Forest Entomology Research Article

Journal of Economic Entomology, 109(4), 2016, 1750–1757 doi: 10.1093/jee/tow102 Advance Access Publication Date: 13 June 2016 Forest Entomology Research article

Influence of Trap Height and Bait Type on Abundance and Species Diversity of Cerambycid Beetles Captured in Forests of East-Central Illinois

Thomas C. Schmeelk,1 Jocelyn G. Millar,2 and Lawrence M. Hanks1,3

1Department of Entomology, University of Illinois at Urbana-Champaign, Urbana, IL 61801 ([email protected]; hanks@life. illinois.edu), 2Department of Entomology, University of California, Riverside, CA 92521 ([email protected]) and 3Corresponding author, e-mail: [email protected]

Received 2 March 2016; Accepted 19 April 2016 Downloaded from Abstract We assessed how height of panel traps above the forest ﬂoor, and the type of trap bait used, inﬂuenced the abundance and diversity of cerambycid beetles caught in forested areas of east-central Illinois. Panel traps were sus- pended from branches of hardwood trees at three heights above the ground: understory (1.5 m), lower canopy

(6 m), and midcanopy (12 m). Traps were baited with either a multispecies blend of synthesized cerambycid http://jee.oxfordjournals.org/ pheromones or a fermenting bait mixture. Traps captured a total of 848 beetles of 50 species in the cerambycid subfamilies Cerambycinae, Lamiinae, Lepturinae, and Parandrinae, and one species in the closely related family Disteniidae. The species caught in highest numbers was the cerambycine Anelaphus pumilus (Newman), represented by 349 specimens. The 17 most abundant species (mean 6 1 SD: 45 6 80 specimens per species) included 12 cerambycine and five lamiine species. Of these most abundant species, 13 (77%) were attracted to traps baited with the pheromone blend. Only the cerambycine Eburia quadrigeminata (Say) was attracted by the fermenting bait. Three species were captured primarily in understory traps, and another five species primarily in midcanopy traps. Variation among cerambycid species in their vertical distribution in forests accounted for simi- by guest on August 11, 2016 lar overall abundances and species richness across trap height treatments. These findings suggest that trapping surveys of native communities of cerambycids, and quarantine surveillance for newly introduced exotic species, would be optimized by including a variety of trap baits and distributing traps across vertical strata of forests.

Key words: wood-boring insect, pheromone, fermenting bait, monitoring, invasive species

The longhorned beetles (Cerambycidae) are among the most diverse valuable tool for managing native cerambycid species that are pests of insect groups, with an estimated 35,000 species worldwide (e.g., Maki et al. 2011), for assessing biodiversity (e.g., Hanks et al. (Sv acha and Lawrence 2014). The endophytic larvae are generally 2014), and for developing methods for conserving endangered na- beneficial in forest ecosystems because they contribute to the decom- tive species (e.g., Ray et al. 2014). Research over the past decade has position of dead woody plants. However, some species become im- revealed that cerambycid species of the subfamilies Cerambycinae, portant pests by killing living trees or damaging trees felled for Lamiinae, and Spondylidinae use aggregation pheromones that are lumber (Craighead 1923, Linsley 1959). Due to their endophytic na- produced by males and attract both sexes, whereas species in the ture, the larvae are readily transported by international commerce, Prioninae and Lepturinae use sex pheromones that are produced by concealed within nursery stock, solid wood packing materials, and females and attract only males (reviewed by Millar and Hanks other wood products (e.g., Haack 2006, Cocquempot and Lindelo¨w 2016). There appears to be substantial pheromonal parsimony 2010). Several species of exotic cerambycids have invaded North within the family in terms of both closely related species (conge- America in recent years, among them the Asian longhorned beetle, neric) and more distantly related species (in different subfamilies) Anoplophora glabripennis (Motschulsky), a devastating pest of producing the same, or very similar, pheromone components. hardwood forests (Haack et al. 2010). Similarity in pheromone composition across taxonomic groups The continuing threat posed by exotic and potentially invasive of the Cerambycidae results in simultaneous attraction of multiple cerambycid species has inspired research on attractants for use as species to traps baited with blends of synthetic pheromones (e.g., trap baits, both to monitor for new incursions (e.g., Narai et al. Hanks and Millar 2013, Handley et al. 2015). The abundance and 2015) and to manage exotic species that already have established diversity of cerambycids that are attracted by such multispecies (e.g., Nehme et al. 2010, Zou et al. 2016). Trapping also may be a pheromone lures can be further improved by combining them with

