Effects of bark beetle pheromones on the attraction of Monochamus alternatus to pine volatiles

Abstract We evaluated the attraction of Monochamus alternatus Hope (Coleoptera: Cerambycidae), Dryocoetes luteus Blandford and Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae) to multiple-funnel traps baited with the pine volatiles, ethanol and (+)-α-pinene and the bark beetle pheromones, ipsenol and ipsdienol. M. alternatus were attracted to traps baited with ethanol and (+)-α-pinene but not those baited with ipsdienol and ipsenol. Ipsdienol and ipsenol decreased catches of M. alternatus in traps baited with ethanol and (+)-α-pinene. Traps baited with either binary combinations of ethanol and (+)-α-pinene or ipsdienol and ipsenol were attractive to D. luteus and O. erosus. The addition of ipsenol and ipsdienol to traps baited with ethanol and (+)-α-pinene synergized attraction of O. erosus but not D. luteus.     

Selective manipulation of predators using pheromones: responses to frontalin and ipsdienol pheromone components of bark beetles in the Great Lakes region

Abstract 1 One proposed approach to improving biological control of bark beetles (Coleoptera: Scolytidae; alt. Curculionidae: Scolytinae) is to manipulate predator movement using semiochemicals. However, selective manipulation is impeded by attraction of both predators and pests to bark beetle pheromones. 2 The primary bark beetle affecting pine plantations in Wisconsin, U.S.A., is the pine engraver, Ips pini (Say). Other herbivores include Ips grandicollis (Eichhoff) and Dryophthorus americanus Bedel (Curculionidae). The predominant predators are the beetles Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae). 3 We conducted field assays using two enantiomeric ratios of ipsdienol, and frontalin plus α‐pinene. Ipsdienol is the principal pheromone component of I. pini, and frontalin is produced by a number of Dendroctonus species. α‐Pinene is a host monoterpene commonly incorporated into commercial frontalin lures. 4 Thanasimus dubius was attracted to frontalin plus α‐pinene, and also to racemic ipsdienol. By contrast, I. pini was attracted to racemic ipsdienol, but showed no attraction to frontalin plus α‐pinene. Platysoma cylindrica was attracted to 97%‐(–)‐ipsdienol and, to a lesser extent, racemic ipsdienol, but not to frontalin plus α‐pinene. Ips grandicollis was attracted to frontalin plus α‐pinene but not to ipsdienol. Dryophthorus americanus was attracted to both ipsdienol and frontalin plus α‐pinene. 5 This ability to selectively attract the predator T. dubius without attracting the principal bark beetle in the system, I. pini, provides new opportunities for research into augmentative biological control and basic population dynamics. Moreover, the attraction of T. dubius, but not P. cylindrica, to frontalin plus α‐pinene creates opportunities for selective manipulation of just one predator. 6 Patterns of attraction by predators and bark beetles to these compounds appear to reflect various degrees of geographical and host tree overlap with several pheromone‐producing species.     

Variation in enantiospecific attraction of Ips avulsus (Coleoptera: Curculionidae) to the pheromone ipsdienol in Georgia

In 2006, we tested the responses of the small southern pine engraver, Ips avulsus (Eichhoff) (Coleoptera: Curculionidae), to multiple-funnel traps baited with (+)-, (-)-, and (+/-)-ipsdienol. Three experiments were conducted in Georgia with all traps co-baited with one of the following lure combinations, respectively: experiment 1, ipsenol; experiment 2, lanierone and (-)-alpha-pinene; and experiment 3, ipsenol, lanierone, and (-)-alpha-pinene. Ipsdienol and lanierone are aggregation pheromones used by I. avulsus, whereas ipsenol is an aggregation pheromone used by the eastern fivespined ips, Ips grandicollis (Eichhoff), a sympatric species also breeding in pines. In all experiments, the highest catches of I. avulsus were obtained in traps baited with (+/-)-ipsdienol. The relative attractiveness of (+)-ipsdienol varied across the three experiments. When traps were co-baited with ipsenol, lanierone, and (-)-alpha-pinene, catches of I. avulsus in traps baited with (+)-ipsdienol were high and no different from those in traps baited with (+/-)-ipsdienol. When traps were co-baited with lanierone and (-)-alpha-pinene, catches in traps baited with (+)-ipsdienol were lower than those in traps baited with (+/-)-ipsdienol by approximately 40%, whereas catches were reduced by approximately 95% when traps were co-baited with ipsenol alone. We hypothesize that the observed variation in enantiospecific responses of I. avulsus to ipsdienol may be explained by variation in semiochemical context (i.e., different co-baits among the experiments) or seasonal variation.     

