Paul E. Tilden

Western pine beetle: field response to its sex pheromone and a synergistic host terpene, myrcene.

In the field, both sexes of the western pine beetle, Dendroctonus brevicomis, are attracted by the female-produced bicyclic ketal exo-brevicomin; this response is enhanced by myrcene (a constituent of the beetles host, ponderosapine), which is not an attractant by itself. This synergism may be part of the phenomenon of the mass attack on its host. Temnochila virescens chlorodia, one of the principal insectan predators of this beetle, is attracted by exo-brevicomin alone.

Field responses of the western pine beetle1 and one of its predators to host- and beetle-produced compounds

The responses of the western pine beetle (Dendroctonus brev-icomis LeConte) andTemnochila chlorodia (Mannerheim) to candidate attractants—exo- andendo-brevicomm, frontalin,trans-verbenol, ver-benone, and ponderosa pine turpentine and its major monoterpene components—were quantified by counts of beetles on traps baited with the various attractants, singly and in combinations released simultaneously. Combinations ofexo-brevicomin and frontalin plus a monoterpene or turpentine were the most attractive toD. brevicomis. The responses to these attractant combinations were reduced when verbenone plustrans-verbenol were present. All single compounds and binary mixtures, exceptexo-brevicomin plus frontalin, were much less attractive.exo-Brevicomin was most attractive toT. chlorodia, and this response appeared to decrease when verbenone plustrans-verbenol were present.

Effects of verbenone andtrans-verbenol on the response ofDendroctonus brevicomis to natural and synthetic attractant in the field

Verbenone andtrans-verbenol were investigated as candidate interruptants for use as tree protectants. Verbenone andtrans-verbenol, pheromones released byDendroctonus brevicomis during host colonization, reduced the trap catch ofD. brevicomis near sources of the attractant composed ofexo-brevicomm, frontalin, and myrcene. Catch reduction at some trap positions was greater at a high release rate than at a low release rate oftrans-verbenol alone and of the combination of verbenone plustrans-verbenol. Verbenone also reduced catches at traps baited with attractive bolts from trees under attack byD. brevicomis. Attempts to use verbenone to protect living trees fromD. brevicomis attack were inconclusive.

Trapping the western pine beetle at and near a source of synthetic attractive pheromone: Effects of trap size and position

Western pine beetles were caught on unbaited sticky traps placed near a source ofexo-brevicomin, frontalin, and myrcene. Size of trap, distance and direction from the source of attractant, and height from the ground were varied. Significant differences in trap catch were observed in relation to each of the variables. Traps close to the source of attractant caught more beetles than traps farther from the source. Traps downwind of the source of attractant caught more beetles than did upwind traps. More males than females were trapped close to the source of attractant.

TrappingDendroctonus brevicomis Changes in attractant release rate, dispersion of attractant, and silhouette

A sticky trap with 3 m2 surface area was modified by changes in attractant release rate, vertical dispersion of the attractant, and addition of a tree trunk silhouette to the trap axis. As attractant release rate increased, the number ofDendroctonus brevicomis caught at the source of attractant and at 1.5 and 5.2 m above ground on two vertical silhouettes 4.5 m away increased. In one experiment, more beetles were caught at a dispersed source of attractant than at a point source. Fewer beetles were caught at the lower traps on the two outlying silhouettes when a silhouette was at the source, than when no silhouette was at the source. As attractant release rate increased, the catch of a predator,Temnochila chlorodia, increased at the source.

Effect of verbenone on response ofDendroctonus brevicomis toexo-Brevicomin, frontalin, and myrcene.

Abstractexo-Brevicomin (E), frontalin (F), and myrcene (M) were released at two rates 10-fold apart with verbenone at four rates 10-fold apart, and without verbenone in plots with one trap on a vertical cylinder at the pheromone source and one trap on each of four cylinders 5 m away. Catch of the western pine beetleDendroctonus brevicomis decreased with increasing levels of verbenone at both release rates of EFM, but not all differences in catch were statistically significant. Significantly more beetles were caught at the high rate of EFM than at the low rate, combining all rates of verbenone. The percent of total beetles caught at the center trap tended to decrease with increasing rates of verbenone, but the only statistically significant differences were at the low rate of EFM.

