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Repellency and Toxicity of Conophthorus Ponderosae Hopkins (Coleoptera: Scolytidae) by the Host Monoterpene Myrcene

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Repellency and Toxicity of Conophthorus Ponderosae Hopkins (Coleoptera: Scolytidae) by the Host Monoterpene Myrcene Repellency and Toxicity of Conophthorus ponderosae Hopkins (Coleoptera: Scolytidae) by the Host Monoterpene Myrcene Brian M. Shirley and Stephen Cook Seed orchards produce high-quality seed from selected tree genotypes. In the intermountain west, Conophthorus ponderosae Hopkins (Coleoptera: Scolytidae) is a pest in seed orchards of ponderosa pine, Pinus ponderosa (Laws). The effect of myrcene as a deterrent to cone attack by C. ponderosae in a ponderosa pine seed orchard was examined. Two factors were considered, timing of cone cluster attack and average brood production per cone cluster. There was a delayed Downloaded from https://academic.oup.com/wjaf/article/22/4/241/4717750 by guest on 01 October 2021 attack by C. ponderosae on cones treated with vials of myrcene attached at the base of cone clusters. During both 2003 and 2004, final brood production per cone was not affected significantly by the presence of myrcene. During 2003, brood production was influenced by the timing of attack, with later attacks resulting in fewer brood adults per cone cluster. The toxicity of myrcene to adult C. ponderosae was examined in a laboratory and compared with that of ABSTRACT (ϩ)-␣-pinene, another host-produced monoterpene that acts as a synergist for the male attractant pheromone pityol. Keywords: Conophthorus ponderosae, monoterpene, ponderosa pine, repellency, myrcene, toxicity onifer seed orchards are used to produce high-quality seed nisms of the tree. The girdling also causes seed abortion, but the from selected tree genotypes. Because the orchards are ex- cones of ponderosa pine typically remain attached to the stem. After Cpensive to establish and require intensive management, successfully attacking a cone, female C. ponderosae release a male-at- there is little tolerance for damage to the seed crop. In Idaho, the tractant sex pheromone, pityol (Birgersson et al. 1995). Typically, ponderosa pine cone beetle, Conophthorus ponderosae Hopkins (Co- from 2 to 21 eggs are deposited in the gallery created by the female, leoptera: Scolytidae), can be a pest in both ponderosa pine [Pinus with the average being approximately 7 eggs per cone (Williamson ponderosa (Laws)] and western white pine [Pinus monticola (Doug- et al. 1966, Kinzer et al. 1970). Both parents usually remain in the las)] seed orchards (Wood 1982). C. ponderosae typically causes little cone throughout the brood development period (Wood 1982). No damage in natural pine stands (Schenk and Goyer 1967). For exam- published data are available on actual brood production per cone, ple, a study conducted over a wide area of north Idaho reported which should be lower than the number of eggs laid because of such damage to the cone crop of 9% by Conophthorus monticolae (equals factors as predation, parasitism, and competition among the devel- C. ponderosae) in natural stands of western white pine, P. monticola oping larvae. Douglas (Williamson et al. 1966). However, C. ponderosae was ob- Many scolytid beetles use a combination of chemical and visual served destroying 24.8% of the potential seed crop of ponderosa cues to locate their respective host trees and for intraspecific com- pine over a 4-year period in New Mexico (Kinzer et al. 1970). A munication (Raffa 2001). The chemical cues used by scolytids are closely related species (Conophthorus resinosae Hopkins) with a life often aggregation or sex pheromones that are frequently synergized history similar to that of C. ponderosae can reduce seed production by the presence of tree monoterpenes. Rappaport et al. (2000) de- by up to 90% in cultivated red pine stands (de Groot and Zylstra termined some of the chemical cues used by male C. ponderosae to 1995). Because of this potentially high level of seed mortality, or- locate mates. Male C. ponderosae exhibit a positive response to the chard managers frequently used insecticides such as carbofuran for female-produced sex pheromone pityol (Rappaport et al. 2000). control of C. ponderosae populations (DeBarr et al. 1982, Valenti et Male attraction to pityol is enhanced or diminished in the presence al. 1990). However, because of the impact of carbofuran on nontar- of specific host- and beetle-produced compounds. For example, get organisms and other negative environmental affects, it is no adding 4-allylanisole to a pityol-baited trap reduces trap catch of longer used against Conophthorus spp. male beetles, indicating some form of interruption in the beetle During the spring and early summer, adult C. ponderosae emerge communication system (Rappaport et al. 2000). Adding the tree- from cones attacked the previous year and disperse to locate new produced compound myrcene to pityol-baited traps also reduces host cones (Wood 1982). Female beetles initiate attacks near the trap catches (Shirley 2005). Similar reduction in trap catches has stem at the base of