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The Capacity of Conophthorin to Enhance the Attraction of Two Xylosandrus Species (Coleoptera: Curculionidae: Scolytinae) To

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The Capacity of Conophthorin to Enhance the Attraction of Two Xylosandrus Species (Coleoptera: Curculionidae: Scolytinae) To Agricultural and Forest Entomology (2013), 15, 391–397 DOI: 10.1111/afe.12026 The capacity of conophthorin to enhance the attraction of two Xylosandrus species (Coleoptera: Curculionidae: Scolytinae) to ethanol and the efficacy of verbenone as a deterrent ∗ ∗ Nicole R. VanDerLaan and Matthew D. Ginzel † ∗Department of Forestry and Natural Resources, Purdue University, 715 W. State Street, West Lafayette, IN 47907, U.S.A. and †Department of Entomology, Purdue University, 901 W. State Street, West Lafayette, IN 47907, U.S.A. Abstract 1 Exotic invasive ambrosia beetles are among the most economically important pests of nursery stock and forest systems in the U.S.A. 2 Conophthorin, a common bark volatile of deciduous trees, acts as an attractant for the exotic black stem borer, Xylosandrus germanus (Coleoptera: Curculionidae: Scolytinae). Nevertheless, the extent to which a congener, Xylosandrus crassiusculus and other ambrosia beetles are attracted to conophthorin remains unclear. It is also unknown whether conophthorin enhances the attraction of these Xylosandrus beetles to traps baited with ethanol. 3 In the present study, we evaluated the extent to which conophthorin enhances the attraction of X. crassiusculus and X. germanus to ethanol-baited traps. We also tested the capacity of verbenone, an anti-aggregation pheromone component of several coniferophagous bark beetles, to deter both species. 4MoreX. crassiusculus were captured in traps baited with both conophthorin and ethanol than in those containing either compound alone, suggesting that conophthorin enhances the response of X. crassiusculus to ethanol. This combination was also attractive to the checkered beetles Madoniella dislocatus and Pyticeroides laticornis (Coleoptera: Cleridae). 5 Verbenone deterred both X. germanus and X. crassiusculus, suggesting that the use of conophthorin and ethanol as an attractant and verbenone as a deterrent can be incorporated into an effective integrated pest management programme. Keywords Conophthorin, ethanol, verbenone, Xylosandrus crassiusculus, Xylosandrus germanus. Introduction individuals are necessary to establish a growing population (Kirkendall & Ødegaard, 2007; Biedermann et al., 2009). Exotic ambrosia beetles are serious wood-boring pests of land- Currently, practical and technological limitations hamper the scape, nursery, orchard and forest trees (Oliver & Mannion, detection and subsequent management of small, newly-founded 2001; Adkins et al., 2010) and among the most commonly populations of exotic beetles (Mehta et al., 2007; Bogich intercepted insects at ports-of-entry in the U.S.A. (Haack, 2001; Kuhnholz¨ et al., 2001; Rabaglia et al., 2008; Marini et al., et al., 2008). Once established, ambrosia beetles are difficult to 2011). As international trade and travel increases, the number control because the majority of their lifecycle is spent beneath of introductions of exotic ambrosia beetles continues to rise at the bark of trees, where they are physically protected from an alarming rate (Haack, 2001). These pests cause substantial sprayed insecticides (Reding et al., 2010). Indeed, conventional environmental damage and economic loss (CABI, 2012). Xyle- contact insecticides provide control only if applied soon before borine beetles also have a symbiotic relationship with ambrosia the adult flight period and may not kill all colonizing adults fungi and, as a result of their haplodiploid lifecycle, only a few (Mizell & Riddle, 2004; Frank & Sadof, 2011). Xylosandrus germanus (Blandford) and Xylosandrus cras- Correspondence: Matthew D. Ginzel. Tel.: (765) 494-9369; fax: siusculus (Motschulsky) (Coleoptera: Curculionidae: Scolyti- (765) 494-0535; e-mail: [email protected] nae), two exotic ambrosia beetles of Asian origin, were first © 2013 The Royal Entomological Society 392 N. R. VanDerLaan and M. D. Ginzel detected in the U.S.A. via infested material in 1932 and 1974, & Miller, 2010). When combined with ethanol, verbenone respectively (Felt, 1932; Hoffman, 1941; Wood, 1982; Weber & also decreases trap catches of X. crassiusculus, Xylosandrus McPherson, 1984; Cote, 2008; CABI, 2012). Both species are compactus (Eichhoff) and Xyleborinus saxesenii (Ratzeburg) considered high-risk, invasive pests and are now established in koa (Acacia koa A. Gray) stands (Burbano et al., 2012); in much of the eastern U.S.A. (CABI, 2012). These beetles however, the bioactivity of verbenone alone has yet to be have a broad host range of over 200 species, including valu- assessed. able hardwoods, and can proliferate rapidly in agricultural and Furthermore, X. germanus appears to be attracted