Journal of Economic Entomology, 109(5), 2016, 2145–2150 doi: 10.1093/jee/tow197 Advance Access Publication Date: 27 August 2016 Forest Entomology Research article

Influence of Fermenting Bait and Vertical Position of Traps on Attraction of Cerambycid to Pheromone Lures

Joseph C. H. Wong and Lawrence M. Hanks1

Department of Entomology, University of Illinois at Urbana-Champaign, Urbana, IL 61801 ([email protected]; hanks@life. illinois.edu), and 1Corresponding author, e-mail: [email protected]

Received 6 June 2016; Accepted 10 August 2016

Abstract Because larvae of cerambycid beetles feed within woody plants, they are difficult to detect, and are readily transported in lumber and other wooden products. As a result, increasing numbers of exotic cerambycid spe- Downloaded from cies are being introduced into new regions of the world through international commerce, and many of these species pose a threat to woody plants in natural and managed forests. There is a great need for effective meth- ods for detecting exotic and potentially invasive cerambycid species, and for monitoring native species for con- servation purposes. Here, we describe a field experiment in east-central Illinois which tested whether attraction

of beetles to a blend of synthesized cerambycid pheromones would be enhanced by volatiles from fermenting http://jee.oxfordjournals.org/ bait composed of crushed fruit, sugars, yeast, and wood chips. A second experiment tested the same treat- ments, but also assessed how trap catch was influenced by the vertical position of traps within forests (under- story versus within the canopy). During the two experiments, 885 cerambycid beetles of 37 species were caught, with colonus (F.) (subfamily Cerambycinae) being the most numerous (52% of total). Adults of several cerambycid species were significantly attracted by the pheromone blend, but the fermenting bait significantly enhanced attraction only for X. colonus and Graphisurus fasciatus (Degeer) (subfamily ). Traps in the forest understory caught the greatest number of X. colonus and G. fasciatus, whereas more adults of the cerambycine Neoclytus mucronatus mucronatus (F.) were caught in the forest canopy rather by guest on November 28, 2016 than the understory.

Key words: Cerambycidae, wood-borer, trap, monitoring, semiochemical

Wood-boring beetles of the Cerambycidae play important roles in Miller and Crowe 2009, Allison et al. 2014, Alvarez et al. 2014, forest ecosystems, especially as nutrient recyclers which initiate the Dodds et al. 2015), which have been rendered even more efficient in breakdown of dead woody plants (Linsley 1959). Because the larvae catching and retaining beetles by coating their surfaces with lubri- feed within woody plants, they are difficult to detect, and are readily cants (e.g., Graham et al. 2010, Allison et al. 2011, Alvarez et al. transported in lumber and other wooden products. As a result, in- 2014). Furthermore, several studies have shown that the vertical po- creasing numbers of exotic cerambycid species are being introduced sition of traps within forests can be an important factor for compre- into new regions of the world through international commerce, and hensive assessment of cerambycid communities, because adults of many of these species pose a threat to woody plants in natural and some species appear to be localized within specific vertical strata of managed forests (Nowak et al. 2001, Brockerhoff et al. 2006). forests (e.g., Vance et al. 2003, Wermelinger et al. 2007, Graham Effective methods for detecting exotic and potentially invasive et al. 2012). cerambycid species, and for monitoring native species for conserva- Many species of cerambycids are known to be attracted to vola- tion purposes (e.g., Ray et al. 2014) would be valuable tools for for- tiles emitted by the host plants of their larvae (reviewed by Hanks est managers and regulatory personnel. and Wang 2016). Synthetic reconstructions of host volatiles en- Over the past decade, there has been a great deal of progress in hanced attraction of some cerambycid species to their pheromones, identifying pheromones of cerambycid beetles for use as lures in increasing the species diversity and number of beetles lured to traps traps (reviewed by Millar and Hanks 2016). Blends of the synthe- (e.g., Sweeney et al. 2004, Silk et al. 2007, Hanks et al. 2012). sized pheromones of several species have been developed as Scattered reports, extending over many decades, suggest that fer- multispecies lures that are broadly attractive to multiple species menting baits, consisting of aqueous blends of sugary substances (e.g., Hanks et al. 2012). In parallel with this pheromone work, and alcoholic beverages which are allowed to ferment, produce vola- more effective traps have been developed (e.g., Bouget et al. 2008, tiles that attract some species of wood-boring (Champlain

