Proc. Indian Acad. Sci. (Anim. ScL), Vol. 97, No. 4, July 1988. pp. 339-345.

© Printed in India.

Functional response of Eocsnthecon« Iurcellst« (Wolff.) (Heteroptera:

Pentatomidae) in relation to prey density and defence with reference to
its prey Latoia lepida (Cramer) (Lepidoptera: Lemacodidae)

R SENRAYAN
Entomology Research Institute, Loyola College, Madras 600 034, India
MS received 8 March 1988; revised 12 May 1988
Abstract. The functional responses of Eocanthecona furcellata (Wolff.) that predates on its
caterpillar prey, Latoia lepida (Cramer), a sporadic pest of many agricultural crop plants are
presented using Holling's 'disc' equation. The results indicate the importance of prey density
and defence in influencing the attack response of.the predator. Further, the phytophagous
habits of this predatory bug and its ability to derive nourishment from plant sources during
periods of prey absence are also discussed.
Keywords. Predator; functional response; pentatomid.

1. Introduction

The potential of Eocantheconafurcellata (Wolff.), a polyphagous insect predator, as a

natural control agent of insect pests has been documented (David and Basheer 1961;
Ghorpade 1972;R.ai 1978). E.furcellata is an effective predator of the slug caterpillar,
Latoia lepida (Cramer), a pest of several crops. However, little is known of its predatory
efficiency and functional response with reference to its prey, L. lepida. Interestingly,
like many predatory heteropteran bugs E. furcellata is also known to feed on plants.
The results presented here pertain to the functional responses of the adults of
Ei furcellata in relation to prey density and defence of L. lepida using Holling (1959)
'disc' equation, besides describing the nature of the phytophagous habit of E.furcellata.

2. Materials and Methods

Nymphs and adults of E. furcellata were collected from Cassia sp. and mango trees
from private gardens in Madras, and a colony of E. furcellata was raised.from these
collections in the laboratory. For culturing the predators glass troughs (25 x 10 cm)
were used and their open tops were covered with muslin cloth for proper ventilation.
Young mango leaves, provided inside the predator rearing troughs, served as
oviposition sites. Bggs and young larval stages of L. lepida were collected from
mango plants and were reared in the laboratory using glass chimneys with mango
foliage. Developmental stages of the prey were determined based on the total length
and head width.

2.1 Functional response tests

The objective of the experiment was to find out (i) the prey consumption rate by the
predator at different prey densities with various stages in a restricted milieu in the
laboratory and (ii) the influence of prey defence on the functional response of the
339
340 R Senrayan

predator. The prey insect were packed loosely in the jar with mango foliage, that
were changed periodically, since the leaves were not preferred when dry. The prey
were randomly distributed in the foliage of each jar and the larvae killed by the
predator were replaced daily and the prey number was maintained at all densities.
Both males and females were taken equally in replicating the tests for each prey
density and at each prey stage level.
To evaluate the influence of prey defence on the functional response of the
predator, experiments were repeated with 'defenceless' larvae. A small metal pin was
used to hold the thoracic segment of each larva so that the larva could not defend
itself. Care was taken not to inflict any injury to the larva and the larvae injured were
discarded. For prey densities beyond 16, the volume of the experimental jars was
reduced instead of increasing the numbers of the prey, so that counting and replacing
the larvae were done easily without any disturbance to the experiment.

2.2 Procedure for application of ,disc' equation

The various parameters followed in the 'disc' equation to describe the functional
response of E. furcellata adults at different prey densities are listed below.
(i) x-Prey density.
(ii) y-Total number of prey killed in a given period of time, Tt.
(iii) xly-The attack ratio.
(iv) Tt-Total time in days when prey was exposed to the predator.
(v) b-Time taken for handling each prey by the predator.
(vi) a-'Rate of discovery' per unit of searching time.
Usually the prey discovery was instantaneous with very little time being required.
Although the parameter 'rate of discovery' (a) is infinite, the predator did spend some
time in searching for the prey at lower prey density. Assuming that the predation is
proportional to the prey density and to the time spent by the predator in searching
prey (Ts) the expression of the relationship is:
y=a Tsx. (1)
But time available for searching is not a constant. It is reduced from the total time
(Tt) by the time spent handling the prey. If we presume that each prey requires a
constant amount of time 'b' for consumption, then
Ts= Tt-by. (2)
Substituting (2) in (1)
Y=a(Tt-by)x, (3)
or
Y= Tt axil + abx (HoIling 1959). (4)

3. Results

E.furcellata is an active predator and it attacks the prey in spite of violent resistances
of the prey. Further, prey density and predator age alter the response of the predator
 Predator-prey interactions 341

at all prey stages. In view of these factors, the functional response of the predator was
studied with reference to the prey density, prey defence and predator age.