to trap forested areas (Wood, 1982; Weber & McPherson, 1984; Oliver trees baited with conophthorin (Dodds & Miller, 2010), a & Mannion, 2001; Adkins et al., 2010; Reding et al., 2010). bark volatile produced by several deciduous trees (Byers Xylosandrus crassiusculus exclusively attacks deciduous trees, et al., 1998; Huber et al., 1999). Conophthorin is also a whereas X. germanus also attacks some Abies, Picea and Pinus pheromone component of some bark beetles within the genera spp. (Cote, 2008; CABI, 2012). Both species overwinter as Conophthorus, Leperisinus and Pityophthorus (Francke et al., adults and emerge in high densities in the spring. These species 1979; Dallara et al., 1995; Birgersson et al., 1995). The addition have two adult flight periods per year in Indiana. Overwin- of conophthorin to flight intercept traps baited with ethanol and tering adults emerge in the spring and the peak flight period Linoprax® (a synthetic pheromone lure for coniferophagous of the subsequent generation occurs in late summer, although bark beetles) reduced the number of X. germanus captured all life stages can be found within galleries during the grow- (Kohnle et al., 1992). Nevertheless, the influence of ethanol ing season (Cote, 2008). Males are flightless and spend their on the attraction of Xylosandrus beetles to conophthorin has entire lifecycle within the host tree (Peer & Taborsky, 2005), yet to be empirically tested. whereas dispersing females travel 100 m or more to locate a In the present study, we tested the hypothesis that conoph- suitable host (Peer & Taborsky, 2005). Colonizing females vec- thorin serves as an attractant to the exotic ambrosia beetles tor pathogenic fungi that disrupt the flow of water and nutrients X. germanus and X. crassiusculus and also acts to enhance their within the xylem, potentially killing the host tree (Cote, 2008; attraction to ethonal-baited traps. Additionally, we tested the Adkins et al., 2010). Both adults and larvae feed on the fun- efficacy of verbenone alone to deter these beetles. The ratio- gal mycelium and not the host tree (Cote, 2008; Adkins et al., nale for such an investigation is that an improved lure for exotic 2010). These fungi are of concern to growers of lumber and ambrosia beetles may lead to early detection of these pests and veneer because they can spread throughout the tree, stain the trigger timely management responses to reduce the extent of wood blue and make it less marketable (Cote, 2008). No aggre- damage and prevent losses. gation or sex pheromone has been identified for any xyleborine species and little is known of their chemical ecology (Hulcr & Cognato, 2010). In fact, ethanol is the only semiochemical Materials and methods lure currently used by United States Department of Agriculture (USDA) Animal and Plant Health Inspection Service (APHIS)- We conducted a field experiment aiming to evaluate the attrac- Plant Protection and Quarantine (PPQ) and the U.S. Forest tion of X. germanus and X. crassiusculus to conophthorin and Service for the early detection of these and other invasive ethanol alone and determine the extent to which conophthorin ambrosia beetles at ports-of-entry (Rabaglia et al., 2008). enhances the response of these beetles to ethanol-baited traps. Ethanol, a common indicator of plant stress (Kimmerer & We also determined the extent to which these beetles were Kozlowski, 1982; Miller & Rabaglia, 2009; Ranger et al., inhibited by verbenone. The experiment was conducted during 2010), is a general attractant for many ambrosia beetles (Mont- the flight periods of both species in the summer of 2011 and gomery & Wargo, 1982; Miller & Rabaglia, 2009) and is 2012 and during the spring in 2012 (for dates and locations, see commonly used to enhance the attraction of woodborers to Table 1). During each flight period, three transects comprised semiochemical lures and trap trees (Montgomery & Wargo, of five traps (spaced 10 m apart) were positioned perpendicular 1982; Miller & Rabaglia, 2009; Ranger et al. 2013a). Ethanol to the direction of prevailing winds at one or two locations in enhances the attraction of both sexes of the ambrosia beetle Indiana with an active ambrosia beetle population (Table 1): Gnathotrichus retusus (Lec.) to traps baited with the aggre- Martell Forest (approximately 172 ha) and the Purdue Wildlife gation pheromone, (S )-(+)-sulcatol (Borden et al., 1980). The Area (approximately 64 ha) both located in Tippecanoe Co.; combination of ethanol and turpentine increases the response and Black Rock Barrens (NICHES Land Trust; approximately of the ambrosia beetle Xyleborus pubescens Zimmerman com- 40 ha) located in Warren Co. Each transect was considered a pared with the individual lures alone (Phillips et al., 1988). replicate; thus, there were nine replicates of each treatment. Additionally, ethanol enhances the attraction of other scolytids, All sites were dominated by oak,
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