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] 1750 Journal of Economic Entomology, 2016, Vol. 109, No. 4 1751 lures that release synergistic plant volatiles such as ethanol and a-pi- Field Experiment nene, and bark beetle pheromones that apparently act as kairo- Beetles were captured with black cross-vane panel traps (Alpha mones (e.g., Miller et al. 2015, Ryall et al. 2015). It has long been Scents, Inc., West Linn, OR) that were coated with Fluon to improve suggested that cerambycids also may be attracted to traps by fer- capture efficiency (Northern Products, Woonsocket, RI; for details, menting baits whose odors mimic those produced by decaying host see Graham et al. 2010). Trap basins contained 300 ml of satu- trees or the fermenting tree sap on which the adults feed (e.g., Frost rated aqueous NaCl solution to kill and preserve captured beetles. and Dietrich 1929, Champlain and Knull 1932, Galford 1980). A set of traps comprised three traps hung in different trees that Since those early studies, however, fermenting baits have rarely been were 12 m apart, one trap at each of three heights: the understory investigated as attractants for cerambycid beetles. (bottom of trap 1.5 m above the ground), lower canopy (6 m), Trapping surveys that target wood-boring insects usually are de- and midcanopy (12 m). Traps were hung from tree branches with ployed in the forest understory simply for convenience (Dodds nylon rope in mid-April 2014, before the trees had leafed out. Traps 2014), but the abundance and species diversity of captured insects in the two highest positions were set up by first throwing a line over may be greatly influenced by the height of traps above the ground the branch with a large slingshot (Big Shot, Sherrill Inc., (e.g., Ulyshen and Hanula 2007, Graham et al. 2012, Dodds 2014). Greensboro, NC) following the methods of Hughes et al. (2014). Adult cerambycids may not be distributed uniformly throughout the Traps were stabilized by attaching a rope from the trap bottom to a forest canopy, but rather may concentrate their activities within spe- stake in the ground. cific vertical strata as they search for mates, and/or hosts that are For each set of three traps, those at the two lower positions were suitable for oviposition. Species whose larvae develop in branches or in shagbark hickories, Carya ovata (Mill.) K. Koch, whereas those twigs of living trees may spend most of their adult lives in the tree at the highest position were in white oaks, Quercus alba L., due to Downloaded from canopy, whereas those that breed in roots of trees, or in fallen trees the availability of suitably high and horizontal branches in that spe- and branches may have no need to fly above the understory. For ex- cies. Although this experimental design confounds the highest trap ample, adults of the European cerambycine Clytus tropicus (Panzer) position with tree species, it was deemed worthwhile because many are restricted to the forest canopy, which has resulted in the false im- species of the local cerambycid fauna are polyphagous, their larvae pression that the species is rare (see Mannerkoski et al. 2010). On feed in both oaks and hickories (see Results; Lingafelter 2007), and the other hand, adults of the North American prionine Orthosoma because the hickory and oak trees that were used were adjacent to http://jee.oxfordjournals.org/ brunneum (Forster) do not venture above the forest understory be- one another. cause they do not feed, and larval hosts are decaying trees on the for- Two sets of three traps (separated by 30 m) were installed at est floor (Ulyshen and Hanula 2007). each of the two field sites, with one set baited with a blend of syn- Here, we assessed how the height of traps in forested areas of east- thetic pheromones and the other set baited with a fermenting bait central Illinois influenced the abundance and diversity of cerambycid mixture. The pheromone blend was a combination of synthetic ag- beetles that they captured. Traps were deployed at three heights above gregation pheromones known to attract many species native to the the forest floor: understory, lower canopy, and midcanopy. In con- study area (Hanks et al. 2014), including pheromones characteristic junction with the trap height effect, we compared attraction to a fer- of cerambycine species (3-hydroxyhexan-2-one, syn-2,3-hexanediol, by guest on August 11, 2016 menting bait versus a blend of synthetic pheromones that was 2-methylbutan-1-ol) and lamiine species ([E]-fuscumol, [E]-fuscu- developed as a multispecies lure. The trap height effect has rarely mol acetate, monochamol; Millar and Hanks 2016). The phero- been tested using synthetic pheromones of cerambycids as bait mone blend was dispensed from resealable polyethylene sachets (Graham et al. 2012, Dodds 2014, Webster et al. 2016), and to our (5.1 7.6 cm, Bagettes model 14770, Cousin Corp., Largo, FL) that knowledge has not been tested using a fermenting bait. contained a cotton roll (1 4 cm dental wick, Patterson