3.1 Prey density

The total time (Tt) during which the prey are exposed to predation was 20 days. The
number of prey killed increase with the prey density upto x = 32, in the case of first to
third instar prey stages but beyond x = 32, no further increase in 'y' was evident.
Hence the predator has all the prey quantity it can feed upon without any search
effort, when the prey density is above 32. On the contrary, the number of prey killed
increased only up to the prey density of 16, in 4th and 5th instar prey stages (figure 1).
The mean maximum predation of defenceless larvae by a predator in 20 days was 91,
68,6, 51,6,42,2 and 34·4 from the 1st to the 5th instar prey respectively. Similarly, for
free larvae the value was 89,9,67,6,44'6,35,1 and 28·5 from the 1st to the 5th instar
prey respectively, indicating the role of prey density and defence in influencing the
functional response of the predator.

3.2 Prey defence

The maximum predation at highest prey density will be represented by 'K' and the
value remains higher in the 1st instar than the 5th instar prey. In defenceless larvae,
the attacking time is taken as 0 in terms of days and the time taken to feed varied
from 0·020 to 0·089 days in the 1st to the 5th instars of the prey. The third
component, in addition to attacking and feeding, viz. the interval between the time of
completion of feeding and the time for the predator to attack again is estimated as,
ob' value-feeding time for all stages. The value of Ob' assumed to be constant for all
prey densities. At prey densities below 32, the predator spent sometime searching its
prey and therefore the searching time was calculated following the formula, Ts = Tt-:

85
75
55
,,_-'0'-------0.----0"
I Inslor It l:n star
"0 m Inslar
a.§
QI

15
'-_""----'-_-'----'-_-'--l
a
'---'-_~.....I----''---'-...J '---:~-'--'---'--'-:-"

u 8 24 64 8 24 64 24 64
>-
l!! - - Defenceless prey
Cl. 451- I-
---- Free prey
25 V;-~:~:~-----
5 I I I
8 24 64 8 24 64
Prey densily Ixl

Figure 1. Functional response of E. furcellata adults to the density of L. lepida, when the
prey is defenceless (D) and free to defend itself (F).
342 R Senrayan

by in terms of days (table 1). Another parameter 'a' the rate of discovery, was also
estimated and the value was higher during the 1st instar stages than the 5th instar.
Unlike defenceless larvae the free larvae showed resistance especially during
mature stage, hence the attacking time of the predator was calculated and the value
was relatively negligible in the lst and 2nd instar larvae. However, the predator took
0'00041,0,00 11,0'0027 days to paralyse the 3rd to 5th instar stages of the prey. The
ob' value of the predator increases from early to mature stages of the prey and from
defenceless to free prey, and this is attributed to the following: (i) greater time is
taken by the predator to paralyse the larger prey, (ii) increased amount of food
consumption in a larger prey and (iii) longer time interval taken by the predator as
resting period to attack another prey due to satiation (table 2). Similarly, the 'a' value
was reduced in free prey as compared with defenceless larvae and from early to
mature prey and it could be due to 3 reasons: (i) 'a' value is the proportion of prey
attacked successfully per unit of searching time, (ii) continuous prey searching in
view of the defensive activities of the larvae and (iii) predator search harder to find
prey at smaller prey stages due to hunger.

3.3 Predator age

The importance of predator age in the functional response of the predator is quite
obvious from the data provided in table 2. The values of 'K', 'a' and 'b' have been
calculated at two age intervals to illustrate the influence of predator age. Higher
values in 'a' and lower values in 'b' was recorded during younger stages of the
predator presumably due to active searching, quicker time taken to paralyse the prey
and shorter time interval between successive attacks. In contrast, increased values in

Table 1. Summary of calculations used in fitting the functional response curves for d~fence-
less prey (D) and free prey (F) for a first instar L. lepida.

Prey Prey Max. Days Days Days Att.

Prey Den. att'd y per y all y's searching ratio yJx 7 T,
type (x) (y) (K) (b) (by) (T,) (yJx) (a)

I 18·6 4{)9 IHI 18-6 1·17

2 35·0 7-70 12030 17'5 1-42
4 61·4 13-51 6·49 15·3 2-37
D 8 78·8 17·32 2-66 9·9 3-70
16 91·4 zo-n 5·7
32 90'6 91·0 0·22 19·93 2-8
64 91'0 20-02 1·4
y= ai Tt>: by)x= 2'17(20-0'22y} x 2'17
I 18·8 4·14 15·86 18·g 1'19
2 35-6 7-83 12·17 17·8 1'46
4 62-0 13·64 6·36 15·5 2·44
F 8 76·4 16·81 3·19 9·5 2·99
16 89'4 19-67 5·5
32 89·8 89·9 0·22 19·76 2·8
64 90·4 19·89 1·4
y=2'02(20-0'22y)x 2'02
Total time 20 days.
 Predator-prey interactions 343

Table 2. Functional response of E. furcellata adults on

various stages of the prey, L. lepida at two different age
levels.

