion of food into body

tissues. Food consumption, in addition to accounting for the

quantitative loss brought about by the pest species also serves

as an indirect measurement of the relative susceptibility of

food plants.

Keeping in mind immense economic importance, pest

status and its ecological success, it has been desired to study

the biology and consumption, assimilation and utilizaton of

castor leaves (Riccinus communis) by the larvae of _D. obliqua.

The present work is divided in two parts, the first one deals

with the biology and the second accounts for the quantitative

food consumption and utilization.

:5;

MATERIALS AND METHODS

The culture was started with the eggs which were

collected from the field during the spring season. The rearing

was done in glass jars measuring 12" x 8". The larvae of

Diacrisia obliqua Wlk. were reared on castor, (Riccinus communis)

leaves at 27±2°C temperature, 65±5 relative humidity and 12:12

hrs. photoperiod. The food was supplied twice a day and the

hygienic conditions were maintained by cleaning the jars daily.

To avoid overcrowding the number of larvae per jar was

restricted to 100-200, I-II instar and 30-50, III-VI instar

larvae. The adults were fed on 10% glucose solution soaked in

cotton changed daily to avoid fungal growth. Folded paper

strips were placed in the jars containing the adults for

oviposition.

The biological cycle of the pest was started from the

freshly hatched larvae (in three replicates of five each) and

was completed on the same stage by passing through a series of

larval instars, pupa, adult and the egg. The individual

specimens were isolated from the mass culture for the detailed

morphological studies.

The quantitative assessment of food consumption

digestion and utilization was done by using Waldbauer (1968)

procedure with slight modification. The food consumed, excreta

voided and the weight gained by individual larvae from III to

VI instars were determined on dry weight basis-

6:

The method involves the isolation of freshly moulted

larvae from the mass culture. These were daily supplied with

fresh castor leaves (weighed wet) for three consecutive days.

The left over wet leaves and the excreta voided were weighed

the following day. The difference between the fresh leaves and

the left over leaves gives the net weight of the leaf consumed.

For finding out the dry weight of the food consumed/ equivalent

amount of fresh leaves were placed in an oven for one hour at

100°C and then at 60°C until the weight became constant. The

excreta voided was also dried by using the same method. The

difference between the amount of food consumed and the feces

produced is the weight of the food digested.

The fresh weight of the larvae was determined at the

beginning of each experiment. The dry weight of each larva was

estimated by using the mean percentage dry matter of an aliquot

of like larvae which were killed by chloroform and then dried

in an oven (at 100°C for 1 hr and at 60°C till the weight

became constant). The difference between the final dry weight

of the larva when sacrificed after the moult and the estimated

initial dry weight was the net weight increased. The amount of

food consumed or digested in relation to the increase in body

weight is a measure of the efficiency of utilization.

The experiment was set in three replicates of five each

for three consecutive days per larval instar at 27±2°C

temperature, 65±5 RH and 12:12 hrs- photoperiod.

In the expression of the result following indices were

used :

The approximate digestibility (A.D.) is calculated as :

A.D. = ^ ~ ^ X 100 F

The growth rate (G.R.) is expressed as :

G.R. T A

The consumption index (CI.) is calculated as

C.I T A

The efficiency of conversion of ingested food to body matter

(E.C.I.) is a measure of the overall ability of the organism to

grow on a given food. It is calculated as :

E.C.I. = p ^ x .100 = f X 100

The efficiency of conversion of digested food to body matter

(E.C.D.) is calculated as :

E.C.D. = - F ^ - X 100

Where F represents the dry weight of the food ingested

per larva per three days, E is the dry weight of the excreta

voided per larva per three days, G expresses the dry weight

gained per larva after feeding/ T is the time of feeding and A

is the mean dry weight of the initial, final and intermediate

weights divided by the number of weighings. Since the larvae on

8:

various fresh leaves ate and grew at different rates over

variable periods, therefore it became necessary to express all

measurements in a form which made possible the comparison of

intake and efficiency of utilization.

REVIEW OF LITERATURE

Diacrisia obliqua Wlk. is an important pest of

various crops in India. The larvae of this moth are

polyphagous and voracious feeders of the foliage. It has been

reported from all over the country to cause considerable

damage to pulse/ cotton/ vegetables/ sorghum/ maize/ ragi/

oil seeds/ small millets/ sugarcane/ paddy/ wheat/ guinae

grass / jute/ sunhemp/ beat root/ potatoes and sweet potatoes

(Mathur, 1962 and Pant, 1964).

It has been reported as the most destructive pest of

soybean (Gangrade/ 1976/ 77; Bhardwaj & Bhalla/ 1977; Ram &

Bhattacharya/ 1978; Khaleque, 1983; Haq et al./ 1984/ 85

and Ali/ 1988). It causes significant damage to oil seed

crops like groundnut/ castor/ mustard and sunflower. It has

been reported on groundnut by Srivastava et al. (1965)/ Sethi

et al. (1979)/ Pachori et al. (1980)/ Islam et al. (1983) and

Vora et al. (1985). It has been reported on mustard by

Bakhetia & Brar (1982) and Bakhetia (1986); on castor by

Srivastava et al. (1972) and Singh & Gangrade (1974) and on

sunflower by Rohilla et al. (1981) and Srivastava & Pandey

(1987).

It has been found damaging jute by Sharif (1962) and

Tripathi (1967)/ on maize by Verma et al. (1977). Among

pulses it has been found causing severe damage to moth bean

(Phaseolus acontifolius Jacq.) (Puttaswami et al. / 1977),

:10:

french bean (jP. vulgaris L.) (Puttaswami & Reddy/ 1981), as

seasonal pest on green gram (£. aurenis Roxb.) (Sinha et al. /

1985) and on mungbean (Lai, 1985). Its incidence has also

been reported on blackgram (Vigna mungo L.) (Dhuri et al.,

1984) on cowpea {V_. sinensis Savi) (Yadav & Yadav, 198 3), on

chicken pea (Cicer arietinum L.) (Deka et al., 1987).

It has been found attacking some ornamental plants

and vegetables (Gargav & Katiyar, 1971 and Sachan, 1981). It

has been reported on Chinese potato (Coleus parviflorus) by

Palaniswami & Pillai (1983), on raddish by Butani & Juneja

(1984). Among ornamental plants, the flowers of Gladiolus and

leaves of globe artichoke (Cynara scalymus) are very prone to

its attack (Krishnah et al., 1975). Its incidence has also

been reported on few medicinal plants by Mathur (1962).

An other species S . lubricipeda has been reported on

apple and pear foliage by Sengalevich (1960) and ^. Virginia

was found very common on weeds (Rizzo, 1978, 79). The larvae

of £. obliqua have been deployed to control the cosmopolitan

weed Lantana (Benson & Chatterjee, 1940).

D. obliqua being a polyphagous pest consumes a

variety of food. These foods have variable effect on their

growth, development, nutrition and reproduction (Babu et al.,

1978; Gupta et al. , 1979; Prasad & Prem Chand, 1980 (a & b) ;

Deshmukh et al. , 1982; Gupta, 1982; Srivastava & Pandey,

1983; Goel et al., 1986 and Kabir & Miah, 1987).

11;

Morphological studies of the adults have been done by

Ahmad & Ahmad (1976) and the larval morphology was studied by

Goel & Kumar (1983)/ biology and larval morphology of S .

Virginia (F.) was studied by Rizzo (1978/ 79).