Dental Supply Inc., St. Paul, MN). The fermenting bait was based on rec- ipes suggested by Steyskal et al. (1986), prepared by combining beer Materials and Methods (355 ml, Rolling Rock, Latrobe Brewing Co., St. Louis, MO), molas- ses (100 ml), half a thinly sliced banana, granulated sugar (50 g), Study Sites and baker’s yeast (34 g, Fleischmann’s Associated British Foods, The two study sites were in Vermilion County, east-central Illinois, London, United Kingdom), diluted to 1 liter with tap water. and were 19 km apart: Forest Glen Preserve (Vermilion Co. Fermenting bait lures consisted of 250 ml of the solution in polyeth- Conservation District, http://www.vccd.org/ last accessed 30 April ylene sachets (10 15 cm, Bagettes model 14772, Cousin Corp.) 2016; 728 ha, 40.015N, 87.568W), and Vermilion River with two holes (5.5 mm) punched in the top above the liquid level Observatory (University of Illinois natural area, http://research.illi to increase release rate of volatiles while excluding insects such as nois.edu/cna/ last accessed 30 April 2016; 193 ha, 40.0656N, 87. moths and wasps that may have been attracted by the odors. 561W). Both sites were secondary forests dominated by species of The experiment was conducted from 5 May to 23 September maples (Acer species), hickories (Carya species), beech (Fagus spe- 2014, and treatments were assigned randomly to trap sets within cies), and oak (Quercus species). sites on the first day. Traps were checked for beetles every 1–3 d, and were rebaited every 2 wk, at which time treatments were switched between sets (within sites) to control for location effects. Sources of Chemicals Taxonomy of captured beetles follows Lingafelter (2007). Racemic 3-hydroxyhexan-2-one, monochamol (2-[undecyloxy]etha- Representative specimens of all species are available from the labo- nol), racemic (E)-fuscumol ([E]-6,10-dimethylundeca-5,9-dien-2- ratory collection of LMH. ol), and racemic (E)-fuscumol acetate ([E]-6,10-dimethylundeca- 5,9-dien-2-yl acetate) were purchased from Bedoukian Research (Danbury, CN), and racemic 2-methylbutan-1-ol was purchased Data Analysis from Sigma-Aldrich (St. Louis, MO). Syn-2,3-hexanediol was syn- Because assumptions of analysis of variance were violated by hetero- thesized as described in Lacey et al. (2004). scedasticity (Sokal and Rohlf 1995), differences between treatments 1752 Journal of Economic Entomology, 2016, Vol. 109, No. 4 in mean numbers of beetles captured (of all cerambycid species com- significantly attracted to traps baited with the pheromone blend, in- bined, and separately for individual species) were tested with the cluding both cerambycines and lamiines (Table 2). Only the ceram- nonparametric Friedman’s test (PROC FREQ with CMH option; bycine Eburia quadrigeminata (Say) was captured in significantly SAS Institute 2011). The experiment was replicated spatially (by greater numbers by traps with fermenting baits (27 of 29 beetles, study site) and temporally (by collection date). Patterns in treatment 93%; Tables 1 and 2). effects were consistent across study sites, so the nonsignificant site Trap catches of about half of the most abundant species were sig- term was dropped from analyses. Pairs of treatment means were nificantly influenced by trap position (Table 3). Traps in the under- compared with the nonparametric Dunn–Nemenyi multiple compar- story captured the greatest numbers of the cerambycines Neoclytus ison test (Zar 2010, Elliot and Hynan 2011). Differences between a. acuminatus (F.) and X. colonus, and the lamiine Astylidius parvus trap height treatments in species richness (i.e., total number of (LeConte), whereas traps in the midcanopy caught the greatest num- cerambycid species captured over the season) were tested with the bers of the cerambycines E. quadrigeminata, Neoclytus scutellaris v2 test (assuming an even distribution across heights). The trap bait (Olivier), Parelaphidion incertum (Newman), and Phymatodes lengi effect was tested separately for the species caught in greatest num- Joutel, and the lamiine Aegomorphus modestus (Gyll.). An excep- bers (total of at least nine specimens), dropping replicates with zero tion was the cerambycine Phymatodes aereus (Newman), which was specimens. In the analysis of the trap height data, we optimized sam- captured in similar numbers by traps in both the understory and ple size per replicate for the two species caught in largest numbers lower canopy. The remaining species showed no significant prefer- (Anelaphus pumilus [Newman] and Xylotrechus colonus [F.], see ence for one trap height over another. Results), while maintaining sufficient replication for a robust analy- Sex ratios of captured beetles were approximately equal across sis (at least 10 replicates) by dropping replicates that had fewer than all treatments, ranging from 46 to 55% for the fermenting bait and Downloaded from a threshold number of specimens (N ¼ 4 and 3, respectively). To pheromone blend treatments, and from 52 to 56% for the three confirm that