Climate change leads to more frequent droughts that may alter multitrophic networks in agroecosystems by changing bottom-up and top-down effects on herbivorous insects. Yet, how bottom-up effects of drought alter tritrophic interactions remains poorly understood. This study investigated two intensities of drought stress in the tritrophic system consisting of sugar beet (Beta vulgaris), an aphid (Aphis fabae), and its parasitoid (Aphidius colemani). We thoroughly investigated each trophic level, examining the performance of plants, pest insects, and parasitoids, as well as the attraction of parasitoids to herbivore-induced plant volatiles (HIPVs). Drought stress negatively affected plant growth but benefited A. fabae, leading to faster development and a higher reproduction rate. Drought-stressed plants also emitted less plant volatiles, which resulted in reduced attraction of A. colemani to aphid-infested plants. Drought indirectly affected parasitoid performance, as evidenced by lower emergence rates and production of fewer females, although mummification rates were higher on drought-stressed plants. Reduced parasitoid attraction and performance on drought-stressed plants may exert lower top-down pressure on aphid populations. Combined with increased aphid performance, this may facilitate aphid outbreaks, which could further weaken drought-stressed plants. Our findings highlight the need to study multiple trophic levels and emphasize the importance of incorporating HIPVs and parasitoid attraction when assessing combined abiotic and biotic stresses in crops.

Foraging animals face the difficult task to find resources in complex environments that contain conflicting information. The presence of a non-suitable resource that provides attractive cues can be expected to confuse foraging animals and to reduce their foraging efficiency. We used the parasitoid Cotesia glomerata to study the effect of non-host-infested plants and associative learning on parasitoid foraging efficiency. Inexperienced C. glomerata did not prefer volatiles emitted from host (Pieris brassicae)-infested plants over volatiles from non-host (Mamestra brassicae)-infested plants and parasitoids that had to pass non-host-infested plants needed eight times longer to reach the host-infested plant compared to parasitoids that had to pass undamaged plants. Contrary to our expectations, oviposition experience on a host-infested leaf decreased foraging efficiency due to more frequent visits of non-host-infested plants. Oviposition experience did not only increase the responsiveness of C. glomerata to the host-infested plants, but also the attraction towards herbivore-induced plant volatiles in general. Experience with non-host-infested leaves on the contrary resulted in a reduced attraction towards non-host-infested plants, but did not increase foraging efficiency. Our study shows that HIPVs emitted by non-host-infested plants can confuse foraging parasitoids and reduce their foraging efficiency when non-host-infested plants are abundant. Our results further suggest that the effect of experience on foraging efficiency in the presence of non-host-infested plants depends on the similarity between the rewarding and the non-rewarding cue as well as on the completeness of information that parasitoids have acquired about the rewarding and non-rewarding cues.

In plant‐associated communities, foraging insects aim to find different resources of interest. Herbivores look for the most nutritious plant tissues, pollinators for sweet nectar and protein‐rich pollen, and carnivores for their preferred prey. Although these foragers look for different resources, they face similar problems: resource cues are often highly variable and need to be detected in complex environments among non‐resources. Search templates—a subset of stimuli that is likely correlated with the occurrence of a particular resource—help foragers across trophic levels to find their resources. However, search‐template‐based foraging can also cause perception errors. Here, we synthesize knowledge on foraging by insects from functional groups belonging to different trophic levels to reveal common problems experienced by foragers and strategies to solve such problems. We focus mostly on volatile‐based foraging because this searching strategy is best studied across functional groups. We argue that search templates are both multimodal (including multiple trait types, e.g. odour and visual) and hierarchical (including multiple foraging steps). Search templates are plastic and updated by experience to match the dynamic foraging environment over time. By comparing insects from different functional groups spanning multiple trophic levels, we have identified important missing gaps on cue use and foraging strategies which should be addressed in the future in order to reduce knowledge asymmetries among functional groups about search‐template‐based foraging. We propose a way to achieve this goal arguing that studies on less well investigated functional groups can advance rapidly by borrowing, testing and adjusting already available hypotheses and theories formulated for other insect groups. Knowledge of search‐template‐based foraging across functional groups will give new insights into the evolution of foraging behaviour in complex ecological communities, help predict ecological consequences of large‐scale human‐made disturbances and help optimize insect‐delivered ecosystem services (pollination and biological control) in cropping systems. Read the free Plain Language Summary for this article on the Journal blog.