Bark beetle pheromones and pine volatiles: attractant kairomone lure blend for longhorn beetles (Cerambycidae) in pine stands of the southeastern United States

In 2006, we examined the flight responses of 43 species of longhorn beetles (Coleoptera: Cerambycidae) to multiple-funnel traps baited with binary lure blends of (1) ipsenol + ipsdienol, (2) ethanol + alpha-pinene, and a quaternary lure blend of (3) ipsenol + ipsdienol + ethanol + alpha-pinene in the southeastern United States. In addition, we monitored responses of Buprestidae, Elateridae, and Curculionidae commonly associated with pine longhorn beetles. Field trials were conducted in mature pine (Pinus pp.) stands in Florida, Georgia, Louisiana, and Virginia. The following species preferred traps baited with the quaternary blend over those baited with ethanol + alpha-pinene: Acanthocinus nodosus (F.), Acanthocinus obsoletus (Olivier), Astylopsis arcuata (LeConte), Astylopsis sexguttata (Say), Monochamus scutellatus (Say), Monochamus titillator (F.) complex, Rhagium inquisitor (L.) (Cerambycidae), Buprestis consularis Gory, Buprestis lineata F. (Buprestidae), Ips avulsus (Eichhoff), Ips calligraphus (Germar), Ips grandicollis (Eichhoff), Orthotomicus caelatus (Eichhoff), and Gnathotrichus materiarus (Fitch) (Curculionidae). The addition ofipsenol and ipsdienol had no effect on catches of 17 other species of bark and wood boring beetles in traps baited with ethanol and a-pinene. Ethanol + alpha-pinene interrupted the attraction of Ips avulsus, I. grandicollis, and Pityophthorus Eichhoff spp. (but not I. calligraphus) (Curculionidae) to traps baited with ipsenol + ipsdienol. Our results support the use of traps baited with a quaternary blend of ipsenol + ipsdienol + ethanol + alpha-pinene for common saproxylic beetles in pine forests of the southeastern United States.     

Role of ipsdienol, ipsenol, and cis-verbenol in chemical ecology of Ips avulsus, Ips calligraphus, and Ips grandicollis (Coleoptera: Curculionidae: Scolytinae)

Stressed or damaged pine (Pinus sp.) trees in the southeastern United States are often colonized simultaneously by three southern Ips species (Coleoptera: Curculionidae: Scolytinae): small southern pine engraver, Ips avulsus (Eichhoff); sixspined ips, Ips calligraphus (Germar); and eastern fivespined ips, Ips grandicollis (Eichhoff). All three species mediate colonization of host material with volatile pheromones. All of the southern Ips produce cis-verbenol, and either ipsdienol or ipsenol, and electrophysiological studies have demonstrated that all three southern Ips are able to detect all three compounds. This study examined the role of ipsdienol, ipsenol, and cis-verbenol in the chemical ecology of the southern Ips in Georgia and Louisiana. The most attractive blends of pheromones, with the fewest number of components, were ipsdienol plus ipsenol for I. avulsus, cis-verbenol plus ipsdienol for I. calligraphus, and either cis-verbenol plus ipsenol or ipsdienol plus ipsenol for I. grandicollis. Cross-attraction of I. grandicollis to the pheromone blend most attractive to I. avulsus was observed. Although the presence of heterospecific pheromone reduced the catches of all three species (i.e., the tertiary blend captured fewer beetles than the most attractive binary blends) in both states (significantly in two cases), high numbers of all three species were still captured in traps baited with all three compounds. These results suggest that the pheromones cis-verbenol, ipsdienol, and ipsenol can be combined for monitoring all three species of the southern Ips simultaneously.     