both sexes were responding to trap bait and height height treatments (v2 tests, P > 0.05). This suggests that males and treatments, beetles were drawn from the statistically significant females were similarly influenced by trap height and trap bait. Sex treatments (species with at least 25 specimens) and sexed. Sex ratios ratios within treatments for the individual most abundant species of captured beetles (% female) in different treatments were com- also approximated 50%, the exception being P. lengi, with 4 males 2 pared statistically using the v test. and 16 females captured by midcanopy traps that were baited with http://jee.oxfordjournals.org/ the pheromone blend (sex ratio 80%, significantly different from 50%, v2 ¼ 7.2, P ¼ 0.0073). Results The traps captured 848 cerambycid beetles of 50 species in the sub- Discussion families Cerambycinae, Lamiinae, Lepturinae, and Parandrinae, including one species in the closely related family Disteniidae Most of the species attracted by the blend of synthetic pheromones (Table 1; Sv acha and Lawrence 2014). Thirty species (60%) were previously had been shown to be attracted by that same blend in by guest on August 11, 2016 represented by five or fewer specimens, of which 19 species were other field studies (Hanks et al. 2012, Hanks and Millar 2013, represented by only a single specimen. The cerambycine A. pumilus Handley et al. 2015). The lamiine Graphisurus despectus (LeConte) was caught in greatest numbers (349 specimens, 41% of the total). also had been reported to be attracted by the blend (Handley et al. Traps baited with the pheromone blend captured 727 beetles 2015), although the sample size in the present study may have been (86% of the total) compared to 121 beetles in traps with ferment- too small to confirm the treatment effect. Attraction of a species to ing bait (means 6 1 SE: 9.6 6 1.9 and 1.6 6 0.24 beetles per repli- the blend is evidence that its pheromone is contained in the blend, cate, respectively; significantly different: Friedman’s Q1,152 ¼ 37.0, which has been confirmed for many of the most abundant species P < 0.0001). However, the two treatments did not differ signifi- (Table 4). Thus, pheromones have not yet been identified for the cantly in species richness, with traps baited with the pheromone cerambycine P. testaceus and the lamiines A. modestus and G. fas- blend or fermenting bait catching beetles of 40 and 36 species, re- ciatus, but their attraction to the blend suggests that at least some spectively (v2 ¼ 0.33, P > 0.05). components of their pheromones are present in the blend (Table 4). Traps positioned at the three heights captured similar numbers The cerambycine E. quadrigeminata was the only species to be of cerambycid beetles, with totals of 276, 285, and 287 beetles in attracted by the fermenting bait mixture in large numbers, and has traps in the understory, and at the lower and midcanopy, respec- long been known to be attracted to various fermenting bait concoc- tively (means 3.6 6 0.71, 3.8 6 0.79, and 3.8 6 0.60, respectively; tions (e.g., Frost and Dietrich 1929, Champlain and Knull 1932, not significantly different: Friedman’s Q2,228 ¼ 3.7, P ¼ 0.15). Galford 1980, Hanks and Millar 2013). Females of this species ovi- Species richness also was not influenced by trap height, ranging posit into wounds in the bark of living trees, which provides the neo- from 30 to 38 species (Fig. 1; v2 ¼ 1.2, P ¼ 0.6). Twenty of the 50 nate larvae access to the heartwood (Craighead 1923). Sap species (40%) were captured by traps at all three heights (Fig. 1). produced from these wounds may ferment, releasing volatiles that Another 12 species were captured only by midcanopy traps, more attract beetles of both sexes to feed, and females to oviposit. Adult than twice the number that were unique to the remaining height E. quadrigeminata apparently were not attracted by any compo- treatments (Fig. 1), but this difference also was not statistically sig- nents of the pheromone blend, nor have they demonstrated evidence nificant (v2 ¼ 5.4, P ¼ 0.066). It should be noted that all of the spe- of attraction to any known cerambycid pheromones in numerous cies that were unique to one trap position were represented by few field bioassays conducted in the study area (see Hanks et al. 2014). specimens (usually N ¼ 1; Table 1), thus any patterns in species rich- It is possible that this species does not have a volatile pheromone, ness across treatments were likely due to sampling error. and location of mates depends on mutual attraction of both sexes to A group of 17 species was designated as most abundant, being host plants (see Millar and Hanks 2016). Consistent with this hy- represented by at least nine specimens (mean 6 1 SD: 45 6 80 beetles pothesis, male E. quadrigeminata lack the prothoracic glands that per species). Of these most abundant species, 13 (77%) were are the source of the hydroxyketones and 2,3-alkanediols that serve Journal of Economic Entomology, 2016, Vol. 109, No. 4 1753