second-year cones. During the attack process, the been observed for other Conophthorus spp. when conophthorin is female girdles the cone, effectively cutting off the defensive mecha- added to traps (Birgersson et al. 1995, Pierce et al. 1995, de Groot et Received April 26, 2006; accepted October 17, 2006. Brian M. Shirley ([email protected]), Department of Forest Resources, College of Natural Resources, University of Idaho, Moscow, ID 83844-1133. Stephen Cook ([email protected]), Department of Forest Resources, College of Natural Resources, University of Idaho, Moscow, Idaho 83844-1133. We thank Sanford Eigenbrode and Elwood Hart for their valuable input and the Bureau of Land Management for the use of their seed orchard. We also appreciate the valuable comments from two anonymous reviewers. The research was supported in part by the Inland Empire Tree Improvement Cooperative and the University of Idaho. Copyright © 2007 by the Society of American Foresters. WEST.J.APPL.FOR. 22(4) 2007 241 al. 1998). However, pityol predominantly attracts male beetles, and examine the impact of the myrcene treatment on mean C. pondero- it is unclear what effect any of these compounds may have on female sae brood production per cone. The C. ponderosae broods in two beetles. Of the three compounds identified as potential repellents to untreated clusters of cones attacked by C. ponderosae were parasit- C. ponderosae, myrcene is produced by host trees, and it has been ized by Cephalonomia conophthori Evans (Hymenoptera: Bethyli- demonstrated to be toxic to other scolytid species (Cook and Hain dae) (Shirley et al. 2004). Because the objective of this study was to 1988, Raffa and Smalley 1995). Many host-produced monoter- determine the effects of myrcene on C. ponderosae brood produc- penes are toxic to insect herbivores, but the level of toxicity is not tion, only cones that had been attacked by C. ponderosae and that consistent across compounds (Cook and Hain 1988, Raffa and were without evidence of predation or parasitism were used in this Smalley 1995, Zou and Cates 1997, Wallin and Raffa 2000). An- analysis. There were 12 treated clusters and 11 untreated clusters other host monoterpene that is also toxic to several species of sco- with attack and no evidence of parasitism. lytids, ␣-pinene (Cook and Hain 1988, Raffa and Smalley 1995), To examine the effect of the attack period on brood production, has also been demonstrated to be a synergist for pityol in attracting the data were reclassified into two periods from the original four, male Conophthorus spp. (Rappaport et al. 2000, de Groot et al. based on the peak time period of beetle capture in pityol-baited traps 1998). within the orchard (Figure 1). The first three sample periods (May The overall objective of this study was to examine the impact of 9–June 23) were combined into a single period, which was the time myrcene on female C. ponderosae when they are attacking host cones period in which the highest daily trap catch of C. ponderosae oc- Downloaded from https://academic.oup.com/wjaf/article/22/4/241/4717750 by guest on 01 October 2021 and on subsequent, within-cone brood production. In addition, the curred. The remaining sample period coincided with the period of toxicity of myrcene to adult C. ponderosae was examined and com- the lowest number of male C. ponderosae trapped, and became the pared with the toxicity of ␣-pinene. new period 2. The reclassification of time periods enabled us to examine the data based on the biology of the beetle. The sample size Materials and Methods for the new analysis had 17 clusters for attack period 1 and 8 clusters In 2003 and 2004, fieldwork was conducted from May to July in for attack period 2 (again, parasitized cones were not included in this the Russell Barr seed orchard in north-central Idaho. The orchard is analysis). Differences between the two periods were compared using owned and operated by the Bureau of Land Management and is Student’s t-test. All statistical analyses were conducted using SAS located on the eastern shore of the Salmon river in Idaho County, Institute (2002). approximately 10 km south of Whitebird, Idaho. The seed orchard During 2004, 20 individual ponderosa pines were selected, and was established in 1977. P. ponderosa is the only species of conifer the control and myrcene treatments were applied to clusters in each present within the orchard. Trees are irrigated and pruned regularly tree and monitored as in 2003. However, during 2004, myrcene to prevent damage to the trees by the frequent prescribed fires ap- treatments were replaced at intervals of approximately 3 week (i.e., plied to the orchard (approximately every 2–3 years). The grasses at the time of cone inspection). The myrcene treatments were re- and forbs growing around the trees are mown frequently (approxi- placed to ensure a continuous presence of myrcene throughout the mately every 2–3 weeks) throughout the growing season. flight season, and the monitoring dates were set earlier to ensure that During 2003, 18 individual ponderosa pine trees with a mini- the myrcene was present prior to any beetles’ flight.
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