Effect of different semiochemicals blends on spruce bark beetle,Ips typographus (Coleoptera: Scolytidae)

The spruce bark beetle, Ips typographus, is a recent new introduction to the Qilian Mountains of China. An outbreak of these beetles has infested over 0.03 million hectares of spruce forests in this area. Although primary attraction to volatiles has been clearly demonstrated for I. typographus, the existence and role of attraction to insect‐produced pheromones have been widely debated. Currently, commercial lures for I. typographus include only the volatiles ipsdienol, cis‐verbenol, trans‐verbenol, 2‐methyl‐3‐buten‐2‐ol and 2‐phenylethanol in Europe. Several potential pheromone candidates have been identified for I. typographus. Our GC–MS and GC–FID analyses volatiles from hindgut extracts of I. typographus in different attack phases demonstrated that the 2‐methyl‐3‐buten‐2‐ol, ipsdienol, cis‐verbenol and trans‐verbenol as major hindgut components, and ipsenol, 2‐phenylethanol, trans‐ myrtenol and verbenone as minor components. We tested various combinations of semiochemical candidates, to determine an optimal blend. Our results suggest that addition of 2‐methyl‐3‐buten‐2‐ol to either ipsenol alone, or to blends of ipsenol and other semiochemical candidates, significantly enhanced attraction of I. typographus. Therefore, a simple lure consisting of ipsenol and 2‐methyl‐3‐buten‐2‐ol would be an optimal blend of I. typographus in the Qilian Mountains, China. We conclude that this optimal semiochemical blend may provide an effective biological pest control method for use in forest ecosystem against I. typographus.     

Aggregation pheromone of the Oriental spruce engraver Pseudips orientalis

• 1 Volatiles from the hindgut extracts of males of the Oriental spruce engraver Pseudips orientalis (Wood & Yin) (Coleoptera: Curculionidae, Scolytinae) of different phases of gallery development were analyzed by gas chromatography‐mass spectrometry‐flame ionization detection (GC‐MS/FID) with both polar and enantioselective columns.
• 2 GC‐MS/FID analyses showed that unmated males or males mated with one female produced approximately 95%‐(?)‐ipsenol and (?)‐cis‐verbenol as major components, as well as (?)‐trans‐verbenol, myrtenol, approximately 70%‐(+)‐ipsdienol and (?)‐verbenone as minor or trace components. The release of these male‐produced compounds was confirmed by GC analysis of an aeration sample of a P. orientalis‐infested spruce log. Mating reduced production of the male‐specific hindgut volatiles.
• 3 A field‐trapping bioassay in Qinghai, China, showed that a ternary blend containing two major components, 97%‐(?)‐ipsenol (i.e. close to naturally produced enantiomeric ratio) and (?)‐cis‐verbenol, plus a minor component (?)‐trans‐verbenol, caught significantly more P. orientalis beetles (♂: ♀ = 1: 2.7) compared with the unbaited control. Subtraction of (?)‐trans‐verbenol from the active ternary blend had no significant effect on trap catches. The addition of (±)‐ipsdienol (at 0.2 mg/day release) to the active ternary or binary blends significantly interrupted their trap catches. Replacing 97%‐(?)‐ipsenol with (±)‐ipsenol in the ternary blend significantly reduced trap catches to a level that was no different from the blank control.
• 4 Pseudips species were sister to all other Ipini genera in a phylogeny reconstructed with mitochondrial cytochrome oxidase I DNA data for 51 Ipini and outgroup species.
• 5 The results obtained suggest that the two major components, 95%‐(?)‐ipsenol and (?)‐cis‐verbenol (at approximately 4–5 : 1), produced by unmated fed males, are probably the primary aggregation pheromone components for P. orientalis. In light of the phylogeny, the use of terpenoid semiochemicals as pheromones probably occurred early in the evolution of Ipini and these semiochemical blends were subsequently modified in the process of speciation.

     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Attractive responses by Monochamus galloprovincialis (Col., Cerambycidae) to host and bark beetle semiochemicals

Abstract:  The pine sawyer Monochamus galloprovincialis is the European vector of the recently introduced pine wood nematode. This nematode is the causal organism of pine wilt disease, a serious tree killer in East Asia. Efficacious baits and traps to monitor and control this beetle are now required. The effect of bark beetle ( Ips spp.) pheromone components, released individually (ipsenol) or in blends (ipsenol, ipsdienol, cis -verbenol and methyl-butenol), together with host volatiles (turpentine or α -pinene and ethanol) on M. galloprovincialis trap catches has been studied in Spain. A kairomonal response by male and female of M. galloprovincialis to Ips semiochemicals was found. Beetles were more attracted to host blends supplemented with bark beetle pheromones than to host volatiles alone. Ipsenol alone was attractive to pine sawyers, and was synergistic with α -pinene and ethanol. The full blend of the four Ips semiochemicals and the host compounds was highly attractive. Multiple-funnel traps were as effective as black cross-vane traps in capturing this insect when the escape of trapped beetles was prevented. Trapping of non-target bark beetle predators was also evaluated. The trogossitid Temnochila coerulea and clerid Thanasimus formicarius were kairomonally attracted to and killed in traps baited with bark beetle pheromones. These results suggest that effective monitoring of M. galloprovincialis would be possible by baiting any of these traps with host volatiles and Ips semiochemicals, but reduction of the lure components and trap modification to minimize impact on predators should be considered.     

