Olfactory recognition and behavioural avoidance of angiosperm nonhost volatiles by conifer-inhabiting bark beetles

When searching for suitable hosts in flight, especially in mixed forests, conifer- inhabiting bark beetles will encounter not only suitable host trees and their odours, but also unsuitable hosts and nonhost trees. Rejection of these trees could be based on an imbalance of certain host characteristics and/or a negative response to some nonhost stimuli, such as nonhost volatiles (NHV). 2 Recent electrophysiological and behavioural studies clearly indicate that conifer- inhabiting bark beetles are not only able to recognize, but also to avoid, nonhost habitats or trees by olfactory means. Green leaf volatiles (GLV), especially C6-alcohols, from the leaves (and partly from bark) of nonhost angiosperm trees, may represent nonhost odour signals at the habitat level. Specific bark volatiles such as trans-conophthorin, C8-alcohols, and some aromatic compounds, may indicate nonhosts at the tree species level. Flying bark beetles are also capable of determining whether a possible host is unsuitable by reacting to signals from conspecifics or sympatric heterospecifics that indicate old or colonized host tree individuals. 3 Combined NHV signals in blends showed both redundancy and synergism in their inhibitory effects. The coexistence of redundancy and synergism in nega- tive NHV signals may indicate different functional levels (nonhost habitats, species, and unsuitable hosts) in the host selection process. Combinations of NHV and verbenone significantly reduced the number of mass attacked host trees or logs on several economically important species (e.g. Dendroctonus ponderosae, Ips typographus, and I. sexdentatus). 4 We suggest a semiochemical-diversity hypothesis, based on the inhibition by NHV of bark beetle host-location, which might partly explain the lower outbreak rates of forest insects in mixed forests. This 'semiochemical-diversity hypothesis' would provide new support to the general 'stability-diversity hypothesis'. 5 Natural selection appears to have caused conifer-inhabiting bark beetles to evolve several olfactory mechanisms for finding their hosts and avoiding unsuit- able hosts and nonhost species. NHV and unsuitable host signals have potential for use in protecting trees from attack. The use of these signals may be facilitated by the fact that their combination has an active inhibition radius of several metres in trap test, and by the observation of area effects for several trees near inhibitor soruces in tree protection experiments. Furthermore, incorpora- tion of negative signals (such as NHV and verbenone) and pheromone-based