Table 1. Taxonomy and numbers of cerambycid beetles that were captured in east-central Illinois by panel traps baited with either a blend of synthesized pheromones or a fermenting bait, and positioned at three heights above the ground: understory (Low), lower canopy (Medium), or midcanopy (High)

Taxonomy Pheromone bait Fermenting bait Total

Low Medium High Low Medium High

Cerambycinae Anaglyptini Cyrtophorus verrucosus (Olivier) 3 1 1 2 7 Callidiini Phymatodes aereus (Newman) 24 20 1 1 46 Phymatodes amoenus (Say) 14 12 17 1 2 46 Phymatodes lengi Joutel 3 1 20 1 25 Phymatodes testaceus (L.) 5 4 3 12 Phymatodes varius (F.) 2 3 5 Clytini Clytoleptus albofasciatus (Laporte & Gory) 1 1 Megacyllene caryae (Gahan) 3 2 5 Neoclytus a. acuminatus (F.) 11 5 3 3 22

Neoclytus m. mucronatus (F.) 5 3 4 3 15 Downloaded from Neoclytus scutellaris (Olivier) 3 1 5 9 Xylotrechus colonus (F.) 33 20 13 11 1 3 81 Eburiini Eburia quadrigeminata (Say) 2 2 2 23 29 Elaphidiini

Anelaphus pumilus (Newman) 98 151 99 1 349 http://jee.oxfordjournals.org/ Anelaphus villosus (F.) 1 12 Elaphidion mucronatum (Say) 2 1 1 1 2 7 Parelaphidion aspersum (Haldeman) 1 7 3 2 13 Parelaphidion incertum (Newman) 1 1 6 1 2 11 Stenosphenus notatus (Olivier) 1 1 Neoibidionini Heterachthes quadrimaculatus var. pallidus Haldeman 1 1 Obriini Obrium maculatum (Olivier) 1 1 by guest on August 11, 2016 Tillomorphini Euderces picipes (F.) 11 Euderces pini (Olivier) 11 Trachyderini Purpuricenus axillaris Haldeman 11 Lamiinae Acanthocinini Astylidius parvus (LeConte) 17 6 4 1 1 29 Astylopsis collaris (Haldeman) 1 1 Astylopsis macula (Say) 1 1 Graphisurus despectus (LeConte) 4 6 4 2 1 3 20 Graphisurus fasciatus (Degeer) 6 9 6 1 3 4 29 Hyperplatys maculata Haldeman 1 1 2 Leptostylus transversus (Gyll.) 1 2 3 Lepturges angulatus (LeConte) 5 7 2 2 16 Lepturges conﬂuens (Haldeman) 2 2 4 Sternidius alpha (Say) 1 3 4 Acanthoderini Aegomorphus modestus (Gyll.) 3 3 8 1 4 19 Dorcaschematini Dorcaschema cinereum (Olivier) 1 1 Monochamini Goes tigrinus (Degeer) 1 1 Microgoes oculatus (LeConte) 1 1 Pogonocherini Ecyrus d. dasycerus (Say) 1 113 Saperdini Saperda discoidea F. 11 Saperda imitans Felt & Joutel 2 2

(continued) 1754 Journal of Economic Entomology, 2016, Vol. 109, No. 4

Table 1. continued Taxonomy Pheromone bait Fermenting bait Total

Low Medium High Low Medium High

Lepturinae Lepturini Bellamira scalaris (Say) 1 1 Brachyleptura rubrica (Say) 1 1 1 3 Stenelytrana emarginata (F.) 11 Strangalia bicolor (Swederus) 11 Rhagiini Gaurotes cyanipennis (Say) 11 Stenocorus cinnamopterus (Randall) 1 1 1 1 1 5 Stenocorus schaumii (LeConte) 1 1 Parandrinae Parandrini Neandra brunnea (F.) 1 1 Disteniidae Disteniini Elytrimitatrix undata (F.) 1 1 1 1 2 6 Total: 246 265 216 30 20 71 848 Downloaded from

