© Benaki Phytopathological Institute
Papanikolaou
et al.
60
Type II functional response seems to be
common among aphidophagous coccinel-
lidae species. Thus, a type II functional re-
sponse is recorded for
Cheilomenes sulphurea
Olivier male adults on
Aphis fabae
Scopo-
li (Hodek
et al
. 1984),
Coelophora inaequalis
F. fourth instar larvae and female adults on
Toxoptera citricida
Kirkaldy (Wang and Tsai,
2001),
Propylea dissecta
Mulsant fourth in-
star larvae and adults on
Aphis gossypii
Glov-
er (Omkar and Pervez, 2004),
Harmonia axy-
ridis
Pallas larval instars and female adults on
Myzus persicae
Sulzer (Lee and Kang, 2004),
Coccinella septempunctata
L. fourth instars
and adults on
M. persicae
(Cabral
et al
., 2009),
H. axyridis
and
C. septempunctata
third in-
star larvae and adults on
Aphis glycines
Mat-
sumura (Xue
et al.
, 2009),
Hippodamia var-
iegata
Goeze larval instars and adults on
A.
fabae
(Forhadi
et al.
, 2010) and
Adalia bipunc-
tata
L. female adults on
M. persicae
(Jala-
li
et al.
, 2010). However, few exceptions have
been reported, as Messina and Hanks (1998)
showed that plant architecture may alter the
type of functional response of
P. quatuor-
decimpunctata
L. fourth instars and adults,
where Lou (1987) reported type I and Hu
et
al.
(1989) type III functional responses for
Harmonia axyridis
Pallas adults.
Coccinellids’ handling time, i.e. the time
a predator spends on pursuing, subduing,
eating and digesting its prey, is not constant
through functional response experiments
(Hassell
et al.
, 1976). Hodek (1996) stated that
hungry aphidophagous coccinellids com-
pletely devour the first prey items they en-
counter and then become less competent at
exploiting prey. According to Papanikolaou
(unpublished data) this trait results from the
digestion process. Nevertheless, the estima-
tion of handling time based on mean values is
substantial for developing general models for
the description of density-dependent preda-
tion in a realistic manner (Hassell
et al.
, 1976).
In a previous study, Papanikolaou
et al.
(2011) showed that larval instars of
P. quatuor-
decimpunctata
exhibit a type II functional re-
sponse on
A. fabae
, when allowed to prey for
24 h. The present study investigated the func-
tional response of the predator on
A. fabae
at
12 h taking into account that handling time
is not constant through functional response
experiments. Furthermore, the appropriate
model was fitted to the data in order to pre-
dict the predation ability of
P. quatuordecim-
punctata
on several densities of
A. fabae
.
Materials and Methods
An
A. fabae
colony (originally from a stock
colony from the Biological Control Laborato-
ry, at the Benaki Phytopathological Institute)
was reared on potted fava bean,
Vicia faba
L., plants at 20 ± 1ºC (65 ± 2% RH, 16:8 (L:D)
h).
Propylea quatuordecimpunctata
was col-
lected in summer 2006 from corn,
Zea mays
L., plants infested by the corn leaf aphid,
Rhopalosiphum maidis
Fitch, in Arta Coun-
ty (Northwestern Greece, 21°0’0’’/39°10’0’’)
and was reared on
A. fabae
in cylindrical
Plexiglass cages (50 x 30 cm) at 25 ± 1ºC, 65
± 5% RH and a photoperiod of 16:8 (L:D) h.
Functional response experiments were
carried out at 20 ± 1°C, 65 ± 2% R.H., with a
photoperiod of 16:8 (L:D) h. The experimen-
tal arena consisted of a plastic container (12
cm height x 7 cm diameter) with a potted
V.
faba
plant (top growth was cut in order to
provide plants of 8-9 cm height), bearing
different densities of
A. fabae
(3-3.5 day-old
immature aphids to avoid reproduction dur-
ing the experiments). A single predator lar-
va of a certain instar, which was previously
starved from food for 12 h, was placed into
the experimental arena for 12 h. The prey
densities tested were 2, 4, 8, 16 and 32 aphid
nymphs for the 1
st
instar larvae of the pred-
ator, 2, 4, 8, 16, 32 and 64 nymphs for the
2
nd
instar larvae, and 4, 8, 16, 32, 64 and 128
aphids for the 3
rd
and 4
th
instar larvae.
In order to test for density-dependent
prey mortality, a logistic regression analy-
sis was conducted on the proportion of prey
eaten as a function of the initial prey densi-
ty, fitting the polynomial function (Juliano,
2001):
)NP NP NP P exp(
1
)NP NP NP P exp(
N
N
3
0 3
2
0 2
0 1
0
3
0 3
2
0 2
0 1
0
0
e
+ + + +
+ + +
=
1...,3,4,5,6,7,8,9,10,11,12 14,15,16,17,18,19,20,21,22,23,...43