VC The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: [email protected] 2145 2146 Journal of Economic Entomology, 2016, Vol. 109, No. 5 and Knull 1932, Linsley 1959, Galford 1980). These baits constitute consisted of 200 ml of the resulting liquid in polyethylene sachets another possible source of attractants for some wood-boring insects, (10 by 15 cm, 0.05 mm thick, Bagette model 14772, Cousin Corp.) including cerambycids. with four holes (5.5 mm) punched below the seal to increase re- In an earlier study, Schmeelk et al. (2016) compared attraction lease rate of volatiles. of cerambycid beetles to traps baited with either a blend of synthe- Experiment 1 compared attraction of beetles to the pheromone sized pheromones or a fermenting bait in forests of east-central blend, fermenting bait, and the two lure types combined using traps Illinois. Traps were positioned at three heights within forests. They deployed 10 m apart in two separate linear transects (15 m apart) found that pheromone-baited traps caught greater numbers of bee- in the forest understory. Traps were hung from inverted L-shaped tles than did traps baited with the fermenting bait. Moreover, some frames constructed of polyvinyl chloride (PVC) pipe (bottom of pan- species were caught in greater numbers by traps in the understory, els 1 m above the ground), with lures hung from the central open- whereas other species were more numerous in traps in the forest can- ing. Treatments were as follows: 1) pheromone blend þ fermenting opy. Here, we describe follow-up experiments which tested whether bait (separate lures), 2) pheromone blend, 3) fermenting bait, and 4) fermentation volatiles enhanced attraction to a pheromone blend, solvent control. The experiment was conducted from 23 June to 11 conducted at one of the same field sites used by Schmeelk et al. September 2015, with treatments randomly assigned to traps (2016), and which also tested the effect of trap height by positioning within transects on the day of setup. Traps were serviced at intervals traps either in the forest understory or at midcanopy height. of 2 d, at which time treatments were rotated one position along transects to control for positional effects. Experiment 2 also tested the attraction of beetles to the phero- Materials and Methods mone blend, fermenting bait, and the combination, but included a Downloaded from trap height treatment. Panel traps of the same type were deployed Sources of Chemicals in pairs such that one trap was hung from a PVC frame in the un- Racemic 3-hydroxyhexan-2-one, 2-(undecyloxy)ethanol (termed derstory (as already described), and above it a trap was suspended monochamol), (E)-6,10-dimethylundeca-5,9-dien-2-ol (fuscumol), from the branch of a sugar maple (Acer saccharum Marsh.) in the and (E)-6,10-dimethylundeca-5,9-dien-2-yl acetate (fuscumol ace- midcanopy (12 m above the ground). Traps in the midcanopy tate) were purchased from Bedoukian Research (Danbury, CT), and were positioned by first throwing a line over the branch with a http://jee.oxfordjournals.org/ racemic 2-methylbutan-1-ol from Sigma-Aldrich (St. Louis, MO). large slingshot (Big Shot, Sherrill Inc., Greensboro, NC), then Syn-2,3-hexanediol was synthesized as described in Lacey et al. hauling the trap up to that height, following the methods of (2004). Hughes et al. (2014). Sugar maples were chosen for this treatment because the midcanopy branches tended to be more nearly hori- Field Bioassays zontal than those of other species of trees, and so were more con- Research was conducted at Forest Glen Preserve (Vermilion venient for setting up traps. In addition to the trap height Co. Conservation District, http://www.vccd.org/; 40.015 N, 87. treatment, the same four trap bait treatments were tested as in 568 W), a 728-ha secondary forest dominated by species of oaks Experiment 1, with each pair of traps (understory and midcan- by guest on November 28, 2016 (Quercus species), hickories (Carya species), maples (Acer species), opy) receiving the same treatment. The experiment was conducted beech (Fagus species), and ash (Fraxinus species). Beetles were from 16 July to 11 September 2015, and traps were serviced at in- caught with panel traps (cross-vane, black corrugated plastic; tervals of 2 d, at which time treatments (per pair of traps) were AlphaScents, Portland, OR) which were coated with the fluoropoly- rotated along transects. mer dispersion Fluon PTFE (AGC Chemicals Americas, Inc., Exton, The of trapped beetles follows Monne´ and Hovore PA) to enhance trapping efficiency (for details, see Graham et al. (2005). Representative specimens of all species are available from 2010). Trap basins were partially filled with saturated aqueous the laboratory of L.M.H., and vouchers have been deposited with NaCl solution as a killing agent and preservative. the Illinois Natural History Survey, Champaign, IL. Traps were baited with either a blend of synthesized pheromones of cerambycids or a fermenting bait. The pheromone treatment com- prised a blend of pheromones of diverse species which had been de- veloped as a multispecies lure (Hanks et al. 2012), including Data Analysis pheromones of species in the subfamily Cerambycinae (racemic For each experiment, data were analyzed separately for the ceram- 3-hydroxyhexan-2-one, syn-2,3-hexanediol, racemic 2-methylbu- bycid species that were caught in greatest numbers ( 14 specimens tan-1-ol) and of species in the Lamiinae ([E]-fuscumol, [E]-fuscumol in total). Differences between treatments in mean numbers of beetles acetate, monochamol; reviewed by Millar and Hanks 2016). Lures caught, blocked by transect and collection date, were tested with the were formulated to contain 25 mg of each pheromone enantiomer nonparametric Friedman’s test (PROC FREQ with CMH option; (i.e., 50 mg of racemic compounds) dissolved in 1 ml of isopropanol. SAS Institute 2011) because data violated assumptions of analysis of Control lures contained 1 ml of neat isopropanol. Pheromone emit- variance (Sokal and Rohlf 1995). Thus, replicates were defined by ters were transparent polyethylene sachets (press-seal bags, Bagette trap transect and collection date. Assuming a significant overall test, model 14770, 5.1 by 7.6 cm, 0.05 mm thick, Cousin Corp., Largo, pairs of treatment means were compared using the Ryan-Einot- FL) which contained a cotton roll (1- by 4-cm dental wick, Patterson Gabriel-Welsch Q (REGWQ) multiple comparison test, which con- Dental Supply, Inc., St. Paul, MN). The recipe for the fermenting trols for maximum experiment-wise error rates (SAS Institute 2011). bait was based on the methods of Galford (1980), as follows: Only replicates containing at least one were included in analy- crushed banana (0.45 kg), brown sugar (0.45 kg), molasses (20 ml), ses. This threshold was increased for species that were caught in baker’s yeast (7 g), and coarsely chipped hickory wood (1.5 liter; great numbers, so as to improve the robustness of the analysis while Full Circle smoking wood for barbecue, Topco Associates LLC, Elk maintaining sufficient replication for good statistical power Grove, IL) were added to 3 liters of water and allowed to ferment (N 14; threshold ¼ 2 for Neoclytus mucronatus mucronatus (F.) for 3 d under ambient laboratory conditions (23C). Lures and 10 for Xylotrechus colonus (F.) in Experiment 1; see Results). Journal of Economic Entomology, 2016, Vol. 109, No. 5 2147

