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Recent studies revealed that parasitoids
with a broader host range seem to use more
general cues than more specialized para-
sitoids (Vet et al.,1993; Hedlund et al., 1996;
Röse et al.,1998; Bruni et al., 2000). However,
in other studies, no or only minor differ-
ences were found (Geervliet et al., 1996;
Cortesero et al., 1997). As yet, not enough
data exist to make a meaningful comparison
of the relative learning ability of specialist
and generalist parasitoids. In addition, most
present data, with the exception of results
from Cotesia species, include work on only
distantly related species. Any comparison
would risk erroneously attributing differ-
ences in learning to differences in diet
breadth when in fact they are due to other
factors, for example, differences in phy-
logeny (Papaj and Prokopy, 1989). We sug-
gest that the lack of consensus with regard
to the learning abilities of specialists and
generalists may be due in part to the failure
to test enough stimuli over the possible
range of response levels.


Concluding Remarks

The effect of experience on the mean and
variability (and thus predictability) of behav-
ioural responses has interesting implications
for the use of parasitoids in biological con-
trol. An improved predictability of natural-
enemy behaviour will stimulate application


of biological control (Lewis et al., 1990).
Unpredictable behaviour can hamper mass
rearing and the development of reliable intro-
duction schemes, lead to disinterest in the
biological-control method and result in the
release of exorbitantly high numbers of ani-
mals of poor quality, leading to high control
costs. The postrelease migration behaviour of
parasitoids away from the target area is con-
sidered to be a special problem (e.g. Ridgway
et al., 1981; Keller et al., 1985). Increasing the
mean and reducing the variability of the
response to target stimuli through experience
could considerably alleviate this problem.
Although our model implies that all learn-
ing in parasitoids can be reduced to simple
associative processes, where reinforcement
increases the response to some other stimu-
lus, it also includes other effects of experience
where an obvious reinforcement is lacking
(e.g. sensitization and habituation).
A behavioural repertoire is a complex
process, influenced by genes, environment,
physiology and experience. Being aware of
this complexity, we merely present a tool to
simplify and clarify the effect of experience
on behavioural responses and variability in
those responses. The simplicity of the model
enables us to formulate clear and testable
hypotheses bearing on the desired or
unavoidable manipulation of natural ene-
mies, interspecific differences in behavioural
plasticity and learning mechanisms.
The information presented in this chapter

Parasitoid Foraging Behaviour 35

Specialist
Generalist

Stimulus rank

Response potential

Fig. 3.4.Differences in response-potential curves between specialist and generalist parasitoid species. See
text for additional explanation.

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