The bioecology of D. obliqua has already been done on

a number of host plants. Temperature has been found as one of

the most important ecological factor which affects the growth

and development of the insects (Singh & Gangrade/ 1974; Bhat

& Bhattacharya/ 1978/ and Chaudhary & Bhattacharya, 1985).

The other ecological factors like photoperiod and light

intensity do have significant effect on the premating

behaviour or the calling rhythm of the arctiid moths (Webster

& Conner, 1986). Details of mating and oviposition of D.

obliqua have been studied by Islam & Alam (1980), Husain

(1982) and Siddiqi (1985/ 88). Observations on the biology,

larval and pupal durations adult emergence/ longevity and

the percentage mortality of adults have been worked out by

Sinha & Gangrade (1974/ 77)/ Babu et al. (1978)/ Deshmukh

et al. (1982)/ Srivastava & Pandey (1983) and Chaudhary &

Bhattacharya (1986).

The efficiencies of consumption and utilization of

ingested and digested food has been worked out in a few

Orthopteroid and Lepidopterous insects (Waldbauer 19(54/ 68).

Among Orthopteroids / the food consumption/ assimilation and

tissue growth have been studied by Chlondy (1967)/ Bennakkars

12

et al. (1970), Mordue & Hill (1970), Mehrotra et al . (1970),

Khalsa (1973), Gupta & Vats (1980), Simpson (1982) and Vir &

Jindal (1983). Among L^idopteran insects the ecological

efficiencies, food consumption, utilization, growth rate and the

efficiencies of conversion of ingested and digested food has

been worked out by Mukerjee & LeRoux (1969), Kogen & Cope

(1969), Mathavan & Bhaskaran (1975), Bhat & Bhattacharya (1978),

Mackey (1978), Haukioja et al. (1978), Ram & Bhattacharya

(1978), Chand (1979), Babu et al. (1979), Prasad & Chand (1980),

Valentine & Talerico (1980), Cohen & Patana (1985), Crocomoard &

Parra (1985), Slansky Jr. (1985), Warrington (1985) and Sharma &

Tara (1988) .

REPRODUCTIVE BIOLOGY

13:

RESULTS

The Bihar hairy caterpillar/ Diacrisia obliqua Wlk.

is commonly found abundant in and around Aligarh. These are

capable of damaging a variety of crops like oil seeds,

vegetables, fibre crops, fruits, ornamental plants and even

the weeds. Because of its significant economic importance,

this moth has been selected for the present study. The moths

are active during night hours, get readily attracted to light

and are strong and steady fliers for short distances.

EGG

The eggs are l a i d in batches e i ther on the paper

s t r i p s , sides of the g la s s j a r s or on the muslin cover in

c a p t i v i t y . The freshly l a i d egg i s firmly glued to the

surface with the st icky s ec re t i ons of the accessory gland.

The newly la id egg i s g l i s t e n i n g , small and spher ica l in

shape. I t measures 0.75 mm in diameter and i s l i g h t green in

colour but undergoes a s e r i e s of changes before ec los ion . The

colour changes from green t o p a l e , redish brown, brown and

f i n a l l y black .(Fig.l&2). The outer covering of the egg becomes hard and

t rans lucent a few hours before hatching and the la rva inside

can eas i ly be recognised from the ex t e r io r . The incubation

period var ies between 173.15:—215.15 hrs . with an average of

-198.33±23.24 hrs. The percentage of eggs successful ly hatched

(eclosion) i s 98.8%. The eggs usual ly hatch in masses .(I^ble - 1 ,

Fig. 13).

: 14;

FIRST INSTAR LARVA

The first instar larva is a minute creature with a

symmetrical body. It is pale yellow in colour with brownish

head and gives a greenish reflection after feeding on the

leaves. They bear five pairs of prolegs from third to sixth

and the last abdominal segment. The body remains covered with

setae and hairs, pre-spiracular wart of the prothorax with

three setae. The setae and hairs are not visible with .the

nacked eyes (Plate-II/ Fig. 3).

The full grown first instar larva measures 3.8 mm in

length and 18.15 mg - in dry weight. The average first instar

larval duration ranges 84.82±20.22 hrs. (Table-1, Fig. 13).

The newly hatched larva shows wriggling movement and

wanders around the container in search of food. It feeds on

the mesophillic tissues of the leaves, leaving the vascular

ones and thus act as skeletonizers. If there is scarcity of

food, these larvae consume the entire leaf and become the

defoliators. When disturbed they hang themselves with a

silken thread which also help them in the dispersal. The

larvae are gregarious in habit.

SECOND INSTAR LARVA

The second instar larva can easily be distinguished

from the first instar by having a characteristic cylindrical

body. The newly moulted ones remain sluggish and are light

15:

yellow in colour but the active feeding stage is bright

yellow with slight orangish tinge. The number and position of

prolegs remains the same/ the chrochets become prominent. The

location and the number of setae oh the prespiracular wart

of the prothorax are similar to the first instar larvae. The

hairs become prominent because of acquiring black colour

(Plate-II, Fig. 4) .

The actively feeding second instar larva measures 5.8

mm in length and 29.64 mg in dry weight. This stage lasts for

91.54±20.37 hrs(Table - 1/ Fig. 13).

The feeding pattern is also very much similar to the

first instar i.e. only consumes the mesophyllic tissues but

the late second instar also consumes the secondary and

tertiary veins (cortical and vascular tissue). When disturbed

it drops suspended with the silken thread which is secreted

by its mandibular glands.

THIRD INSTAR LARVA

The third instar larva is very slender/ orange in

colour with black spots on all the thoracic and the last two

abdominal segments. The number and position of the prolegs,

and the setae on the prespiracular wart is similar to the

first and the second instar larvae. Warts of every segmen;

become prominent because of the thickening of the cuticle in

that area. The hairs and the setae can be distinguished into

: 16:

two t y p e s i . e . smoo th and t h e s p i n o s e ( P l a t e - I I I / F i g . 5 ) .

The a c t i v e t h i r d i n s t a r l a r v a m e a s u r e s 1 1 . 4 mm in

l e n g t h and 3 8 . 5 2 mg i n dry w e i g h t . T h i s i n s t a r l a s t s f o r

1 2 7 . 0 0 ± 1 2 . 2 9 h r s { T a b l e - 1 , F i g . 1 3 ) .

These a r e g r e g a r i o u s and v o r a c i o u s f e e d e r s of t h e

f o l i a g e c a u s i n g s e v e r e damage t o t h e c r o p s . They consume

e n t i r e l e a f l e a v i n g o n l y t h e m i d r i b . When d i s t u r b e d they t r y

t o d r o p t h e m s e l v e s suspended w i t h t h e s i l k e n t h r e a d which

soon b r e a k s up b e c a u s e of a s i g n i f i c a n t i n c r e a s e i n t h e i r

body w e i g h t .

FOURTH INSTAR LARVA

The body i s c y l i n d r i c a l / a p p e a r s a s a s t r o n g / s t o u t

and a ha rdy c r e a t u r e which a r e d e n s e l y h a i r y . The c o l o u r i s

f u r t h e r d a r k e n e d t o w a r d s o r a n g e w i t h v e r y p rominen t b l a c k

s p o t s on t h e t h o r a c i c and t h e l a s t two abdomina l segments

( P l a t e - I l l , F i g . 6 ) .

The a c t i v e l y f e e d i n g l a r v a l s t a g e m e a s u r e s 21.8 mm in

l e n g t h and 5 0 . 0 6 mg i n d ry w e i g h t . T h i s l a r v a l i n s t a r l a s t s

f o r 1 2 8 . 3 6 ± 9 . 8 1 h r s ( T a b l e - 1, F i g . 1 3 ) .