Results Table 1. Number of cerambycid beetles caught by panel traps in forested study sites of east-central Illinois during Experiments 1 During the two experiments, traps caught 885 cerambycid beetles of and 2 36 species in the subfamilies Cerambycinae, Lamiinae, Lepturinae, Parandrinae, and Prioninae, and one species in the closely related Taxonomy Expt. 1 Expt. 2 Total family Disteniidae (Table 1). Many of the species (N ¼ 18, or Cerambycinae 49%) were represented by five or fewer specimens, with nine spe- Anaglyptini cies represented by only a single specimen. Xylotrechus colonus,of Cyrtophorus verrucosus (Olivier) 2 2 the subfamily Cerambycinae, was caught in greatest numbers in both experiments (457 specimens, or 52% of the total; Table 1), Neoclytus a. acuminatus (F.) 13 6 19 followed by the cerambycine N. m. mucronatus, and three species in Neoclytus m. mucronatus (F.) 63 28 91 the subfamily Lamiinae: Graphisurus fasciatus (Degeer), Astylidius Neoclytus scutellaris (Olivier) 8 8 parvus (LeConte), and Lepturges angulatus (LeConte). Sarosesthes fulminans (F.) 17 17 In Experiment 1, 28, 21, 15, and 13 species of cerambycid bee- Xylotrechus colonus (F.) 354 103 457 tles were caught by traps baited with pheromone þ fermenting bait, Eburiini Eburia quadrigeminata (Say) 6 3 9 pheromone, fermenting bait, and control lures, respectively. Elaphidiini Nevertheless, traps baited with the pheromone blend caught signifi- Anelaphus pumilus (Newman) 19 19 cantly more cerambycid beetles in general (i.e., totaled across spe- Anelaphus villosus (F.) 12 12 cies) than did control traps (Fig. 1A), and trap catch was even Elaphidion mucronatum (Say) 11 1 12 greater for the combination of pheromone þ fermenting bait Parelaphidion aspersum (Haldeman) 4 3 7 Downloaded from (Friedman’s Q3,212 ¼88.4, P < 0.0001), suggesting that the ferment- Parelaphidion incertum (Newman) 1 1 ing bait acted additively. When tested alone, however, the ferment- Neoibidionini ing bait attracted few beetles (Fig. 1A). Heterachthes quadrimaculatus Haldeman 1 4 5 The significant lure treatment effect in Experiment 1 was due Tillomorphini primarily to the cumulative responses of the most numerous species, Euderces picipes (F.) 8 2 10 Lamiinae http://jee.oxfordjournals.org/ none of which were significantly attracted by fermenting bait alone (Table 2). Adults of X. colonus were significantly attracted by the Astyleiopus variegatus (Haldeman) 5 5 pheromone blend, and traps baited with both pheromones and fer- Astylidius parvus (LeConte) 16 19 35 menting bait caught twice as many beetles on average (Table 2). On Astylopsis macula (Say) 6 6 the other hand, adults of G. fasciatus were significantly attracted Graphisurus despectus (LeConte) 15 12 27 only by the combination of pheromones and fermenting bait. Adults Graphisurus fasciatus (Degeer) 39 14 53 of N. m. mucronatus, A. parvus, and L. angulatus apparently were Hyperplatys maculata Haldeman 3 3 equally attracted to the pheromone blend with and without ferment- Lepturges angulatus (LeConte) 35 35 ing bait (Table 2), indicating that the fermentation volatiles were Lepturges confluens (Haldeman) 1 2 3 by guest on November 28, 2016 not affecting attraction. The number of trapped beetles was not Sternidius alpha (Say) 5 1 6 Sternidius misellus (LeConte) 1 1 great enough to statistically discriminate between treatments for the Urgleptes foveatocollis (Hamilton) 1 1 cerambycines Anelaphus pumilus (Newman), Sarosesthes fulminans Urgleptes querci (Fitch) 1 1 2 (F.), and the lamiine Graphisurus despectus (LeConte) (Table 2). Acanthoderini During Experiment 2, the number of cerambycid species repre- Aegomorphus modestus (Gyllenhal) 9 7 16 sented in the trap catches again was not significantly influenced by neither the lure nor the trap height treatments, with cerambycid bee- Eupogonius pauper LeConte 1 1 tles of 13, 16, 10, and 6 species being found in traps baited with the Psenocerus supernotatus (Say) 1 1 pheromone þ fermenting bait, pheromone, fermenting bait, and con- Dorcaschematini trol treatments, with 16 species each in the understory versus Dorcaschema cinereum (Olivier) 1 1 midcanopy treatments. Traps baited with pheromones again caught Saperdini Saperda lateralis F. 1 1 more beetles than those baited with fermenting bait, or control Lepturinae traps, but in this case attraction was not significantly influenced by Lepturini the fermenting bait (Fig. 1B; Q3,92 ¼45.2, P < 0.0001). Adults of Brachyleptura rubrica (Say) 1 1 X. colonus again were significantly attracted by the pheromone Typocerus lugubris (Say) 4 4 blend (Table 2), and attraction was enhanced by the fermenting Rhagiini bait. Adults of N. m. mucronatus were significantly attracted only Gaurotes cyanipennis (Say) 1 1 by the combination of pheromones and fermenting bait, while adults Parandrinae of A. parvus were significantly attracted by the pheromone blend Parandrini and apparently not influenced by fermentation volatiles (Table 2). Neandra brunnea (F.) 2 2 Too few adults of G. fasciatus were caught to statistically discrimi- Prioninae Prionini nate between treatments. Orthosoma brunneum (Forster) 4 2 6 During Experiment 2, traps in the forest understory caught sig- Disteniidae nificantly greater numbers of beetles than did traps in the canopy Elytrimitatrix undata (F.) 4 1 5 (means 6.0 6 1.2 and 3.7 6 0.75 beetles per replicate, respectively; Total 661 224 885 Q1,46 ¼5.0, P ¼ 0.03). Among the most numerous species, two were trapped in greater numbers in the forest understory: X. colonus (3.9 6 0.6 and 1.3 6 0.3 beetles per replicate for understory and 2148 Journal of Economic Entomology, 2016, Vol. 109, No. 5