Geographic variation in prey preference in bark beetle predators

Abstract 1. Bark beetles and their predators are useful systems for addressing questions concerning diet breadth and prey preference in arthropod natural enemies. These predators use bark beetle pheromones to locate their prey, and the response to different pheromones is a measure of prey preference. 2. Trapping experiments were conducted to examine geographic variation in the response to prey pheromones by two bark beetle predators, Thanasimus dubius and Temnochila virescens. The experiments used pheromones for several Dendroctonus and Ips prey species (frontalin, ipsdienol, and ipsenol) and manipulated visual cues involved in prey location (black vs. white traps). The study sites included regions where the frontalin‐emitter Dendroctonus frontalis was in outbreak vs. endemic or absent. 3. There was significant geographic variation in pheromone preference for T. dubius. This predator strongly preferred a pheromone (frontalin) associated with D. frontalis at outbreak sites, while preference was more even at endemic and absent sites. No geographic variation was found in the response by T. virescens. White traps caught fewer insects than black traps for both predators, suggesting that visual cues are also important in prey location. 4. The overall pattern for T. dubius is consistent with switching or optimal foraging theory, assuming D. frontalis is a higher quality prey than Ips. The two predator species partition the prey pheromones in areas where D. frontalis is abundant, possibly to minimise competition and intraguild predation.     

Traps and attractants for wood-boring insects in ponderosa pine stands in the Black Hills, South Dakota

Recent large-scale wildfires have increased populations of wood-boring insects in the Black Hills of South Dakota. Because little is known about possible impacts of wood-boring insects in the Black Hills, land managers are interested in developing monitoring techniques such as flight trapping with semiochemical baits. Two trap designs and four semiochemical attractants were tested in a recently burned ponderosa pine, Pinus ponderosa Dougl. ex Laws., forest in the Black Hills. Modified panel and funnel traps were tested in combination with the attractants, which included a woodborer standard (ethanol and alpha-pinene), standard plus 3-carene, standard plus ipsenol, and standard plus ipsdienol. We found that funnel traps were equally efficient or more efficient in capturing wood-boring insects than modified panel traps. Trap catches of cerambycids increased when we added the Ips spp. pheromone components (ipsenol or ipsdienol) or the host monoterpene (3-carene) to the woodborer standard. During the summers of 2003 and 2004, 18 cerambycid, 14 buprestid, and five siricid species were collected. One species of cerambycid, Monochamus clamator (LeConte), composed 49 and 40% of the 2003 and 2004 trap catches, respectively. Two other cerambycids, Acanthocinus obliquus (LeConte) and Acmaeops proteus (Kirby), also were frequently collected. Flight trap data indicated that some species were present throughout the summer, whereas others were caught only at the beginning or end of the summer.     

Evaluation of attractants and traps for monitoring small banded pine weevil Pissodes castaneus

This study aimed to develop a semiochemical‐baited trapping system to monitor the populations of small banded pine weevil, Pissodes castaneus, a serious pest in Pinus sylvestris young stands that are weakened by biotic and abiotic factors. The scope of the work included the development of a dispenser for compounds (ethanol and α‐pinene) emitted by P. sylvestris and the pheromones of P. castaneus: grandisol and grandisal. Additionally, the effectiveness of beetle catches in different types of traps (unitrap, cross‐unitrap and long and short pipe traps) baited with a dispenser was assessed. The olfactometric studies showed that most of the newly hatched beetles that had not fed were attracted by a mixture of grandisol and grandisal. However, in the group of feeding beetles, half were attracted by a mixture of ethanol and α‐pinene. These results indicated that both pheromones and α‐pinene plus ethanol should be useful for capturing P. castaneus beetles. In the field trials, the highest efficiency was found in baited unitraps that caught up to several hundred P. castaneus beetles, while the baited cross‐unitraps caught up to a few dozen beetles. No insects were found in either type of baited pipe trap or in any of the unbaited control traps. The baited unitraps and cross‐unitraps also collected, with varied intensity, Hylobius abietis beetles, a serious pest of reforestations. These results indicate the possibility of using a unitrap baited with a 4‐component attractant for monitoring P. castaneus in integrated pest management for the protection of young forests.     

