No Evidence of Responding Individuals Constraining the Evolution of the Pheromone Signal in the Pine Engraver Ips avulsus.

Chemical signals are important mediators of interactions within forest ecosystems. In insects, pheromone signals mediate intraspecific interactions such as mate location and acceptance. The evolution of pheromones in insects has been mostly studied from a theoretical perspective in the Lepidoptera. With this study, we aimed to broaden our understanding of pheromone communication in bark beetles. We first demonstrated that the enantiomeric ratios of ipsdienol produced by male I. avulsus, showed little variation. Subsequently, with field trapping trials we characterized the influence of the enantiomeric ratio of ipsdienol (pheromone component of I. avulsus) on I. avulsus captures and observed a great amount of variation in the receiver preference function. Most importantly, we demonstrated that responding individuals responded indiscriminately to all the enantiomeric ratios produced by the emitting individuals. These observations are consistent with the asymmetric tracking model which postulates that if the limiting sex is the emitting sex, responding individuals should not discriminate between emitted ratios. Consequently, responding individuals do not constrain the evolution of the signal. Our data suggest that, in I. avulsus, the composition of the aggregation pheromone signal might be more responsive to external selection forces, such as predation and metabolic constraints, as suggested by the asymmetric tracking model.