Dodds 2014, Webster et al. 2016). In contrast, other species that

were caught in large numbers, such as A. pumilus, were evenly dis- http://jee.oxfordjournals.org/ tributed across trap heights, indicating that they move freely between ground level and the canopy. The approximately equal sex ratio in traps at all heights for almost all the most abundant species argues against height preference being based solely on females seeking oviposition sites. Some adult cerambycines feed on pollen or tree sap, whereas others do not feed as adults (Hanks and Wang 2016). Among the cerambycines caught in significant numbers in the present study, only A. pumilus is by guest on August 11, 2016 known to feed on pollen, whereas N. a. acuminatus, N. m. mucronatus, P. lengi, P. testaceus, and X. colonus do not (L.M.H., unpub. data), although N. a. acuminatus has been reported to feed on tree sap (Waters 1981). Pollen feeding by A. pumilus apparently does not limit the height at which adults fly because they were equally abundant across the low, medium, and high forest strata. However, this indiscriminate distribution may be due to adults of both sexes alternately moving up into the canopy in search of flowers for feeding, and down into the understory when seeking mates and oviposition sites.

Fig. 1. Venn diagram showing the total number of cerambycid species cap- Most cerambycine species mate on the larval host (Linsley tured by panel traps that were positioned at three heights in the forest canopy 1959). Therefore, the vertical distribution of adults may be associ- (regardless of trap bait): understory (1.5 m above the ground), lower canopy ated with the spatial distribution of larval hosts, especially for spe- (6 m), and midcanopy (12 m). Also shown is the number of species cap- cies that do not feed as adults. Many of the most abundant species tured only at one trap height (in parens), and the number captured at multiple in the present study are moderately to highly polyphagous on hard- heights. woods, including oaks and hickories which were common at the study sites, the exceptions being P. amoenus (larvae feeding within vines of grape, Vitis species) and P. lengi (larval hosts unknown; as pheromones for many species of cerambycines (Ray et al. 2006). Lingafelter 2007). It is not known whether adult P. amoenus feed, Nevertheless, cerambycines of some species produce pheromones of but their even distribution across forest strata could be due to the different structures which apparently are released by glands else- climbing nature of the larval hosts. Capture of adult P. lengi primar- where on the body of males (e.g., Ray et al. 2009). ily in the midcanopy suggests that their larvae may develop in About half of the most abundant species appeared to prefer spe- branches or twigs of living trees well above the ground, which could cific vertical strata in the forest, with some species being caught in explain why larval hosts of this species have not yet been identified. traps below the canopy (N. a. acuminatus, P. aereus, X. colonus, There is limited published information on the spatial distribution A. parvus) whereas others were more active within the canopy of larval hosts for the most abundant species, to our knowledge, and (E. quadrigeminata, N. scutellaris, P. incertum, P. lengi, A. modes- so effects on the distribution of adults are difficult to assess. tus). These findings corroborate previous reports that activities of Nevertheless, the hypothesis that adults prefer certain forest strata cerambycid species are stratified within forests (Graham et al. 2012, based on the distribution of larval hosts is supported by the field Journal of Economic Entomology, 2016, Vol. 109, No. 4 1755 experiments of Waters (1981) who studied host choice in ceramby- study. She also reported that adult N. scutellaris almost exclusively cids of Alabama by mounting sticky cards on felled hardwood trees, landed on standing trees, consistent with its being absent from un- and on trees that were standing, but which had been girdled and derstory traps in the present study. Finally, she found that adult N. were dying. Waters (1981) found that adults of both N. a. acumina- m. mucronatus showed no preference for felled or standing trees, tus and X. colonus were caught primarily on felled trees, consistent consistent with their even distribution across trap heights in the pre- with their being most abundant in the understory in the present sent study. The four species of most abundant lamiines varied as to whether they were most abundant in the canopy, the understory, or showed no pattern in their vertical distributions. Lamiine species invariably Table 2. Mean (6 1 SE) number of beetles per trap of the ceramby- feed as adults, often on tender bark or leaves of the same tree species cid species caught in greatest numbers by panel traps baited with as the larval hosts (Hanks and Wang 2016), and thus both sexes a blend of synthetic pheromones (3-hydroxyhexan-2-one, syn-2,3- must be active within the tree canopy for at least part of their lives. hexanediol, 2-methylbutan-1-ol, [E]-fuscumol, [E]-fuscumol acetate, monochamol) or fermenting bait in forests of east-central All four of the abundant lamiine species are highly polyphagous on Illinois hardwoods (Lingafelter 2007), but the condition of larval hosts re- quired by the larvae (i.e., whether hosts are healthy, stressed, or dy- Taxonomy Pheromone Fermenting Friedman’s Q ing) is not known. Of these species, only G. fasciatus was trapped blend bait by Waters (1981) in sufficient numbers to assess host preference. Cerambycinae This species showed no preference for felled or standing trees, as Anelaphus pumilus 12.4 6 3.4 0.04 6 0.04 Q1,56 ¼ 46.0*** would be predicted from its even distribution across trap heights in Downloaded from Eburia quadrigeminata 0.17 6 0.17 2.25 6 0.73 Q1,24 ¼ 13.2*** the present study. Neoclytus a. acuminatus 1.2 6 0.12 0.18 6 0.18 Q ¼ 20.7*** 1,34 Differences among cerambycid species in their preference for Neoclytus m. mucronatus 1.1 6 0.16 0.27 6 0.20 Q1,22 ¼ 8.7**