The f e e d i n g b e h a v i o u r i s v e r y much s i m i l a r t o t h e

t h i r d i n s t a r . When d i s t u r b e d they t r y t o e s c a p e from che a r e a

by f a s t c r a w l i n g movements .

: 1 7 :

FIFTH INSTAR LARVA

The f i f t h i n s t a r l a r v a i s c h a r a c t e r i z e d by t h e tough

and compact body h a v i n g more d e n s e h a i r s a s compared t o t h e

f o u r t h i n s t a r . The h a i r s a p p e a r i n g r o u p s on t h e w a r t s

(become t u f t e d ) and a r e v e r y l o n g . S i n g l e h a i r measures 6-8

mm i n l e n g t h . The c o l o u r i s b r i g h t o r a n g e w i t h s i m i l a r b l a c k

s p o t s a s on t h e e a r l i e r i n s t a r s b u t t h e i r i s a p r o m i n e n t

b l a c k band i n t h e i n t e r s e g m e n t a l a r e a ( P l a t e - IV/ F i g . 7 ) .

The f u l l g rown, a c t i v e l y f e e d i n g f i f t h i n s t a r l a r v a

m e a s u r e s 2 8 . 6 mm i n l e n g t h and 7 7 . 9 3 mg i n d r y w e i g h t . Th is

s t a g e l a s t s f o r 1 3 3 . 5 8 ± 4 . 7 2 h r s ( T a b l e - 1 , F i g . 1 3 ) .

The f e e d i n g b e h a v i o u r and t h e p a t t e r n i s a lmos t t h e

same a s i n t h e t h i r d and t h e f o u r t h i n s t a r s but t h e amount

and t h e r a t e of t h e food consumed i s s i g n i f i c a n t l y i n c r e a s e d .

SIXTH INSTAR LARVA

The s i x t h i n s t a r l a r v a i s d i s t i n g u i s h e d by a s l e n d e r ,

s t r o n g and s t o u t b o d y , r e s e m b l i n g v e r y much w i t h t h e f i f t h

i n s t a r l a r v a i n t h e c o l o u r , s p o t s and t h e p o s i t i o n of h a i r s .

Because of t h e h a i r s be ing t u f t e d t h e r e i s a c l e a r m i d - d o r s a l

s t r e a k which i s s l i g h t l y p a l e i n c o l o u r v e n t r a l l y . The l a r v a

i s p a l e y e l l o w w i t h p r o m i n e n t i n t e r s e g m e n t a l a r e a s ( P l a t e -

IV, F i g . 8 ) .

The l a s t i n s t a r l a r v a m e a s u r e s 3 0 . 7 5 mm in l e n g t h and

1 1 7 . 6 1 mg i n d r y w e i g h t . The e n t i r e l a r v a l p e r i o d i n c l u d i n g

18:

the pre-pupal period under controlled conditions varies

between 212.44±5.88 hrs. Duration of the last instar alone is

138.85±2.54 hrs (Table - 1, Fig. 13). The feeding behaviour

is similar to the earlier instars. It mostly feeds during

night hours or in dim light and are voracious feeders and

capable of devoring every soft part of the plant.

When the last instar larva attains the maximum size

it ceases feeding and comes to rest/ and after a break of 4-6

min. starts moving all round the container in search of

suitable site for pupation.

PUPA

After selecting the suitable site the larva becomes

confined and the structural modifications start appearing.

The size of the last larval instar is almost reduced to halt,

the intersegmental grooves are deepened giving it an annular

appearance. The hairs and streakes have almost faded out/

the abdominal prolegs are reduced and the abdomen is

tapering/ this stage is known as pre-pupa.

The pupation generally takes place in the debris of

the dead and decaying leaves/ it has also been found

pupating on the ground/ in the soil (in field conditions), on

the muslin cover or even on the sides of the glass jars in

captivity. The pupation occurs within the silken cocoon

interwoven with the shed off hairs of the last instar larva.

The cavity inside is large enough to accomodate the pupa. The

: 19:

cocoon is very loosely woven and therefore in most of the

cases the pupa comes out of it. There are pulsatory

movements in the pre-pupa within the last larval moult. The

posterior extremity of the pupa within the pre-pupal cast

rotates, thus helping in the removal of the cast.

The newly formed pupa is tapering in structure and is

pale yellow in colour but turns to leathery brown within 1-2

hrs.The abdominal segments start tanning first followed by

the thoracic and cephalic segments. The female pupae are

comparatively more tanned (Plate - V/ Fig. 10). The full

grown female pupa measures 19.6 ram in length and 3176.6 mg in

weight, the male pupa being smaller in size measures 15.4 mm

in length and 1750.4 mg in weight. There are-.4-8 tubercles at

the posterior extremity of the pupa in both the sexes (Plate-

V, Fig. 9 & 10). The average pupal duration is 309.11±12.4

hrs. (Table - 1, Fig. 13). Hours before moulting the imago

inside the pupa starts exerting pressure on the pupal moult

and as a result prothorax splits mid-dorsally followed by the

splitting of meso- and meta-thorax posteriorly and the vertex

anteriorly.

ADULT

The body of the adult is robust and hairy, the head

small inconspicuous, proboscis well developed and the palpi

are short. They are bright orange in colour with a mid-dorsal

and lateral rows of black spots from the second to the sixth

:20:

abdominal segments with terminal spots becoming more

prominent. Wings are buff orange in colour with minute black

spots on the fore and very prominent spots on the lower

margin of the hind wings (Plate - VI, Fig. 11 & 12).

Both the sexes are almost similar, the males are

characterized by the presence of the pectinnate antenae. They

are comparatively shorter than the females and measure about

15.2 mm in length and 36.8 mm wing expanse. Plate - VI / Fig.

11). The females are much bigger than the males and measure

18.6 mm in length and 45.6 mm wing expanse (Plate - VI, Fig.12).

The newly emerged adults take about 15-30 min. for

the complete expansion of the wings. On their first flight

they discharge a brownish fluid. The entire longevity of the

adults can be divided in three stages:

i) Pre-Oviposition [pre-mating for the male]

ii) Oviposition [Post-mating for the male]

i i i ) Post-Oviposition

Pre-Oviposition :

The pre-oviposition period is the time taken by the

adults to reach the sexual maturity, the mating and the

interval between the cesation of mating and the actual

oviposition. The mating does not occur just after emergence

of the adults. It takes about 12 hrs. time and a 10% glucose

solution meal to attain the sexual maturity. The unfed

females fail to produce viable eggs. When the matured male

21:

recognises the presence of the female, it becomes sexually

excited. The state of excitation was reflected by some of its

behavioural responses. The males start moving actively with

rapid fluttering of antennae and wings. The abdominal tip is

moved up and placed down repeatedly. These activities

continue for 10-35 min.

The females in response show two types of behaviour,

some of them start fluttering their wings with slight

rhythmic movement (contraction and relaxation) in the anal

area while others start moving slowly with their abdominal

tips pressed on the surface of the container. Finally, the

individuals of the opposite sex approximate each other by

bringing their abdominal tips closed and then copulate in end

to end position, remain interlocked for 0.10-1.15 hr. They

can easily be separated by a slight pressure. The mating

generally occurs during night or early morning hours. The

young moths emerged, mate on the subsequent nights. The

observations also reveal that the females mate only once in

their life time whereas the males can mate more than once.

The pre-oviposition period lasts for 19.11±0.42 hrs. (Table - 2,

Fig. 14).

