Foraging behaviour of parasitoids in multi-herbivore communities

Foraging behaviour of parasitoids in multi-herbivore communities Parasitic wasps, or parasitoids, use herbivore-induced plant volatiles and infochemicals produced directly by the herbivore to locate their herbivorous hosts. This process could be interrupted by the presence of herbivores that are not suitable for the development of parasitoid offspring. These non-host herbivores could affect the behaviour of parasitoids both when parasitoids are foraging for host-infested plants by using plant volatiles and when parasitoids are foraging for hosts on the plant by using herbivore infochemicals. The aim of this thesis was to study the impact of non-host presence on the parasitoid-host-food plant complex of the parasitoid Cotesia glomerata with its host caterpillar Pieris brassicae and a monoculture of the cultivated plant Brassica oleracea. To study the influence of non-hosts on the plant-volatile-based searching behaviour of the parasitoid, a wind-tunnel set-up in the laboratory was used. In this set-up, the parasitoids were given a choice between two plants or between the leaves of one plant. The plant/leaf on which the parasitoid landed was considered the preferred plant/leaf. A second laboratory set-up was used to study the influence of non-host herbivores on the host-infochemical-based searching behaviour of the parasitoid. In this on-plant experiment, the behaviour of the parasitoid was observed after landing on the plant. The influence of non-hosts on the combination of plant-volatile-based and host-infochemical-based searching, i.e. the total foraging efficiency of the parasitoid, was investigated using an outdoor tent set-up in an agricultural field. In this semi-field experiment, parasitoids were allowed to parasitize their hosts in a non-host environment for one to three days. This thesis firstly shows that the feeding guild of non-host herbivores influenced the foraging behaviour of C. glomerata. Leaf-chewing non-hosts negatively impacted the plant-volatile-based searching behaviour of the parasitoid, whereas phloem-feeding non-hosts positively impacted the host-infochemical-based searching. The resulting host-finding efficiency was in general positively affected by phloem-feeding non-hosts. Secondly, the position of host and non-host herbivores on the plant affected the plant-volatile-based and the host-infochemical-based foraging behaviour of the parasitoid, but not the host-finding efficiency. An unnatural distribution of herbivores over the plant (host feeding on old leaf, non-host feeding on young leaf), negatively affected the choice of the parasitoid for a leaf to land on, i.e. the parasitoid more often landed on the non-host infested leaf. Combined feeding by the host and non-host on one leaf positively affected the number of hosts parasitized on that plant compared with the number of hosts parasitized when herbivores were separated. However, the parasitoid was able to compensate for these effects as a result of which the foraging efficiency was unaltered. Thirdly, the density of non-hosts did influence the plant-volatile-based searching of the parasitoid. A high non-host density negatively affected parasitoid preference for host-infested plants. However, the host-infochemical-based foraging and the total foraging efficiency remained unaffected. Fourthly, rather than the species diversity, the species identity of non-host herbivores had an influence on parasitoid host-infochemical-based searching. One of the tested non-host species negatively affected the behaviour of the parasitoid when searching on the plant. However, neither non-host species identity nor diversity affected plant-volatile-based searching of the parasitoid. Fifthly, this thesis investigated if a parasitoid could learn to associate non-host cues with the presence of hosts and if the parasitoid changed the parasitization preference accordingly. After receiving a learning experience, the parasitoid showed an altered landing preference for infested plants according to the learned cues. However, in an outdoor tent set-up, the parasitoid did not show an altered parasitization preference. The results of this thesis show that non-host herbivore traits can affect the different phases of parasitoid foraging either positively, negatively or neutrally. The non-host effect on the total foraging efficiency is not necessarily a result of the sum of the effects on the first and the second foraging phase. In fact, the results of two out of three outdoor-tent experiments that investigated the foraging efficiency of the parasitoid showed no non-host effect, while the separate foraging phases were affected by non-host presence. It is concluded that the foraging efficiency of the parasitoid C. glomerata when searching for its host P. brassicae is not strongly affected by non-host herbivore presence. The use of herbivore-induced plant volatiles by C. glomerata during this foraging process is not interrupted by non-host herbivores. It is advised to consider all phases of the foraging process in studies of parasitoid foraging behaviour, preferably in one experiment that covers the whole searching process. Altogether, this thesis gives a clear and comprehensive overview of the impact of non-host presence on a parasitoid-host-food plant complex and it thereby contributes to the fundamental knowledge of insect foraging in a multi-herbivore context.