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) .
ft) U 4-> Q) (U 4J U >« U (0 X <" (0
D ^ Ky
-O - H (U rH S 12 3 0 m c , 8 Q I
0 ^ 0 0
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+1
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a: M 3 O C!i O
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+1
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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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Atwal, A.S. (1986). Agricultural pests of India and South East Asia. Kalyani Publishers, New Delhi - Ludhiana, pp. 507.
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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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: 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 .
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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 .
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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 .
L a i , S . S . ( 1 9 8 5 ) . A r e v i e w of i n s e c t p e s t s of mung bean and t h e i r c o n t r o l i n I n d i a . T r o p . P e s t Manag . / 3 1 ( 2 ) : 105-114 .
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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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R i z z o , H.F. ( 1 9 7 9 ) . B i o l o g y of S p i l o s o m a v i r g i n i c a ( F . ) ( L e p i d o p t e r a : A r c t i i d a e ) . Rev. S o c . E n t . A r g . , 38 : 8 3 -84 .
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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
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