Oviposition

The phenomenon of egg laying does not occur just

after the cesation of mating. It takes about 1-3 hr. for the

females to get ready for oviposition. Before performing the

:22:

actual act of oviposition the females show a series of

behavioural responses. They become very active, start

fluttering their wings, show rhythmic movements in the anal

area and move fast all around the container till they become

exhausted. Then after a pause of 10-30 min. start moving

slowly with slight protrusion of the anal area and subsequent

touching of the substratum to select the suitable

oviposition site. After the selection of the ovisite the

female stays there and shows peristaltic movement in the

abdominal region before egg laying.

For the deposition of the egg the female brings

abdominal tip close to the substratum and descends an egg

out/ then the abdomen is retracted with a slight jerk and the

wings are slightly raised. The eggs get adhered to the

substratum because of a sticky coating of the internal

secretions. After laying an egg the female moves her abdomen

on either side of it and lays the next egg. In this way the

eggs are laid in an organised fashion in several rows with

even spacing between the eggs.

In the case of virgin females the mechanism of

oviposition is same but the pattern is entirely different- In

most of them the eggs are laid scattered or in heaps which

is an abnormal characteristic for this species. Most of the

virgins do not have the urge of oviposition. The average ovi­

position period in the mated females is 117.17±41.5 hrs. (Table-

2, Fig. 14).

23:

Post-Oviposition :

The pos t -ovipos i t ion period i s the durat ion between

the cesation of egg laying and the mortal i ty of the adu l t s .

I t i s very short and va r i e s between 30.59±20.23 hrs . (Table -

2, F ig . 14).

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:24:

DISCUSSION

In majority of moths if suitable conditions are

available the pre-oviposition period is mostly very short as

most of the eggs attain maturity during the later stage of

pupal life. In jD. obliqua this period varies from 19.11±0.42

hrs .Among other moths/ the pre-oviposition period lasted 1-2

days in Thiacidas postica (Mehra & Shah, 1970) and Macalla

monucusalis (Krishnamurthy et al. / 1974). The preoviposition

period varies between 1-3 days in Nephopteryx leucocephalla

(Shah & Mehra/ 1966)/ 54 hr. in Spodoptera mauritia (Murad/

1969)/ Anomis flava (Rao & Patel/ 1973) and Cnaphalocrocis

medinalis (Velusamy & Subramanium, 1974). The pre-oviposition

period of 1-4 and 2-18 days in Heliothis armigera (Singh &

Singh/ 1975)/ Amarasca bigutella bigutella (Singh/ 1978)

respectively, and varies between 2-3 days in £. obliqua

(Siddiqi, 1986).

The moths lay their eggs either singly or in clusters

and a few show both the patterns. In the present study the

females of D. obliqua laid their eggs in batches like that of

Andraca bipunctata (Banerji/ 1971), Chilo zonellus iTrehan &

Butani/ 1949)/ T_. postica (Mehra & Shah/ 1970) and £, obliqua

(Siddiqi, 1986). On the other hand the females of Parnara

mathias (Teotia & Nand, 1966), Hemithea tritonaria (Mehra &

Shah/ 1966) and H_. armigera (Singh & Singh, 1975) laid their

eggs singly. The pattern of both the type was reported in the

:25:

females of Phyllocnistis citrella (Pandey & Pandey, 1964), M.

moncusalis (Krishnamurthy etal., 1974), Polymatus boeticus

(Pandey et al., 1978).

The females of ^. obliqua laid their eggs during

night hours. The nocternal ovipositional behaviour has also

been reported in £. citrella (Pandey & Pandey, 1964), R.

tritonaria (Mehra & Shah, 1966), N . leucocephalla (Shah &

Mehra, 1966), ^. postica (Mehra & Shah, 1970), A . bipunctata

(Banerji, 1971), M. moncusalis (Krishnamurthy et al., 1974),

E. armigera (Singh & Singh, 1975) and D_. obliqua (Siddiqi,

1986) . Where as in C. zonellus/ the eggs were laid only in

evening (Trehan & Butani, 1949) and females of P . boeticus

oviposited in the bright day light (Pandey et al., 1978).

In the present study the oviposition period in £.

obliqua was 117.17±41.58 hrs. Whereas the duration for the

deposition of eggs in a single batch was 20-268 min. but it

corresponds to the number of eggs in them. The oviposition

period in moths is subject to great variation. It is 1-3 days

in C. zonellus (Trehan & Butani, 1949), 1-11 days in H.

tritonaria (Mehra & Shah, 1966), 2-4 days in £. mathias

(Teotia & Nand, 1966) 4-5 days in T . postica (Mehra & Shah,

1970), 2-14 days in A. flava (Rao & Patel, 1973), C.

medinalis (Velusamy & Subramanium, 1974), 2-5 days in fi.

armigera (Singh & Singh, 1975) and £. boeticus (Pandey et

al., 1978), 4-28 days in A. bigutella bigutella (Singh, 1978)

2 6 :

and 10.75±0.41 days i n D. obl iqua ( S i d d i q i , 1986) . In the

p re sen t study t h e i n c u b a t i o n per iod i n £ . obl iqua i s

190.33±23.24 h r s . a g a i n s t t h e e a r l i e r 7.8 days a t 25''C (Singh

& Gangrade, 1974), 2 . 5 - 3 days (Lai & Mukher j i , 1978) . In _H.

armigera t h i s p e r i o d v a r i e s between 2 . 6 - 3 . 6 days with an

average of 3 .26±0.15 days (Singh & Singh, 1975) and 33 h r s .

in ^ . maur i t i a (Murad, 1969) .

The average l a r v a l d u r a t i o n i s s u b j e c t to g rea t

v a r i a t i o n among Lep idop te r ans depending on v a r i o u s parameters

l i k e the environmenta l c o n d i t i o n s , q u a l i t y and q u a n t i t y of

food and the p o p u l a t i o n d e n s i t i e s . In t h e p r e s e n t work the

average l a r v a l d u r a t i o n i s 705.15±69.95 h r s . when the

popula t ion d e n s i t y was 30 i n s t a r s / j a r (measuring 12" x 8" ) ,

f r e sh and excess ive amount of food was supp l i ed twice a day

a t 27±2°C t empera tu r e , 65±5 RH and 12:12 p h o t o p e r i o d . Whereas

i t i s 33.7 days a t 25°C (Singh & Gangrade, 1974) , 19.9 days

on Chenopodium album and 32 .1 days i n B r a s s i c a rugosa

(Rathore & Sachan, 1978) , f l u c t u a t e s between 17-21 days in

v a r i o u s v a r i e t i e s of b l ack gram (Yadav e t a l . , 1978), 20.88

days on sunflower and 24 .5 days on groundnut (Prasad &

Premchand, 1980), 28 .5 days a t 25°C and 24 .5 days a t 30°G

(Deshmukh et a l . , 1982) and 34.34±0.38 days in Hel ianthus

annulus and 40.33±0.54 days on Chrysanthemum f ru tescens

(S r ivas t ava & Pandey, 1987) . .: In £ . m a u r i t i a t he average

d u r a t i o n was 351 h r s . (Murad, 1969), H_. a rmigera 10.8 + 0.73

days (Singh & S ingh , 1975) , Agrot is i p s i l o n 25-35 days

27:

{Nikolov, 1980) and in _H. virescens 15-21 days (Martinez et

al., 1906).

