Verbenone-the universal bark beetle repellent? Its origin, effects, and ecological roles

Bark beetles (Curculionidae: Scolytinae) spend most of their life in tissues of host plants, with several species representing economically relevant pests. Their behaviour is largely guided by complex olfactory cues. The compound verbenone was discovered early in the history of bark beetle pheromone research and is now sometimes referred to as a 'universal bark beetle repellent'. However, some studies aiming to protect trees with verbenone have failed. In fact, most research effort has gone into applied studies, leaving many questions regarding the ecological functions of verbenone for various species unanswered. Here, we review and analyse the scientific literature from more than 50 years. Behavioural responses to verbenone are common among pest bark beetles (< 1% of scolytine species studied so far). Indeed, attraction is inhibited in 38 species from 16 genera, while some secondary species are unaffected or even attracted to verbenone. It is not clear whether the beetles can control the biosynthesis of verbenone; its release may not be an active signal by the beetles, but a passive cue resulting from microorganisms during host colonisation. In this context, we advocate to recognise a bark beetle and its microbiome as an entity ('holobiont'), to better understand temporal release patterns and deduce the specific function of verbenone for a given species. Surprisingly, natural enemies are not commonly attracted by verbenone, but more taxa need to be studied. A better understanding of the ecological functions of verbenone will help to make verbenone-based tools more effective and improve integrated pest management strategies.