canopy, respectively; Q1,50 ¼14.2, P > 0.01), G. fasciatus (1.5 6 0.4

and 0.25 6 0.16 beetles, Q1,16 ¼4.5, P ¼ 0.03). On the other hand, most adults of N. m. mucronatus were caught by traps in the canopy

(0.23 6 0.1 and 1.9 6 0.5 beetles, Q1,26 ¼6.23, P ¼ 0.01). Adults of A. parvus were caught in similar numbers by traps at the two heights

(1.0 6 0.29 and 1.1 6 0.5 beetles; Q1,18 ¼0.01, P > 0.05).

Discussion The species that were trapped in greatest numbers during the present study already were known to be attracted by the same blend of synthesized pheromones from previous studies (e.g., Hanks et al. 2012, Hanks and Millar 2013, Handley et al. 2015). Adults of X. colonus were attracted by the blend because it contains all four components of the pheromone produced by males of this species, including (R)- and (S)-3-hydroxyhexan-2-one, and (2R,3R)- and (2S,3S)-2,3-hexanediol (Lacey et al. 2009, Hanks et al. 2012). (R)-3-Hydroxyhexan-2-one also comprises the sole or dominant

pheromone component of A. pumilus, N. m. mucronatus, and S. ful- Downloaded from minans (Lacey et al. 2007, 2009; Mitchell et al. 2015). Finally, the fuscumol and fuscumol acetate components of the blend are known to be pheromone components of the lamiines A. parvus and L. angu- latus (Meier et al. 2016), and also are likely pheromone components of G. fasciatus (Mitchell et al. 2011). http://jee.oxfordjournals.org/ The only species that were significantly influenced by volatiles from the fermenting bait were the cerambycine X. colonus and the lamiine G. fasciatus, with the fermentation volatiles enhancing at- traction to the pheromone blend. This finding suggests that adults of X. colonus feed on fermenting tree sap, as do adults of some other species of cerambycines (Linsley 1959). Adults of both X. colonus and G. fasciatus have been reported to be attracted by fermenting bait (e.g., Galford 1980), although the fermenting bait used in the Fig. 1. Mean (6 1 SE) number of cerambycid beetles (all species combined) by guest on November 28, 2016 caught per replicate during Experiment 1 (A) and Experiment 2 (B). Means present study attracted few beetles of any species when tested alone. with different letters within experiments are significantly different (REGWQ This inconsistency likely is due to differences between studies in the test, P < 0.05). formulation of their fermenting baits. Different combinations of

Table 2. Mean (6 1 SE) number of beetles caught per replicate by experimental treatment for the cerambycid species represented by the greatest numbers of specimens (N 14) in Experiments 1 and 2, and results of Friedman’s tests

Species Pheromones þ fermenting baita Pheromones Fermenting bait Control Friedman’s Q (df)b