In the present study the pre-pupal and the pupal

period last for 73.22±1.24 hrs. and 308.9±12.74 hrs.

respectively. The pre-pupal period was 12 hrs. in ^. mauritia

(Murad/ 1969)/ 1-2 days in H_. armigera (Singh & Singh/ 1975),

1.5-1.8 days in £. obliqua (Rathore & Sachan, 1978) and 2-3

days in PJ. virescens (Martinez et al./ 1986). The pupal

period in £. obliqua is 10.5-8.3 days (Singh & Singh; 1974),

8.8-9.6 days (Rathore & Sachan, 1978)/ 9.5-11 days in

different varieties of black gram (iTadav et al. / 1978)/ 12

and 29.3 days on groundnut and cotton (Prasad & Prem Chand,

1980), 11.4-12.6 days at 25-30°C (Deshmukh et al., 1982) and

12.43±0,19 days on C. f rutescens and 19.17±0-66 days on C.

indicum (Srivastava & Pandey, 1987). In S_. mauritia 164 hrs.

(Murad/ 1969)/ R. armigera 5-8 days (Singh & Singh, 1975) and

9-17 days ii. virescens (Martinez et al. , 1986).

The adults of £. obliqua emerge at dusk or at night

as reported by Islam & Alam (1979) and Siddiqi (1985). The

premating time in £. obliqua varies between 8-15 hrs. whereas

it has been reported as 18 hrs. by Siddiqi (1985). On the

other hand the moths of A . bipunctata mated immediately

after emergence (Banerji, 1971). Whereas the moths of P_.

citrella (Pandey & Pandey/ 1964), Lamporosema indicata

(Kapoor et al./ 1972) and H . armigera (Singh & Singh, 1975)

matured for mating in about 14-24 hrs,/ one day after

:28:

emergence and 35 min. to 32 hrs. respectively. It takes

1-2.5 hrs. for Hyphantria cunea to be ready for mating

(Aral & Mabuchi, 1979). D. obliqua generally mate at night or

at dusk i.e. their mating time coincides- with the emergence

time. Similar observations have been reported by Islam & Alam

(1979) and Siddiqi (1985). In majority of the moths mating

generally occurs during night such as £. citrella (Pandey &

Pandey/ 1964)/ £. medinalis (Velusamy & Subramanium/ 1964)/

A. bipunctata (Banerji/ 1971) and _L. indicata (Kapoor et al. ,

1972). The Euxoa bilitura always prefers to mate and oviposit

during light (Ripa, 1980).

The activities of excitement were shown by both the

males and the females. Only the recognition of the presence

of the female in the vicinity excites the male and the

behaviour is reflected by its activities. In D_. obliqua the

fast movement and fluttering of wings occurs before mating.

Similar responses have also been reported by Aral & Mabuchi

(1979), Islam & Alam (1979), Kitamura & Koyama (1984) and

Siddiqi (1985). The mating occurs in end to end position in

D_. obliqua. Similar observations have been found in Polytela

gloreosae (Sachan & Srivastava, 1965), S . mauritia (Murad,

1969) and A. bigutella bigutella (Singh, 1978).

Mating in D. obliqua occurs only once in its life

cycle which takes place before first egg laying. Similar

observations were reported by Siddiqi (1985). Single mating

throughout the life was also reported in P. citrella (Pandey

:29

& Pandey, 1964) and E_. armigera (Singh & Singh/ 1975) but the

moths of L_. indicata mated more than once during their life

time (Kapoor et al., 1972).

Plate-I

Fig. 1. Freshly laid egg batch of D. obliqua (X 1.3 Hun).

Fig. 2. Egg batch of D, obliqua just before hatching (X 1.3

mm).

F16.1

^

i FIG.2

Plate-II

Fig. 3. First instar larvc of D. obliqua (X 2.1 mm).

fig. 4.. Second instaj .1;) va of D. obiigua (X 3.4.') mm)

FIG.3

FIGA

Pla t_e - i : r i

F.-ig. 5 . T h i r d i n s t a r J cva of Do oL * Lqu<,« (X ^.li» jmO

F i g . G. t oov t h i n t t a r I f u v a £>f h^ <' vigvci. (X 2 .61 nm^.

F1G.5

FIG.6

P l a t e - i y

I if;» ?„ Pifi„h i n s t a r larva cf D„ f ^xii.cjua (T. 2 . 1 3 am)

F i g . 8 . r..^:;i:ir' i r i s c a r ?.arva cf P., '',«liq^M.,. (it 2c 7;; - m)

FIG.7

FIG. 8

Plate-V

Fig. 9. Male pupa of D. obliqua (X O.JG mm).

Fig. 10, Femalr' znipa ";" 1= .-' iA'*H£ ^^ 0.. t> mm)

FIG.9

FIG.10

Plate-VI

Fig. 11. Adult male of D. obiiqua (X 0.36 mm).

Fig. ]?.. Adult f.emi-.d e r^f D. obJiqua (X 0.43 mm).

FIG-11

FIG.12

FOOD CONSUMPTION AND UTILIZATION

30:

Insects as a group feed upon a remarkably diverse

list of organic substances. At the same time most species

show a high degree of selectivity in their choice of food.

According to Gordon (1959) "competition and natural selection

gradually drive and bind each species to a specialised food

supply that it can utilize more efficiently than any of its

competitors". Although the quantitative nutritional

requirements of growing insects seems to be uniform (Fraekel/

1953, 59; House, 1962). It seems apparent that adaptive

nutritional differences must be sought on a quantitative

level and that a meaningful comparative nutrition of insects

will not emerge until quantitative studies are emphasised

(Waldbauer, 1968).

The nutritional ecology of insects is the study

related to the success of the insect in the ecosystem, host

plant relationship and the survivalance of the pest in case

of D_. obliqua most of the work done in the area of

nutritional ecology is on the food preference or on the

insect plant relationship. But for the better understanding

of the life history phenomenon like growth development and

reproduction, it becomes essential to study the food

consumption and utilization indices.. The amount, rate and the

quality of the food consumed by the larvae and the adults

determine its performance of mating success, timing and

extent of reproduction, dispersal ability and the probabilitv

:31:

of survival and the quality of off springs produced. Thus the

consumption and utilization link the physiological,

behavioural, ecological and the evolutionary aspects of

insect life. Looking into the immense importance of the

nutritional ecology, the present work was undertaken.

:32:

RESULTS

Consumption Index (C.I.) :

The consumption index explains the rate at which

nutrients enter into the digestive system. The feeding is

basically governed by the massiveness of the food/ water

content and other physico-chemical properties of the food

material. Therefore the rate of intake of food is a function

of response to feeding. The amount of food consumed

increases from third to the last instar with slight decrease

in the fifth instar (Table - 3, Fig. 20) the consumption! index

shows an alternating decrease and increase in successive

developmental stages (Table - 4, Fig. 15).

Growth Rate (G.R.) :

The growth rate explains how much of dry matter

increased in the body of the insect per day per mg of the

body weight. It directly affects the speed of development

which depends on the quality of the food/ physiological stage

and the environmental factors. The initial instars show

higher growth rate which gradually goes on decreasing in

successive developmental stages therefore it is negatively

related to the stage of development (Table - 4, Fig. 16).

33:

Efficiency of conversion of ingested food (ECI) :

The efficiency of conversion of ingested food

basically depends on the amount of food eaten and the weight

gained by the larvae. The ECI values decrease in the

subsequent developmental stages proceeding from third to the

sixth instar larvae with a slight increase in the fifth

instar larvae. The third instar larvae show the maximum ECI

value thus making an alternating pattern of increase and

decrease (Table - 4, Fig. 18).

