The erroneous courtship hypothesis: do insects really engage in aerial wars of attrition?

Males of various flying insects perform conspicuous aerial interactions around their mating stations. The broadly accepted interpretation of their aerial interaction is a war of attrition, where two contestants perform costly displays, and the one that reaches its cost threshold earlier gives up. The implicit but important requirement in this model is that some forces that match the intensity of display of the two contestants are necessary, and failure to enforce matching allows foul contestants that delay or stop their display to avoid paying contest costs. In addition, wars of attrition require flying insects to distinguish the sex of flying conspecifics because their aerial interactions begin when intruders fly into the territory. We investigated past research on the behaviour of odonates and butterflies aiming to clarify whether the two prerequisites of wars of attrition are fulfilled: (1) contestants can inflict substantial costs on nondisplaying opponents and (2) contestants can discriminate the sex of flying conspecifics. In odonates, we found an abundance of evidence suggesting that contests involve physical attack and that the ability of sexual discrimination is sufficient. Therefore, wars of attrition may occur in territorial odonates. In butterflies, however, we could not find any evidence that the two prerequisites are filled. The aerial interactions of butterflies are better interpreted as courtship between sexually active males (the erroneous courtship hypothesis). Based on these results, we discuss future directions of research on the aerial contests of flying insects.