Neoclytus scutellaris 0.38 6 0.18 0.75 6 0.25 Q1,16 ¼ 1.3 specific forest strata during the present study account for the ab-

Parelaphidion aspersum 1.38 6 0.32 0.25 6 0.16 Q1,16 ¼ 6.8** sence of vertical trends in the total number of beetles captured by Parelaphidion incertum 1.0 6 0.33 0.38 6 0.18 Q1,16 ¼ 2.0 traps, and species richness (also see Dodds 2014). Variation in dis- Phymatodes aereus 5.6 6 2.1 0.13 6 0.13 Q1,16 ¼ 11.4*** persal behavior among cerambycid species within communities is http://jee.oxfordjournals.org/ Phymatodes amoenus 6.1 6 1.6 0.42 6 0.30 Q1,14 ¼ 8.0** likely responsible for the lack of consensus among published stud- Phymatodes lengi 2.4 6 0.40 0.10 6 0.10 Q1,20 ¼ 15.0***

Phymatodes testaceus 1.5 6 0.27 0 Q1,16 ¼ 10.8** ies which have tested the trap height effect, with overall abundance Xylotrechus colonus 1.6 6 0.19 0.37 6 0.12 Q1,82 ¼ 31.5*** and/or species diversity of cerambycids increasing with trap height Lamiinae (Vance et al. 2003, Ulyshen and Hanula 2007, Maguire et al. Aegomorphus modestus 0.93 6 0.18 0.33 6 0.13 Q ¼ 5.8* 1,30 2014), decreasing with trap height (Wermelinger et al. 2007, Astylidius parvus 1.8 6 0.48 0.13 6 0.09 Q1,30 ¼ 16.6***

Graphisurus despectus 1.2 6 0.27 0.50 6 0.20 Q1,24 ¼ 3.4 Dodds 2014), or showing no relationship with trap height Graphisurus fasciatus 1.2 6 0.19 0.44 6 0.17 Q1,36 ¼ 7.8** (Graham et al. 2012). Nevertheless, the present study has corrobo- Lepturges angulatus 1.27 6 0.33 0.18 6 0.12 Q1,22 ¼ 9.0** rated that a thorough assessment of species richness and abun- by guest on August 11, 2016 dance of cerambycids can best be accomplished by positioning Means in bold are signiﬁcantly greater than the other means within species. traps throughout the vertical strata of forests, and by baiting traps Asterisks indicate signiﬁcance of Friedman’s Q:*,P < 0.05; **, P < 0.01; ***, P < 0.001. with a variety of attractants.