Interruption of response ofDendroctonus brevicomis to its attractive pheromone by components of the pheromone.

The catch of the western pine beetle,Dendroctonus brevicomis, at an attractive source of racemicexo-brevicomin, racemic frontalin, and myrcene was reduced by surrounding the source with a grid of 48 stations releasing all three compounds together, orexo-brevicomin alone or myrcene alone. Each compound was released at the rate of 2 mg/24 hr/station. The catch at an attractive bolt cut from a tree being colonized byD. brevicomis was not reduced byexo-brevicomin, but was reduced by the combination ofexo-brevicomin, frontalin, and myrcene in one of two tests. When a transect of traps was placed across a 0.81-hectare plot at six of the 48 stations releasing all three compounds, more beetles were caught at outer than at inner traps. More beetles were caught at unbaited traps on trees in a plot when the three compounds were released than when onlyexo-brevicomin or no compounds were released. A few trees were attacked byD. brevicomis in some of the plots. The antiattractant verbenone released from 48 stations at the rate of 4 mg/24 hr/station did not reduce the catch at an attractive tree bolt.

Field response ofDendroctonus brevicomis toexo-brevicomin, frontalin, and myrcene released at two proportions and three levels

Racemicezo-brevicomin, racemic frontalin, and myrcene were released at two proportions (5∶1∶400 and 1∶1∶1), each at three levels (1 ×, 10 ×, and 100 ×) in a ponderosa pine forest in central California. The 5∶1∶400 mix was based on an estimate of the relative amounts released from a ponderosa pine under attack by the western pine beetle,Dendroctonus brevicomis. MoreD. brevicomis were trapped at a source of the three compounds released at 5∶1∶400 than were trapped at a source released at 1∶1∶1, at all three levels, but this difference was statistically significant only at the 1 × and 10 × levels. Sex ratio of trapped beetles and distribution of catch at the source of attractant and 5 m away apparently did not differ between relative release rates.

Behavior of the western pine beetle during host colonization

After living ponderosa pines were baited with either female-infested bolts or synthetic pheromones,Dendroctonus brevicomis were caught on sticky screens throughout trapping periods of 15–46 days; however, large numbers of beetles were trapped during only a small portion (5–10 days) of these trapping periods. The most attractive portions of trees attacked contained 3–6 beetles dm2, in galleries ca. 2 cm long. Catch increased following addition of males to female-infested bolts, supporting the hypothesis that male-produced frontalin is an attractive pheromone of the western pine beetle. Catch at bolts removed from trees under attack was strongly dependent upon levels of boring activity. We found no evidence of interruption of the response to attractants during host colonization.

Interruption of response ofDendroctonus brevicomis to attractive pheromone by release of pheromone at several rates and spacings

The number of western pine beetles,Dendroctonus brevicomis, trapped at the center of three 90 × 90-m plots was reduced during a 16-hr period after surrounding an attractive source ofexo-brevicomin, frontalin, and myrcene with 16, 48, or 168 sources of the same attractant. Compounds were released from the center of the plot at 1 mg/16 hr/compound, and from the surrounding sources at 4–280 mg/16 hr/compound. About half of these amounts was released between 1530 hr and 1930 hr, the period of peak beetle flight. No treatment differences were apparent in reduction of catch within the range of release rates and spacings tested. When compounds were released continuously from 168 stations for 17 days, catch at the center of a plot was generally lower than catch before or after this period, but fluctuated daily. More beetles were caught on traps hung on ponderosa pines within a plot and the number of these traps catching beetles was greater when compounds were released from all stations than from only the center station. Beetles caught on traps were attracted into the plots from the surrounding forest, but appeared to be dispersed within the plot when compounds were released from many stations.D. brevicomis attacked at least 91 trees in the plots, of which 25 were killed during two summers.