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.     

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.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Field response of non‐target beetles to Ips sexdentatus aggregation pheromone and pine volatiles

Although mass trapping cannot be a definitive control measure, it is one of the few ones available to contain the destruction of millions of cubic metres of conifer forests perpetrated every year worldwide by bark beetles. However, using bark beetle aggregation pheromones during both monitoring and control programs may negatively affect other saproxylic insects. The aim of this study was to describe the response of both Ips sexdentatus and its saproxylic beetle associates, especially predators, to traps baited with a commercial blend of I. sexdentatus aggregation pheromone. Furthermore, the usefulness of adding pine volatiles, such as (?)‐α‐pinene and ethanol, to the pheromone was discussed. The commercial blend proved to be attractive to I. sexdentatus adults, both when used alone and together with pine volatiles. Pheromone attractiveness, however, was lessened by the addition of the volatiles. The pheromone blend proved to be attractive to Thanasimus formicarius, as well as to other predator species. Overall, although during our study, traps baited only with (?)‐α‐pinene and ethanol attracted some predator specimens, I. sexdentatus pheromone traps were more attractive. Our study confirms that calendar differences in flight activity between the bark beetle and its predators are substantial; therefore, they should be taken into account when planning control measures. According to our data, the commercial blend of I. sexdentatus pheromone seems to be the most effective, among the baits used, in catching I. sexdentatus adults, while reducing the impact on T. formicarius.     

Response of the whitespotted sawyer beetle, Monochamus s. scutellatus, and associated woodborers to pheromones of some Ips and Dendroctonus bark beetles

The response of whitespotted sawyer beetle, Monochamus s. scutellatus, to pheromones of the bark beetles, Dendroctonus rufipennis, Ips pini, Ips perturbatus and Ips latidens, and α‐pinene was investigated with field‐trapping experiments. Traps baited with ipsenol caught significantly more M. s. scutellatus than unbaited traps, whereas the other compounds (ipsdienol, ipsdienol plus lanierone, ipsdienol plus cis‐verbenol or frontalin) did not. Combining α‐pinene with ipsdienol, ipsdienol plus lanierone, ipsdienol plus cis‐verbenol or with frontalin did not increase captures of M. s. scutellatus above those of α‐pinene alone, whereas the combination of α‐pinene with ipsenol did. When α‐pinene was combined with ipsdienol or frontalin, trap captures of Monochamus mutator were significantly higher than unbaited traps or traps baited with frontalin but were not higher than traps baited with α‐pinene. The combination of ipsenol and α‐pinene was significantly more attractive to Monochamus notatus than unbaited traps; however, traps containing either ipsenol or α‐pinene were as attractive as the combination. None of the species of Buprestidae (Buprestis maculativentris and Chalcophora virginiensis) responded significantly to any of the treatments.     

Attraction of the southern pine beetle, Dendroctonus frontalis, to pheromone components of the western pine beetle, Dendroctonus brevicomis (Coleoptera: Curculionidae: Scolytinae), in an allopatric zone

Subtle differences in pheromone components of sympatric species should be attractive only to the producing species and unattractive or repellent to the nonproducing species, and thereby maintain reproductive isolation and reduce competition between species. Bark beetles Dendroctonus brevicomis and D. frontalis (Coleoptera: Curculionidae) are known to have common pheromone components, except for exo-brevicomin, which is produced by D. brevicomis. We predicted that D. frontalis would not respond to exo-brevicomin outside of the zone of sympatry with D. brevicomis. We conducted a field experiment to determine the effect of exo-brevicomin on attraction of D. frontalis and associated species in Mississippi. We determined whether D. frontalis pheromone production differed inside and outside the sympatric zone and compared the pheromone profiles with D. brevicomis within the sympatric zone. Trapping studies revealed that D. frontalis can perceive and respond positively to exo-brevicomin, an aggregation pheromone of a sympatric congener (D. brevicomis), at locations hundreds of kilometers outside the sympatric zone. Qualitative pheromone profiles showed that both species emit similar pheromone components: frontalin, endo-brevicomin, exo-brevicomin, trans-verbenol, verbenone, and myrtenol. Although not previously reported, D. frontalis males from Arizona produced exo-brevicomin. The predator Thanasimus dubius did not discriminate traps baited with exo-brevicomin and was most attracted to traps with frontalin. Hylastes beetles were significantly attracted to traps baited with exo-brevicomin in combination with other compounds. Our results raise new practical and evolutionary questions on the role of exo-brevicomin in the behavioral ecology of D. frontalis. The addition of exo-brevicomin to the current lure might increase the efficiency of trapping programs in the southeastern United States.     

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.     

Specialization of the olfactory receptor cells in the bark beetleIps typographus and its predatorThanasimus formicarius to bark beetle pheromones and host tree volatiles

Summary Olfactory receptor cells of the spruce bark beetle,Ips typographus, and its predator, the clerid beetleThanasimus formicarius, were studied using electrophysiological techniques. Recordings were made of nerve impulses from single cells and of the summated receptor potential (electroantennogram).Information from bark beetle pheromones and host volatiles is detected by separate olfactory receptor cells inI. typographus. Those which detected bark beetle pheromones responded to only one key substance. Some receptor cells which responded to spruce bark volatiles were strongly activated by one of the synthetic host compounds tested. However, too few host compounds were tested to reach definite conclusions about the specialization of host odour cells. T. formicarius has evolved olfactory receptor cells for bark beetle pheromones. These have similar specificities (specialist types) to those of the bark beetles. Furthermore, the predator has olfactory receptor cells for many bark beetle pheromones. This indicates thatT. formicarius is able to detect and discriminate between many bark beetle species. No significant differences were found between prey and predator cells which responded to host volatiles.     

