Abstract
The existence of different male morphologies within a single species is associated with alternative mating tactics across different animal groups, offering textbook examples of evolutionary biology. The European stag beetle Lucanus cervus is a flagship species which has long fascinated naturalists for its charismatic appearance and behavior, with males possessing large mandibles used in ritualized ground fights for females. Males are polymorphic, with larger males possessing disproportionately larger mandibles and being better fighters, while smaller males require less food to develop and may have better chances to escape predation. We discovered an alternative mating tactic that opens a new interpretation on the evolution of the smaller males, based on aggregations of flying males competing to catch flying females in mid-air. Flight aggregations occur earlier in the season and in sites with sparser canopy cover as compared to ground fight aggregations, but most importantly, smaller males tend to prevail due to their greater agility in flight. Competition between males is crucial for reproductive success in this species, and alternative mating tactics involving flight were unknown in beetles, advocating for further research on the ecology and evolution of the iconic male polymorphism of stag beetles.