Outdoor flight activity and immigration of Rhyzopertha dominica into seed wheat warehouses

The flight activity of lesser grain borer, Rhyzopertha dominica F. (Coleoptera: Bostrichidae), was monitored at two Foundation seed wheat warehouses during the 2003 and 2004 field seasons, using pheromone‐baited Lindgren funnel traps positioned indoors and outdoors. General stored‐product insect activity was also monitored using unbaited sticky traps positioned inside the warehouses around overhead doors. Pheromone‐baited traps were useful for monitoring R. dominica activity, however insect captures decreased when lures were not changed weekly. Flight peaks were documented in early May and again from September through October, and insect captures inside warehouses correlated with timing of outdoor captures. Multiple regression analyses showed that slightly more than half of the variability in R. dominica captures could be explained by mean ambient air temperature and wind speed during the 2 h preceding sunset. Stored‐product Coleoptera captured on unbaited glue boards around overhead doors included Ahasverus advena, Cryptolestes ferrugineus, R. dominica, Sitophilus oryzae, Tribolium castaneum, Trogoderma variabile, and Typhaea stercorea. Door gaskets significantly reduced the number of insect captures on glue boards placed around the overhead doors, and generally restricted their entry to ground level. These studies demonstrated that outdoor pheromone‐baited traps are effective monitoring tools for determining when grain‐handling facilities are most susceptible to infestation and that exclusion may be an effective component of a pest management program.     

Field evaluation on the synergistic attractiveness of 2‐(1‐undecyloxy)‐1‐ethanol and ipsenol to Monochamus saltuarius

To develop an optimal attractant for Monochamus saltuarius (Gebler) (Coleoptera: Cerambycidae), the synergistic effects of a few potential attractants (ethanol and α‐pinene as host‐plant volatiles, and ipsenol and ipsdienol as bark beetle pheromones) were tested in a pine forest combined with 2‐(1‐undecyloxy)‐1‐ethanol (monochamol), the aggregation pheromone of Monochamus species, for two consecutive years, 2014 and 2015. Total number of catches was 65 and 33 in 2014 and 2015, respectively. Ethanol or ethanol + monochamol (a base blend) were not attractive to M. saltuarius with no difference from the control. Addition of α‐pinene and ipsdienol to the base blend did not significantly increase catches. However, ipsenol was significantly synergistic to the base blend in attracting M. saltuarius in 2014, and the blend (ipsenol + base blend) attracted meaningfully higher numbers of M. saltuarius in 2015. Our study illustrates the potential for monochamol and ipsenol baits for monitoring and trapping of M. saltuarius in the field.     

Combining pheromone and kairomones for effective trapping of the pine sawyer beetle Monochamus galloprovincialis

Monochamus galloprovincialis (Olivier) (Coleoptera: Cerambycidae) is a secondary wood borer that acquired primordial importance since it was identified as the European vector of Bursaphelenchus xylophilus, the causal agent of pine wilt disease (PWD). An effective trapping method is needed as a tool for managing this insect vector and allowing early detection of nematode transportation. Among effective attractants identified in recent years are the specific M. galloprovincialis aggregation pheromone, host pine kairomones such as α‐pinene and bark beetle kairomones like ipsenol and methyl‐butenol. The main objective of this study was to optimize the combination of these volatiles to improve lure attractiveness and specificity. Based on ten complementary field experiments, we found a pheromone dose‐response of trap catches. The best combination of attractants was the aggregation pheromone plus two bark beetle kairomones, ipsenol and methyl‐butenol. Addition of pine terpenes, such as α‐pinene, did not significantly improve M. galloprovincialis trap capture, but did increase catch of non target species, including natural enemies. The use of pine terpenes would be advisable only if priorizing to maximize removal of vectors. While this research has lead to the development a new, highly attractive commercial lure for mature pine sawyers, none of the tested blends were successful in attracting immature pine sawyer adults. Further investigation is needed to develop attractants for these beetles.     