Efficiency of conversion of digested food (ECD) :

The efficiency of conversion of digested food meaning

the amount of energy devoted for the maintenance of

physiological functions of the insect. The observations

reveal that the maximum amount of energy is used in the

fourth instar larvae and the least is consumed by the third

instar larvae (Table - 4, Fig. 19). The ECD values

considerably decrease in the fourth instar making the

remaining values higher, otherwise the pattern is a decrease

down the table.

Approximate Digestibility (A-D.) :

The approximate digestibility is the function of the

food ingested and the waste ejested. The AD value is lov/est;

in the third instar which increases significantly in the

34:

fourth instar and then decreases in the following instars

(Table - 4, Fig. 17). Although the food consumption generally

increases with age but the digestibility fluctuates. The

shoot up in the AD values can be attributed to the required

storage of the food for the prepupal and the pupal stages

during which they do not feed.

The dry weight of the subsequent larval instars show

a gradual increase but the difference between the minimum and

the maximum values is not very high. Whereas the increase in

the fresh weight of the larval instars initially increases

slowly but it shoots up suddenly in the last two i.e. V and

VI instars. This shoot up is probably because of the excess

food consumed which contains very high water content (Fig.

21).

The relative increase in the size and weight do not

follow the Dyars law and Pirzibrams rule. The size increases

slowly in the beginning and shoots up in the III/ IV and the

V instar than slightly increases in the last instar. Whereas

the weight increases gradually with a constant speed upto the

IV instars and then shoots up in the last two instars (Fig.

22) .

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:35:

DISCUSSION

It has been observed that the food intake and the

weight gained by the larvae is directly correlated with the

age. Thus as the larvae grow older and enter into subsequent

instarsI gain comparatively more weight and consume more food

(Bhat & Bhattachacya, 1978; Prasad & Chand, 1980; Warrington,

1985 and Sharma & Tara, 1988). Similar results have been

found in the present study but there is slight fluctuation in

the food consumption value of the fifth instar larva.

The consumption index, which is the rate of food

intake per unit weight per day gradually decreases with age

(Babu et al., 1979). However, in the present study there is

a slight increase in the CI value of the fourth and the sixth

instar thus presenting an alternating pattern of increaseand

decrease. The decreasing trend has been reported in Podius

maculiventris (Mukerjee & LeRoux, 1969), Schistocerca

gregaria and Locusta migratoria (Mehrotra et al., 1972) in

Spodoptera litura (Prasad, 1973; Bhat & Bhattacharya, 1978)

and in ^. litura and D. obliqua (Sharma & Tara, 1988).

The growth rate determines how much of dry matter

increased in the body of the organism per day per unit weight

of the body. In the present study the growth rate show

decreasing trend with the advancement of age. Similar

patterns have been observed in L . decemlineata (Chlondy,

1967), _L. migratoria (Bennakkars et al . , 1970), L. migracoria

36:

and S . gregaria (Mehrotora et al . , 1972), S . litura (Bhat &

Bhattacharya/ 1978) and in £. litura and £. obliqua (Sharma &

Tara, 1988).

The approximate digestibility in Lepidopterans is

found to fall in the subsequent instars (Waldbauer, 1968

Kogan & Cope, 1974; Mathavan & Bhaskaran, 1975; Mackey, 1978

Haukioja et al., 2978; Valentine & Talerico, 1980

Warrington, 1985). Similar pattern have also been reported

among Acridids (Mordue & Hill, 1970; Mehrotra et al., 1972;

Khalsa, 1973; Gupta & Vats, 1980 and Vir & Jindal, 1983). But

in the present study the AD value increases in the fourth

instar and then decreases in the remaining instars.

Fluctuations in AD values have also been reported for

Atractomorpha precautionis (Khalsa, 1973), Pseudoplusia

includens (Kogen & Cope, 1974), S_. litura (Bhat &

Bhattacharya, 1978), S-. litura and _D. obliqua (Sharma & Tara,

1988).

The efficiency of conversion of ingested food varies

considerably among the insects because it depends on the food

consumed which is quite variable. In the present study there

is a gradual decrease in the ECI values of the subsequent

instars with a slight shoot up in the fifth instar. Similar

patterns have been reported in A.. precautionis (Khalsa,

1973), £. includens (Kogen & Cope, 1974) and S_. litura (Bhat

& Bhattacharya, 1978). In most of the Lepidopterans the ECI

value tends to decrease VNrith the advancement of aqe

:37

(Waldbauer, 1968; Mackey, 1978 and Scriber, 1979) but this is

not always the case, the ECI fluctuates irregularly with the

final instar having the highest value (Mathavan & Bhaskaran,

1975). But in S . litura there is a gradual increase in ECI

values but a sudden decrease in the final instar larva

whereas in £. obliqua initially the value increases gradually

and then decrease occurs in the fourth instar (Sharma & Tara,

1988).

The efficiency of conversion of digested food

expresses the amount of energy devoted for the maintenance of

physiological functions of the insect. In the present study

the ECD value decreases with the advancement of developmental

stages but their occurs slight increase in the fifth instar

larva/ it means the fifth instar requires more energy for its

biological maintenance. Similar observations were reported

for _S. litura and _D. obliqua (Bhat & Bhattacharya/ 1978;

Sharma & Tara, 1988). It has also been found that the ECD

values fluctuate irregularly in the initial instars but

increases in the fourth, fifth and the sixth instar (Mackey,

1978). The ECD gradually increases in later instars

indicating that more energy is left for the growth and

reproduction (Banerjee & Haque, 1984).

REFERENCES

38:

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Ali, M.I. (1988). A survey of the insect pests of soybean in North Bangladesh, their damage and occurrence. Trop Pest. Manag . , 34(3): 328-330.

Aral, T. and Mabuchi, M. (1979). Effects of light conditions on the timing of adult activities in Hyphantria cunea Drury (Lepidoptera : Arctiidae). Jap. J. Appl. Zool . , 23(3): 162-169.

Atwal, A.S. (1986). Agricultural pests of India and South East Asia. Kalyani Publishers, New Delhi - Ludhiana, pp. 507.

Babu, M.H.; Bhattacharya, A.K. and Rathore, Y.S. (1978). Developmental behaviour of three Lepidopterous pests on soybean and green gram. Z. Agnew. Ent ., 85(1): 108-112.

Babu, M.H.; Bhattacharya, A.K. and Rathore, Y.S. (1979), Rate of intake, growth and digestibility of three Lepidopterous insects. Z. Agnew. Ent . , 87(3): 322-327.

Bakhetia, D.H.C. and Brar, K.S. (1982). Pest control in rapeseed and mustard. Ind. Farm. , 32(8): 60-63.

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Banerjee, T.C- and Hague, N. (1984). Dry matter budgets for Diacrisia casignetum larvae fed on sunflower leaves. J. Insect Physiol. , 30(11): 861-866.

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^Banerji, B. (1971). Eclosion and oviposition rhythms of Andraca bipunctata Walker (Bombycidae : Lepidoptera) with a model for determining theoretical moth densities from eclosion rate. Indian J. Ent . , 33(4): 411-418.

Bennakkars, M.A.; Meison, H.Q. and Scheras, J.M.J.D. (1970). Influence of temperature and food on growth and digestion of fifth instar larvae and adult of Locusta. J. Insect Physiol., 17(11): 871-883.

: 3 9 :

*Benson, C . F . C . and C h a t t e r j e e , N.C. ( 1 9 4 0 ) . P o s s i b i l i t i e s of c o n t r o l of Lan tana (Lantana a c u l e a t a L i n n . ) by i n d i g e n o u s i n s e c t p e s t s . I n d . F o r . R e c . (N.S . E n t . ) . , 6 ( 3 ) : 3 9 - 8 4 .

