In recent years it has become apparent that when studying plant-herbivore interactions, it is important to also consider the multitrophic context, i.e. to what extent generalist and specialist enemies act as bodyguards for the plant. I have performed a field experiment with 25 genets, varying in plant defense levels, of meadowsweet (Filipendula ulmaria) and its herbivore, the chrysomelid beetle Galerucella tenella. Each genet was treated with four different density manipulations of G. tenella (5, 15, 30, and 60 larvae per genet). The experiment was set up on an archipelago island with natural levels of a specialist hymenopteran parasitoid Asecodes parviclava and generalist natural enemies. I set up the following hypotheses: 1) larval survival will be higher on less defended plants than on more defended plants, 2) the proportion of parasitoids will be higher on more defended plants, and 3) larval survival will increase with increasing larval densities. The most important findings of the experiment were an overall very low total survival of larvae (G. tenella pupae and A. parviclava mummies), a generally high parasitism by A. parviclava on more defended plants, and that two groups of generalist predators (hunting and web-forming spiders) were more frequent on less defended plants. My study suggests that herbivore survival may not be higher on less defended plants under situations with abundant occurrence of generalist predators, as they are able to trace high-quality foraging sites.

In host-parasitoid systems, there is a continuous coevolutionary arms race where each species imposes a strong selection pressure on the other. The host needs to develop defence strategies in order to escape parasitism and the parasitoid must evolve counter-defence strategies in order to overcome the host’s immune defence and successfully reproduce. This makes host-parasitoid systems excellent model systems for understanding evolutionary processes underlying host race formation and speciation. In order to gain a better understanding of the complexity of host-parasitoid interactions several aspects must be considered, such as search behaviour and host selection in the parasitoid, the development of immune responses in the host and counter-defence strategies in the parasitoid. In this thesis, I investigate interactions and coevolution in a natural host-parasitoid system, consisting of five species of Galerucella leaf beetles and three species of Asecodes parasitoids, by combining behavioural ecology with chemical ecology and immunology. In the studies performed, I found that pheromone production and responses in the beetles are connected to the phylogenetic relatedness between the Galerucella species (Paper I). I found no evidence that Asecodes exploits the adult pheromone to locate host larvae, but observed an ability in the parasitoids to distinguish a better host from a less suitable one based on larval odors (Paper II). The studies also revealed large differences in immune competence between the Galerucella species, which were linked to differences in hemocyte composition in the beetle larvae (Paper III, IV). Further, the results suggest that parasitism success in polyphagous Asecodes is strongly affected by former host species of the parasitoid (Paper IV). In conclusion, the results of this thesis suggest an on-going evolution in both parasitoid virulence and host immune responses in the Asecodes-Galerucella system. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 4: Manuscript.</p>