Bottom-up effects on a plant-endophage-parasitoid system: the role of flower-head size and chemistry

The effects of water and nutrient addition on a trophic chain were studied in a plant-endophage-parasitoid system comprised of insects associated with flower heads of Chromolaena squalida (Asteraceae). Nine species of endophages associated with C. squalida flower heads were found, belonging to two families of Diptera - Tephritidae (Cecidochares sp1, Cecidochares sp2, Xanthaciura biocellata, X. chrysura, X. sp. and Neomyopites sp.) and Agromyzidae (Melanagromyza sp.), and two families of Coleoptera - Apionidae (Apion sp.) and Anthicidae (Anthicidae sp.). A factorial field experiment with water and nutrient addition showed that resource availability can affect the developmental process of flower heads. Fertilization increased flower-head diameter and nitrogen and alkaloid concentrations. Although nutrient availability affected the size and chemistry of flower heads, endophage species did not respond consistently to the experimental treatments. This is contrary to other studies where endophages showed preference for larger flower heads. Thus, the plant vigour hypothesis was not corroborated for our study system. Our results also showed that coupled responses of plants to resource availability (i.e. tissue nutritional quality and investment in growth of the structure that serves as shelter for endophages) can represent distinct kinds of indirect interactions with opposing effects on the herbivore-parasitoid interaction.