Experiment 1 Cerambycinae Anelaphus pumilusc 2.5 6 1.3 2.3 6 1.3 0 0 8.36 (16)* Neoclytus m. mucronatus 1.9 6 0.5a 1.4 6 0.3a 0.06 6 0.06b 0b 30.4 (64)*** Sarosesthes fulminansc 1.0 6 0.41 0.78 6 0.04 0.11 6 0.01 0 9.8 (36)* Xylotrechus colonus 10.6 6 1.1a 5.3 6 0.7b 0.79 6 0.3c 0.43 6 0.2c 35.5 (56)*** Lamiinae Astylidius parvus 0.46 6 0.2ab 0.69 6 0.1a 0.08 6 0.08bc 0b 15.4 (52)** Graphisurus despectus 0.77 6 0.3 0.33 6 0.2 0.22 6 0.2 0.33 6 0.2 3.65 (36) Graphisurus fasciatus 1.4 6 0.37a 0.61 6 0.2b 0.11 6 0.1b 0.06 6 0.06b 20.0 (72)** Lepturges angulatus 0.92 6 0.3a 0.92 6 0.2a 0b 0b 22.0 (52)*** Experiment 2 Cerambycinae Neoclytus m. mucronatus 1.2 6 0.38a 0.69 6 0.2ab 0.23 6 0.2b 0b 13.9 (52)** Xylotrechus colonus 3.35 6 0.5a 1.5 6 0.4b 0.2 6 0.09c 0.15 6 0.08c 40.4 (80)*** Lamiinae Astylidius parvus 1.0 6 0.5a 1.11 6 0.2a 0b 0b 17.9 (36)** Graphisurus fasciatus 0.63 6 0.26 0.5 6 0.27 0.38 6 0.26 0.25 6 0.16 0.44 (32)

