Fight and rescue or give up and flee? Behavioural responses of different ant species tending the mutualist walnut aphid Panaphis juglandis to native and exotic lady beetles.

Mutualism between ants and honeydew-producing hemipterans is a highly successful evolutionary innovation that attains the status of ecological keystone across many terrestrial ecosystems, involving a multitude of actors through direct or cascading effects. In these relationships, ants often protect their hemipteran partners against their arthropod natural enemies, sometimes interfering with the biological control of pest species. However, the dynamics of these interactions are highly variable based on the specific identity of all the actors involved, and baseline data remain scarce. We performed a field experiment exposing colonies of the walnut aphid Panaphis juglandis attended by five European ant species (Camponotus piceus, Ca. vagus, Crematogaster scutellaris, Dolichoderus quadripunctatus, Lasius emarginatus) to a native and an exotic lady beetle (Adalia bipunctata and Harmonia axyridis), documenting the behavioural interactions between these insects and the performance of ants in the protection of the aphids. Our results reveal a significant behavioural diversity among the ant species involved, with D. quadripunctatus and L. emarginatus being the most aggressive and having the best performance as aphid defenders, and Ca. piceus being least effective and often fleeing away. Cr. scutellaris displayed a rare rescue behaviour attempting to pull away the aphids that the lady beetles grabbed. On the other hand, behavioural responses to A. bipunctata and H. axyridis were similar. Further investigations are needed to understand the eco-ethological implications of these differences, while a better understanding of ant behavioural diversity may help refine biological control strategies.