B h a r d w a j , S . P . and B h a l l a , O . P . ( 1 9 7 7 ) . R e c o r d of i n s e c t p e s t s of s o y b e a n i n Himachal P r a d e s h . I n d i a n J . E n t . , 3 8 ( 3 ) : 2 8 6 - 2 8 9 .

B h a t , N . S . and B h a t t a c h a r y a , A.K. ( 1 9 7 8 ) . Consumpt ion and u t i l i z a t i o n of soybean by S p o d o p t e r a l i t u r a ( F a b r i c i u s ) a t d i f f e r e n t t empera t u ' r e s . I n d i a n J . E n t . , 4 0 ( 1 ) : 1 6 - 2 5 .

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Chand/ P. ( 1 9 7 9 ) . U t i l i z a t i o n of food by B i h a r h a i r y c a t e r p i l l a r / D i a c r i s i a o b l i q u a W a l k e r . I n d i a n J . E n t . , 4 1 ( 3 ) : 2 3 7 - 2 3 9 .

Chaudha ry , R . R . P . and B h a t t a c h a r y a , A.K. ( 1 9 8 6 ) . Bioecology of L e p i d o p t e r o u s i n s e c t s on winged bean , P sophoca rpus t e t r o g o n o l o b u s ( L i n n a e u s ) DeCandole . Mem. I n t . S o c . I n d . , 1 1 : p p . 130 .

^Ch londy , J . ( 1 9 6 7 ) . The amount of food consumed and p r o d u c t i o n o u t p u t of l a r v a e of C o l o r a d o b e e t l e ( Lep t inb ta r sa d e c e m l i n e a t a ) S a y . ) . E k o l o g i a Po l ska -S e r i a A . , 1 5 : 5 3 1 - 5 4 1 .

Cohen, A.C. and P a t a n a , R. ( 1 9 8 4 ) . E f f i c i e n c y of food u t i l i z a t i o n by H e l i o t h e s zea ( L e p i d o p t e r a N o c t u i d a e ) fed on a r t i f i c i a l d i e t s o r g r e e n b e a n s . Can. E n t o m o l . , 1 1 6 ( 2 ) : 139 -146 .

Crocomoard , W.B. and P a r r a , J . R . P . ( 1 9 8 5 ) . Consumpt ion and u t i l i z a t i o n of m i l l e t , wheat and sorghum by Spodop te r a f r u g i p e r d a (Smi th) ( L e p i d o p t e r a : N o c t u i d a e ) . Rev . B r a s . E n t o m o l . , 2 9 ( 2 ) : 2 2 5 - 2 6 0 .

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Deshmukh, P . D . ; R a t h o r e , Y . S . and B h a t t a c h a r y a , A.K. ( 1 9 8 2 ) . E f f e c t of t e m p e r a t u r e on t h e g r o w t h and deve lopment of D i a c r i s i a o b l i q u a (Walker) on f i v e h o s t p l a n t s . I n d i a n J . E n t . , 4 4 ( 1 ) : 2 3 - 3 2 .

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Dhuri, A.v.; Singh, K.M. and Singh, R.N. (1984). Incidence of insect pests in blackgram, Vigna mungo (L.) Hepper. Indian J. Ent . , 46(3): 270-276.

*Gangrade, G.A. (1976). Terminal technical report on the project on assessment of effects on yield and quality of soybeans caused by major Arthropod pests Jabalpur, India. J.N. Krishi Vishwa Vidyalaya; 5; pp. 144.

Gangrade, G.A.; Singh, O.P. and Matkan, S.M. (1977). Soybean yield losses in response to damage by varying levels of three Lepidopterous larvae. Indian J. Ent . / 37(3): 225-229.

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Goel, S.C; Kumar, A. and Singh, J. (1986). Variation in nutritional behaviour of hairy caterpillars, Diacrisia obliqua (Wlk.) (Arctiidae) and Lymantria marginata Wlk. (Lymantriidae). U.P.J. 2ool~ 6(2) : 215-223.

*Gordon, H.T. (1959). Minimal nutritional requirements of the German roach, Blatella germanica L., Ann. A.Y. Acad. Sci., 77: 290-351.

Gupta, A. and Vats, L.K. (1980). Food consumption, assimilation and tissue growth in Phlaeoba infumata (Orthoptera : Acrididae). Indian J. Ecol . , 7(1): 61-70.

Gupta, B.M.; Dabi, R.K.; Gupta, H.C. and Sharma, S.K. (1979). Effect of host plant on the larval development of Diacrisia obliqua Walker. J . Entomol. Res., 3(2): 224-226.

Gupta, S.C. (1982). Effect of food on the development of Diacrisia obliqua Walker. Geobios., 9(5): 215-219.

Haq, M.; Karim, A.N.M.R. and Alam, S. (1984). Preliminary study of varietal reaction of soybean cultivars to leaf defoliators. Trop. Grain, Leg. Bull., 29: 35-37.

: 4 1 :

Haq, M.; Karim, A.N.M.R- and Alam, S . ( 1 9 8 5 ) . P r e l i m i n a r y s t u d y of v a r i e t a l r e a c t i o n of s o y b e a n c u l t i v a t o r s t o l e a f d e f o l i a t o r s . T r o p . G r a i n , Leg . B u l l . / 30 : 2 9 - 3 0 .

* H a u k i o j a , E . ; N i e m e l a / P . ; I s o - I v o r i , L., O j a l a / H. and Aro/ E.M. ( 1 9 7 8 ) . B i r c h l e a v e s a s a r e s o u r c e for h e r b i v o r e s I . V a r i a t i o n i n t h e s u i t a b i l i t y of l e a v e s . R e p o r t of t h e Kevo S u b a r t i c R e s . S t a t . / 14: 5-12-

*House/ H.L. ( 1 9 6 2 ) . I n s e c t N u t r i t i o n . R e v . B i o c h e m . , 3 1 : 6 5 3 - 6 7 2 .

H u s s a i n , T. ( 1 9 8 2 ) . Ma t in g b e h a v i o u r and o v i p o s i t i o n of D i a c r i s i a o b l i q u a Wlk. ( L e p i d o p t e r a : A r c t i i d a e ) . Pak. J . S c i . I n d . R e s . / 2 5 ( 6 ) : 2 3 8 .

I s l a m / B.N. and Alam, M.Z. ( 1 9 8 0 ) . Mating b e h a v i o u r of J u t e h a i r y c a t e r p i l l a r / D i a c r i s i a o b l i q u a Walker ( L e p i d o p t e r a : A r c t i i d a e ) I I . R a t e of c a t c h of male moth by v i r g i n f e m a l e t r a p s . A p p l . E n t . Z o o l . / 1 5 ( 3 ) : 2 9 9 - 3 0 9 .

I s l a m , W.; Ahmad, K-N. ; N a r g i s , A. and I s l a m , U. ( 1 9 8 3 ) . O c c u r r e n c e , a b u n d a n c e and e x t e n t of damage caused by i n s e c t p e s t s of g r o u n d n u t ( A r a c h i s h y p o g a e a ) . Malays ia A g r i l . J . , 5 4 ( 1 ) : 1 8 - 2 4 .

K a b i r , A. and M i a h , H.A. ( 1 9 8 7 ) . E f f e c t o f c e r t a i n food p l a n t s on t h e g r o w t h and deve lopmen t of J u t e h a i r y c a t e r p i l l a r , D i a c r i s i a o b l i q u a W a l k e r . B a n g l a d e s h J . Z o o l . , 1 5 ( 2 ) : 1 7 7 - 1 8 2 .