a Means with different letters within species are significantly different (REGWQ test, P < 0.05). b Asterisks indicate significance level of Friedman’s Q:*P < 0.01; ** P < 0.001; *** P < 0.0001. c Overall analyses for A. pumilus and S. fulminans in Experiment 1 were significant at P < 0.01, but the means separation test failed to detect differences be- tween pairs of means. Journal of Economic Entomology, 2016, Vol. 109, No. 5 2149 fruit, sources of sugar and yeast, and alcoholic beverages undoubt- pine wood nematode vector Monochamus galloprovincialis. J. Appl. edly will yield unique volatile profiles as they ferment, which may Entomol. 139: 618–626. result in their attracting different species of insects (Galford 1980). Bouget, C., H. Brustel, A. Brin, and T. Noblecourt. 2008. Sampling saproxylic  Moreover, the volatile profiles of these baits will evolve over time, beetles with window flight traps: Methodological insights. Rev. Ecol. (Terre Vie), Suppt. 10: 21–32. beginning with anaerobic production of ethanol, followed by acidic Brockerhoff, E. G., A. M. Liebhold, and H. Jactel. 2006. The ecology of forest fermentation and production primarily of acetic acid, and then pu- invasions and advances in their management. Can. J. For. Res. 36: trefaction (Frost and Dietrich 1929, and references therein). Thus, 263–268. standardizing volatile profiles, both in terms of the initial formula- Champlain, A. B., and J. N. Knull. 1932. Fermenting bait traps for trapping tion and its fermentation in the field, poses a challenge to developing Elateridae and Cerambycidae (Coleop.). Entomol. News 43: 253–257. these types of baits as reliable trap attractants. Inconsistency in vola- Dodds, K. J., J. D. Allison, D. R. Miller, R. P. Hanavan, and J. Sweeney. 2015. tile profiles of fermenting baits may explain why the bait used in the Considering species richness and rarity when selecting optimal survey traps: earlier study of Schmeelk et al. (2016), which was tested at the same Comparisons of semiochemical baited flight intercept traps for site as used in the present study, attracted adults of the cerambycine Cerambycidae in eastern North America. Agric. For. Entomol. 17: 36–47. Eburia quadrigeminata (Say), whereas few beetles of this species Frost, S. W., and H. Dietrich. 1929. Coleoptera taken from bait-traps. Ann. Entomol. Soc. Am. 22: 427–436. were caught during the present study. Galford, J. R. 1980. Research Note NE-293 Bait bucket trapping for red oak The apparent preference in adults of X. colonus for traps in the borers (Coleoptera: Cerambycidae). U.S. Department of Agriculture of forest understory also is consistent with the findings of Schmeelk Forest Services, Broomall, PA. et al. (2016). Adults of this species likely prefer this height because Gardiner, L. M. 1960. Descriptions of immature forms and biology of males and females aggregate on larval hosts such as downed hard- Xylotrechus colonus (Fab.) (Coleoptera: Cerambycidae). Can. Entomol. 92: Downloaded from wood trees and fallen branches. Moreover, adults of X. colonus do 820–825. not feed (Gardiner 1960, Waters 1981, L.M.H., unpublished data), Graham, E. E., R. F. Mitchell, P. F. Reagel, J. D. Barbour, J. G. Millar, and L. and so need not disperse into the forest canopy in search of food. M. Hanks. 2010. Treating panel traps with a fluoropolymer enhances their Preference in adults of G. fasciatus for understory traps, and in efficiency in capturing cerambycid beetles. J. Econ. Entomol. 103: 641–647. N. m. mucronatus for midcanopy traps, are difficult to interpret be- Graham, E. E., T. M. Poland, D. G. McCullough, and J. G. Millar. 