Attraction of southern pine engravers and associated bark beetles (Coleoptera: Scolytidae) to ipsenol, ipsdienol, and lanierone in southeastern United States

We determined the response of the small southern pine engraver, Ips avulsus (Eichhoff); eastern fivespined ips, Ips grandicollis (Eichhoff); sixspined ips, Ips calligraphus (Germar); and pine engraver, Ips pini (Say) to the pheromones (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone in the southeastern United States. Catches of I. avulsus and I. grandicollis to baited multiple-funnel traps were increased by (+/-)-ipsenol and (+/-)-ipsdienol in Florida, Georgia, Louisiana, and North Carolina. In all four localities, the highest numbers of I. avulsus were caught in traps baited with the combination of (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone. In Florida, the highest numbers of I. grandicollis were captured in traps baited with the combination of (+/-)-ipsenol and (+/-)-ipsdienol (with or without lanierone). In the remaining three localities, the largest catches of I. grandicollis occurred in traps baited with (+/-)-ipsenol alone or the combination of (+/-)-ipsenol and (+/-)-ipsdienol (with or without lanierone). (+/-)-Ipsdienol was the only consistent attractant for I. calligraphus and I. pini. Attraction of I. pini in North Carolina to (+/-)-ipsdienol-baited traps was synergized by lanierone but interrupted with (+/-)-ipsenol. The interruptive effect of (+/-)-ipsenol on attraction of I. pini to (+/-)-ipsdienol was negated by lanierone. (+/-)-Ipsdienol was attractive to black turpentine beetle, Dendroctonus terebrans (Olivier), in Florida but not North Carolina, whereas (+/-)-ipsdienol was attractive to I. calligraphus in Louisiana, Georgia, and Florida. Both (+/-)-ipsenol and (+/-)-ipsdienol affected catches of Gnathotrichus materiarus (Fitch) in North Carolina. Trap catches of Hylurgops rugipennis pinifex (Fitch), Hylastes salebrosus Eichhoff, and Hylastes tenuis Eichhoff were unaffected by the pheromone treatments. The combination of (+/-)-ipsenol, (+/-)-ipsdienol, and lanierone may be a cost-effective general lure for I. avulsus, I. grandicollis, and I. pini.     

Field responses of the Asian larch bark beetle,Ips subelongatus,to potential aggregation pheromone components: disparity between two populations in northeastern China

Abstract Behavioral responses of the Asian larch bark beetle, Ips subelongatus Motsch. to three potential aggregation pheromone components, ipsenol (racemic or [?]‐enantiomer), ipsdienol (racemic or [+]‐enantiomer) and 3‐methyl‐3‐buten‐1‐ol, were tested using partial or full factorial experimental designs in two provinces (Inner Mongolia and Jilin) of northeastern China. Our field bioassays in Inner Mongolia (Larix principis‐rupprechtii Mayr. plantation) clearly showed that ipsenol, either racemic or 97%‐(?)‐enantiomer, was the only compound that significantly attracted both sexes of I. subelongatus, while all other compounds (singly or in combinations) were unattractive. There were no two‐ or three‐way synergistic interactions. However, in Jilin Province (L. gmelini[Rupr.] Rupr. Plantation), all the individual compounds tested were inactive, except a very weak activity by 97%‐(?)‐ipsenol in 2004 when the beetle population was very high. While a combination of ipsenol and ipsdienol (racemates or enantiomerically pure natural enantiomers) showed a significant attraction for both sexes of I. subelongatus, indicating a two‐way synergistic interaction between these two major components, addition of 3‐methyl‐3‐buten‐1‐ol to these active binary blend(s) did not have any effects on trap catches, suggesting that ipsenol and ipsdienol are the synergistic aggregation pheromone components of I. subelongatus in Jilin Province. It seems that 97%‐(?)‐ipsenol in Inner Mongolia or the binary blend of 97%‐(?)‐ipsenol and 97%‐(+)‐ipsdienol in Jilin Province are superior to their corresponding racemates, which might be due either to weak inhibitory effects of the antipode enantiomers or to reduced release rates of the active natural enantiomer(s) in the racemate(s). Our current bioassay results suggest that there is a strong geographical variation in aggregation pheromone response of I. subelongatus in northeastern China. Future research on the pheromone production and response of I. subelongatus from different regions in Northeast Asia will surely improve our understanding of the dynamic aggregation pheromone system of this economically important forest pest insect.