Kapoor / K.N. ; G u j a r a t i , J . P . and G a n g r a d e , G.A. ( 1 9 7 2 ) . Bionomics of Lamprosema i n d i c a t a F a b r i c i u s ( L e p i d o p t e r a : P y r a l i d a e ) a p e s t of s o y b e a n in Madhya P r a d e s h . I n d i a n J . E n t . , 3 4 ( 2 ) : 1 0 2 - 1 0 5 .

K h a l e q u e , M.A. ( 1 9 8 3 ) . Soybean seed q u a l i t y and s t a n d a r d e s t a b l i s h m e n t i n B a n g l a d e s h . P r o c . Conf. S c i t . A s i a . 2 5 - 3 1 .

*Kha l sa / M.S. ( 1 9 7 3 ) . Hos t p l a n t s e l e c t i o n b e h a v i o u r of Autographa p r e c a u t i o n i s (Guence) ( L e p i d o p t e r a : N o c t u i d a e ) . P h . D . t h e s i s / Univ . of I l l i n o i s , Urbana, I l l i n o i s , U . S . A . , p p . 2 8 6 .

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: 4 2:

* K r i s h n a h , K. ; P r a s a d , V.G. and Mohan, N . J . ( 1 9 7 5 ) . New r e c o r d s of a l t e r n a t e h o s t s of Tobacco c a t e r p i l l a r , Spodop te ra l i t u r a ( F a b . ) ( L e p i d o p t e r a : N o c t u i d a e ) and B i h a r h a i r y c a t e r p i l l a r , D i a c r i s i a o b l i q u a Wlk. ( L e p i d o p t e r a A r c t i i d a e ) . C u r r t . S c i . , 44(4) : 3 0 4 - 3 1 2 .

^ K r i s h n a m u r t h y , M.M.; S a y i , I . V . and Rao , B.H.K.M. ( 1 9 7 4 ) . Notes on t h e b i o l o g y of s h o o t and b l o s s o m webber, Mocal la m o n c u s a l i s Walker ( L e p i d o p t e r a : P y r a l i d a e ) of c a shew. I n d i a n J . E n t . , 3 6 ( 1 ) : 7 6 - 7 7 .

L a i , L. and M u k h e r j i , S . P . ( 1 9 7 8 ) . Growth p o t e n t i a l of D i a c r i s i a o b l i q u a Walker in r e l a t i o n t o c e r t a i n food p l a n t s . I n d i a n . J . E n t . , 4 0 ( 2 ) : 1 7 7 - 1 8 1 .

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M a r t i n e z , E . ; Novo P a d r i n o , J .M. and H e r n a n d e z , H . F . ( 1986 ) . L i f e c y c l e of H e l i o t h e s v i r e s c e n s F a b r i c i u s ( L e p i d o p t e r a : N o c t u i d a e ) under a m b i e n t c o n d i r i c n s in t h e l a b o r a t o r y . C e n t r e A g r i c o l a , 1 1 ( 2 ) : 3 9 - 4 5 .

*Mathavan , S. and B h a s k a r a n , R. ( 1 9 7 5 ) . Food s e l e c t i o n and u t i l i z a t i o n i n a Danid b u t t e r f l y . O e c o l o g i a ( B e r l . ) , 18 : 5 5 - 6 2 .

*Mathur , A.C. ( 1 9 6 2 ) . Food p l a n t s p e c t r u m of D i a c r i s i a o b l i q u a Wlk. ( L e p i d o p t e r a : A r c t i i d a e ) i n f e s t i n g m e d i c i n a l p l a n t s i n Jammu. I n d i a n J . Ent . , 2 4 ( 3 ) : 2 8 6 - 2 8 7 .

*Mehra , B.P. and S h a h , B-N. ( 1 9 6 6 ) . B ionomics of Hemithea t r i t o n a r i a Walker ( L e p i d o p t e r a : G e o m e t r i d a e ) , a p e s t of Moghania m a c r o p h y l l a ( W i l d . ) O. K t z e . Ind i an J . E n t . , 2 8 ( 4 ) : 3 4 5 - 3 5 0 .

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43:

*Mordue (Luntz), A.J. and Hill, L. (1970). The utilization of food by adult female desert locust/ Schistocerca gregaria. Entomol. Exp. Appl., 13: 352-358.

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: 4 4 :

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:45:

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*Not seen in original

Fig. 13. Showing the incubation period/ larval and

pupal durations in D. obliqua.

330-

300-

270-

210-

180-

150-

< a: Q 1201

SO-

SO-

3 0 -

L_L

i-

11 INCUBATION I

*—

II III IV Y VI PRE--lorval duration ^ PUPA

PUPA

FIG.13

Fig. 14. Showing the preoviposition, ovipositiori/

postoviposition periods and the longevity of

unmated and mated adults of D. obliqua.

AOO-

350-

300-

• 250-

2 ZOO-

ISO-

100-

50-

1

ir 1

PRE- OVr- POST- MALE FEMALE MALE FEMALE OVf POSITION (unmated) (mated)

FIG-U

Fig. 15. Showing the relationship between the

consumption index (C.I.) and the larval

stages of D. obliqua.

lU •^

X UJ Q

2-OH

1-6 H

Q. 1-2H

CO

o <-> 0-8 H

0-4-4

—r-III IV

-r V

—r VI

LARVAL INSTARS

FIG.15

Fig- 16. Showing the relationship between the growth

rate (G.R.) and the larval stages of D.

obliqua.

o-4i

} -<

0-3H

5 o g 0.2-

0-H

in —T"

IV - r V

—r VI

LARVAL INSTAR

F I G . 1 6

Fig. 17. Showing the relationship between the

approximate digestibility (A.D.) and the

larval stages of D. obliqua.

80-

Q

< 60-

^0-

2 0-

—r-

in —r VI IV

1^

V

LARVAL INSTARS

FIG.17

Fig. 18. Showing the fluctuations in the values of

efficiency of conversion of ingested food

(E.C.I.) in the subsequent larval instars of

D. obliqua.

25 1

o UJ

20-

15

10-

5 -

—r-III IV

T V

LARVAL INSTAR5

VI

FIG.19

Fig. 19. Showing the fluctuations in the values of

efficiency of conversion of digested food

(E.C.D.) in the subsequent larval instars of

D. obliaua.

Q

U lis

1A0-

120-

100-

8 0 -

6 0 -

^ 0 -

2 0 -

—I r-III IV

- r V

—r VI

LARVAL INSTARS

FIG.19

Fig. 20. Showing the relationship of food consumed in

the subsequent larval instars of D. obliqua.

£

Q

ID

O O Q O

2

500-

^00-

300-

200-

100-

—r-III

—r IV

Y —r-VI

LARVAL INSTARS

FIG. 20

Fig. 21. Showing the relationship between fresh and

dry weight in the subsequent larval instars

of D- obliqua.

600- dry weight

X ^ fresh weight

en g

a: <

X h-U-

o X o UJ

500-

/»00-

300-

2 200-

100-

-1-

11 III T IV V vr

LARVAL INSTARS

FIG-21

Fig. 22. Showing the relationship between the larval

size and weight in the subsequent larval

instars of D. obliqua.

35H h700

'S ize

E J UJ N

in

30H

25H

20-

15

10-

X «• weight -600

h500

Moo

h300

h200

-m

£

X

o

5:

II m —r-IV Y

— I —

VI

LARVAL INSTAR5

FIG.22