2012. A comparison of trap type and height for capturing cerambycid beetles cause there is no published information on how their host plants are http://jee.oxfordjournals.org/ (Coleoptera). J. Econ. Entomol. 105: 837–846. distributed in forests, to our knowledge. Handley, K., J. Hough-Goldstein, L. M. Hanks, J. G. Millar, and V. D’Amico. The findings of this study provide further evidence that the spe- 2015. Species richness and phenology of cerambycid beetles in urban forest cies diversity of cerambycid communities, and population densities fragments of northern Delaware. Ann. Entomol. Soc. Am. 108: 251–262. of their species, would be most accurately assessed by positioning Hanks, L. M., and J. G. Millar. 2013. Field bioassays of cerambycid phero- traps at multiple heights within the forest canopy. The lack of signif- mones reveal widespread parsimony of pheromone structures, enhancement icant attraction to fermenting bait, and the fact that it moderately by host plant volatiles, and antagonism by components from heterospecifics. enhanced attraction to pheromones for only two species, suggests Chemoecology 23: 21–44. that such baits hold little promise for improving methods of trap- Hanks, L. M., and Q. Wang. 2016. Reproductive biology of cerambycid bee- by guest on November 28, 2016 ping cerambycids. Nevertheless, it should be noted that we tested tles. In Q. Wang (ed.), Cerambycidae of the world: Biology and manage- ment. CRC Press/Taylor & Francis, Boca Raton, FL. only one type of fermenting bait and one blend of synthesized phero- Hanks, L. M., J. G. Millar, J. A. Mongold-Diers, J.C.H. Wong, L. R. Meier, P. mones. Other fermenting formulations, which have different odor F. Reagel, and R. F. Mitchell. 2012. Using blends of cerambycid beetle pher- profiles, could be more effective in attracting wood boring insects. omones and host plant volatiles to simultaneously attract a diversity of Moreover, other blends of synthesized pheromones may attract a cerambycid species. Can. J. For. Res. 42: 1050–1059. different set of cerambycid species that may be more strongly influ- Hughes, C. C., R. C. Johns, and J. D. Sweeney. 2014. A technical guide to in- enced by volatiles from fermenting bait. stalling beetle traps in the upper crown of trees. J. Acad. Entomol. Soc. 10: 12–18. Lacey, E. S., M. D. Ginzel, J. G. Millar, and L. M. Hanks. 2004. Male- produced aggregation pheromone of the cerambycid beetle Neoclytus acu- Acknowledgments minatus acuminatus. J. Chem. Ecol. 30: 1493–1507. Lacey, E. S., J. A. Moreira, J. G. Millar, A. M. Ray, and L. M. Hanks. 2007. We thank J. A. Mongold-Diers for assistance in preparing trap lures, Male-produced aggregation pheromone of the cerambycid beetle Neoclytus S. Vachula for assistance with field work, and J. G. Millar for helpful com- mucronatus mucronatus. Entomol. Exp. Appl. 122: 171–179. ments on an early draft of the manuscript. We appreciate funding support Lacey, E. S., J. G. Millar, J. A. Moreira, and L. M. Hanks. 2009. Male- from The Alphawood Foundation of Chicago (to L.M.H.), and USDA produced aggregation pheromones of the cerambycid beetles Xylotrechus National Institute of Food and Agriculture (grant 2012-67013-19303; to J. G. colonus and Sarosesthes fulminans. J. Chem. Ecol. 35: 733–740. Millar and L.M.H.). Linsley, E. G. 1959. Ecology of Cerambycidae. Annu. Rev. Entomol. 4: 99–138. Meier, L. R., Y. Zou, J. G. Millar, J. A. Mongold-Diers, and L. M. Hanks. References Cited 2016. Synergism between enantiomers creates species-specific pheromone Allison, J. D., C. W. Johnson, J. R. Meeker, B. L. Strom, and S. M. Butler. blends and minimizes cross-attraction for two species of cerambycid beetles. 2011. 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