Category
Prey stage Dor F KIT, b a
10 days 4·86 0·206 3·095
D 20 days 4·55 (}220 2·174
I instar
10 days 4·90 (}204 2-491
F 20 days 4-49 (}220 2·021
10 days 3-88 0·258 3·256
D 20 days 3-43 (}290 2·460
n instar
10 days 3-83 (}261 2·232
F osoo 2·560
20 days 3·38
10 days 2-72 0·368 30202
D 20days 2·58 0·390 3'520
HI instar
10 days 2-39 (}419 2-091
F 20 days 2·23 0·450 1'627
10 days 2·26 (}443 1-740
D 1"888
20 days 2'11 0'470
IV instar
10 days 1·99 (}SOI 1·366
F 20 days 1·75 (}570 1"438
10 days 1"87 (}534 2-493
D 1"830
20 days 1·72 0·580
V instar
10 days 1-61 0·622 1·248
F
20 days 1"43 0·700 1·090

'b' and decreased values in 'a' during older age of the predator is due to longer time
interval between successive attacks and reduced searching ability.

3.4 Plant feeding

E. furcellata also feeds to some extent on plants and the frequency of plant feeding is
higher at older age and during prey scarcity. The estimates of 'a' and 'b' values are
slightly biased because of time spent on plant feeding has not been deducted from the
time available for searching and further the process of plant feeding itself is not
continuous. Supplementary plant feeding reduced the nymphal developmental period
considerably so also highest percentage survival in nymphs. Plant diet alone proved
insufficient for normal development and survival of the predator, but however plant
feeding sustained the nymphs and adults for a considerable period during prey
absence.

4. Discussion

Predators are always selective in the choice of their prey, from broad levels of
preference for particular prey species to fine levels of preferred shapes, sizes, colours
344 R Senrayan

and palatabilities (Law 1979). To describe and explain the functional response
HoIling (1959) proposed a 'disc' equation and successfully applied it to data on the
following predator-prey systems: Dohlbominus fulginosus (Nees) vs. N eodiprion
sertifer (Geoff.); (Bumett 1951, 1954); D. fulginosus vs. Neodiprion lecontei (Fitch)
(Bumett 1958); Chelonus taxanus Cress. vs. Anagasta kuhniella (Zell.) (Ullyett 1949a);
Cryptus inornatus Pratt vs. Loxostege sticticalis (L) (Ullyett 1949b); Nasonia
vitripennis (Wlk.) vs. Musca domestica L. (Debach and Smith 1941). Mukerji and
LeRoux (1969) explained the functional response of the predator Podisus
maculiventris (Say) on the prey GaLleria meLlonella (L) in relation to its size of the prey
and explained the positive applicability of Hoiling 'disc' equation. Morris (1963) also
applied the equation to the predator P. maculiventris fed on larvae of Hyphantria
cunea at different densities and confirmed its applicability within certain range of
prey density and time. However in contrast to these, Haynes and Sisojevic (1966)
were unable to apply 'disc' equation successfully to the data obtained on the
predator Philodromus rufus in relation to the densities of the prey. In the present
investigation the Holling 'disc' equation finds its applicability with reference to prey
density and defence upto a range of prey density. The results indicate that the prey
density is an important factor which modifies the rate of attack of the predator.
Further, prey defence also plays a vital role in influencing the response of the
predator at all prey densities and the rate of predation by E. furcellata decreases
'consistently with the age of the adult, presumably as a result of physiological changes
associated with lesser food requirements.
E. furcellata draws moisture and nourishment from plants in addition to prey
feeding, which possibly enable better survival rates at times of prey scarcity, since the
ability of a predator to derive moisture and nourishment from plant materials is
considered to be an adaptation that allows the predator to exploit the readily
available resource without starving during prey scarcity (Sweet 1960). Plant feeding
also helps highly mobile predators, providing food during migration in areas where
the prey may not be abundant (Waddill nd Shepherd 1974). Further, plant feeding
nature of predatory bugs increase the survival rate, reproductive capability and body
size (Kiman and Yeargon 1985; Niranjo and Stimac 1985; Ruberson et al 1986).
Stoner (1972) pointed out that the plant feeding nature of hemipteran predators
would enable them to persist in agricultural habitats when prey becomes scarce.
Hence, the supplementary plant feeding nature of E. furceLlata is an adaptive habit
for sustaining during the periods of prey scarcity or mobility in an agricultural
habitat. Further studies on the individual chemical components of prey and foliage
would enable us to device chemically defined artificial diets, so that mass rearing of
the predator can be achieved.

Acknowledgement

The author is thankful to Prof. T N Ananthakrishnan for guidance.

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 Predator-prey interactions 345

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