Response of the mountain pine beetle, Dendroctonusponderosae Hopkins (Coleoptera: Scolytidae), to five semiochemicals in British Columbia lodgepole pine forests

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Abstract
Five principal semiochemicals were field tested in multiple funnel traps for behavioral activity against Dendroctonusponderosae infesting lodgepole pine, Pinuscontorta var. latifolia Engelmann, in British Columbia. The basic experimental design was to test each compound by adding it in varying concentration or enantiomeric composition to a blend of other semiochemicals. (−)-trans-Verbenol was attractive to both sexes. (±)-exo-Brevicomin and (±)-frontalin (in one of two experiments) were attractive with other semiochemicals to females at a release rate of 0.05 mg/24 h. At higher release rates (±)-exo-brevicomin was inhibitory to males, and frontalin was inhibitory to both sexes. Thus both serve as multifunctional pheromones. For neither exo-brevicomin nor frontalin were the separate enantiomers attractive at the low release rate, suggesting that they had an additive effect. However, at the high release rate both enantiomers mimicked the inhibitory effect of the racemates. Verbenone acted as an antiaggregation pheromone for both sexes. Increasing the release rate of myrcene from 18 to 150 mg/24 h to approximate the release rate from a newly attacked tree had the effect of doubling the catch of responding beetles. A conceptual model is proposed for the sequential interaction of these semiochemicals in the mass attack of a tree. Pioneer females release trans-verbenol, which acts in combination with myrcene from the host tree to attract mainly males. The responding males release exo-brevicomin and later frontalin, which in combination with trans-verbenol and myrcene attract mainly females. Meanwhile autoxidation of α-pinene in the host resin results first in the production of predominately trans-verbenol, which supplements that produced by the beetles. Later, autoxidation of α-pinene and microbial conversion of cis- and trans-verbenol result in the production of the antiaggregation pheromone verbenone. This compound, in combination with large amounts of exo-brevicomin and frontalin as the tree becomes fully occupied, results in the close-range redirection of responding beetles toward nearby trees.