Table 3. Mean (6 1 SE) number of beetles per trap of the cerambycid species caught in greatest numbers by panel traps positioned at three heights about the ground in forests of east-central Illinois

Taxonomy Understory (1.5 m) Lower canopy (6 m) Midcanopy (12 m) Friedman’s Q

Cerambycinae

Anelaphus pumilus 5.9 6 1.7 9.1 6 2.0 5.4 6 2.1 Q2,48 ¼ 4.4

Eburia quadrigeminata 0.17 6 0.11 0.17 6 0.11 2.1 6 0.74 Q2,36 ¼ 16.2*** Neoclytus a. acuminatus 0.82 6 0.21 0.29 6 0.11 0.18 6 0.13 Q2,51 ¼ 8.64*

Neoclytus m. mucronatus 0.45 6 0.16 0.27 6 0.14 0.63 6 0.20 Q2,33 ¼ 2.0 Neoclytus scutellaris 0 0.13 6 0.13 1.0 6 0.19 Q2,24 ¼ 15.4*** Parelaphidion aspersum 0.13 6 0.13 0.88 6 0.40 0.63 6 0.18 Q2,24 ¼ 4.2

Parelaphidion incertum 0.13 6 0.13 0.25 6 0.16 1.0 6 0.27 Q2,24 ¼ 8.0* Phymatodes aereus 3.0 6 1.3 2.5 6 1.3 0.25 6 0.16 Q2,24 ¼ 8.8* Phymatodes amoenus 2.1 6 0.80 1.7 6 0.42 2.7 6 1.2 Q2,21 ¼ 0.18

Phymatodes lengi 0.30 6 0.15 0.10 6 0.10 2.1 6 0.48 Q2,30 ¼ 16.6*** Phymatodes testaceus 0.63 6 0.26 0.50 6 0.38 0.38 6 0.18 Q2,24 ¼ 0.80

Xylotrechus colonus 2.5 6 0.50 0.80 6 0.25 0.50 6 0.22 Q2,30 ¼ 10.5** Lamiinae

Aegomorphus modestus 0.27 6 0.12 0.20 6 0.11 0.80 6 0.15 Q2,45 ¼ 10.8**

Astylidius parvus 1.2 6 0.28 0.47 6 0.27 0.27 6 0.12 Q2,45 ¼ 10.3** Graphisurus despectus 0.50 6 0.15 0.58 6 0.20 0.58 6 0.23 Q2,36 ¼ 0.04

Graphisurus fasciatus 0.39 6 0.18 0.67 6 0.14 0.56 6 0.12 Q2,54 ¼ 3.8 Lepturges angulatus 0.45 6 0.21 0.64 6 0.36 0.36 6 0.15 Q2,33 ¼ 0.03

Means in bold are signiﬁcantly greater than other means within species (Dunn–Nemenyi test, P < 0.05). Asterisks indicate overall signiﬁcance of Friedman’s Q: *, P < 0.05; **, P < 0.01; ***, P < 0.001. 1756 Journal of Economic Entomology, 2016, Vol. 109, No. 4

Table 4. Response of the cerambycid species caught in greatest numbers to a blend of synthesized pheromones (from Table 2), and the composition of their pheromones

Taxonomy Attraction to pher. blend Pheromone composition References

Cerambycinae Anelaphus pumilus Yes 3R-ketone Mitchell et al. 2015 Eburia quadrigeminata No Unknown Neoclytus a. acuminatus Yes (2S,3S)-2,3-hexanediol Lacey et al. 2004 Neoclytus m. mucronatus Yes 3R-ketone Lacey et al. 2007 Neoclytus scutellaris No 3R-ketone Ray et al. 2015 Parelaphidion aspersum Yes 3R-ketone, decan-2-one Mitchell et al. 2013 Parelaphidion incertum No Unknown Phymatodes aereus Yes 3R- and 3S-ketone Mitchell et al. 2013 Phymatodes amoenus Yes 3R-ketone, Mitchell et al. 2015 (R)-2-methylbutan-1-ol Phymatodes lengi Yes 3R-ketone, Mitchell et al. 2015 (R)-2-methylbutan-1-ol Phymatodes testaceus Yes Unknown Xylotrechus colonus Yes 3R- and 3S-ketone, Lacey et al. 2009, Mitchell et al. 2015 2,3-hexanediols

Lamiinae Downloaded from Aegomorphus modestus Yes Unknown Astylidius parvus Yes (R)-, (S)-fuscumol, L. R. Meier, unpub. data (R)-fuscumol acetate Graphisurus despectus No Unknown Graphisurus fasciatus Yes Unknown

Lepturges angulatus Yes (R)-, (S)-fuscumol acetate L. R. Meier, unpub. data http://jee.oxfordjournals.org/

All listed pheromone components are contained in the synthetic blend. Chemical abbreviations: 3R-ketone, (R)-3-hydroxyhexan-2-one; 3S-ketone, (S)-3- hydroxyhexan-2-one.

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