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.     

Predation and bark beetle dynamics

Bark beetle populations may undergo dramatic fluctuations and are often important pests in coniferous forests. Their dynamics are thought to be primarily driven by factors affecting the resistance of the host tree to attack, i.e., bottom-up forces, while natural enemies are usually assigned a minor role in these systems. I present behavioral experiments that suggest that the clerid beetle Thanasimus dubius may be an important source of mortality for the bark beetle Dendroctonus frontalis during attack of the host tree, and determine the nature of the functional response of T. dubius under conditions close to natural. I also examine the numerical response of T. dubius to large-scale fluctuations in D. frontalis density, and the relationship between bark beetle population trends and predator density, and find that beetle populations tend to decline when predator densities are high. Combined with the effects of clerid larvae on bark beetle broods, these results suggest that top-down forces generated by natural enemies could also be an important component of bark beetle dynamics. The implications of these results for bark beetle dynamics are discussed in relation to the prolonged life-cycle of clerid beetles. Received: 23 January 1997 / Accepted: 5 April 1997     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.     

The enemy of my enemy is still my enemy: competitors add to predator load of a tree‐killing bark beetle

1 The mountain pine beetle Dendroctonus ponderosae is a major tree‐killing bark beetle in North America. We evaluated how the subsequent arrival of a competing bark beetle Ips pini influences the arrival of predators and their impact on both species. 2 The predators Temnochila chlorodia and Enoclerus sphegeus were strongly attracted to pheromones of D. ponderosae. By contrast, Enoclerus lecontei was mostly attracted to I. pini pheromones. The host compound myrcene synergized attraction of both D. ponderosae and E. sphegeus to the pheromone of D. ponderosae. However, it inhibited attraction of both I. pini and E. lecontei to I. pini’s pheromone. 3 Dendroctonus ponderosae were more attracted to trees than logs treated with its pheromones, whereas I. pini were more attracted to logs than trees treated with its pheromones. Some 78% of T. chlorodia were captured at hosts baited with D. ponderosae pheromones, whereas 83% of E. lecontei were captured at hosts baited with I. pini pheromones. We characterized the sequence of arrival to live trees baited with pheromones of D. ponderosae as: D. ponderosae, T. chlorodia, E. sphegeus, I. pini, E. lecontei. 4 Various combinations of I. pini and predators were added to logs colonized by D. ponderosae in the above sequence of arrival observed in live trees baited with D. ponderosae aggregation pheromones. Ips pini reduced D. ponderosae adult brood production. However, the combination of I. pini and E. lecontei did not raise D. ponderosae brood production above that observed with only I. pini present. Similarly, the combination of I. pini and T. chlorodia did not reduce D. ponderosae brood production below that observed with I. pini alone. By contrast, the combination of I. pini, T. chlorodia and E. lecontei caused more brood loss to D. ponderosae than I. pini alone. 5 Enoclerus lecontei did not reduce brood production by T. chlorodia, whereas T. chlorodia substantially reduced brood production by E. lecontei. 6 Secondary bark beetles that exploit the resource created by primary tree‐killing species exert negative effects through both competition and increased predator load. Implications to the population dynamics, ecology and evolution of tree‐killing bark beetles are discussed.     

Synergistic effects of α-pinene and exo-brevicomin on pine bark beetles and associated insects in Arizona

The southern pine beetle (Dendroctonus frontalis) and western pine beetle (Dendroctonus brevicomis) cause significant mortality to pines in the southern and western United States. The effectiveness of commercial lures at capturing these bark beetles in Arizona has not been tested and may vary from other regions of their distribution. We conducted experiments using baited Lindgren funnel traps to investigate (i) if D. frontalis is more attracted to the standard commercial lure for D. brevicomis (frontalin + exo‐brevicomin + myrcene) than the D. frontalis lure (frontalin + terpene blend), (ii) whether replacement of myrcene with α‐pinene changes trap catches of Dendroctonus and associated insects, and (iii) whether the attraction to these lures varies across the geographical range of ponderosa pine forests throughout Arizona. In 2005, we tested various combinations of frontalin, exo‐brevicomin, myrcene and α‐pinene to D. frontalis, D. brevicomis and associated species. Dendroctonus frontalis, D. brevicomis and the predator Temnochila chlorodia were most attracted to lures with exo‐brevicomin. The replacement of the myrcene component with α‐pinene in the D. brevicomis lure resulted in the capture of twice as many bark beetles and Elacatis beetles. However, T. chlorodia did not differentiate between monoterpenes. In 2006, traps were set up in 11 locations around Arizona to test the relative attraction of lure combinations. In 9 out 11 locations, the D. brevicomis lure with α‐pinene was more attractive than the lure with myrcene or a terpene blend. These results suggest that the D. brevicomis lure with α‐pinene rather than myrcene is more effective lure to capture D. brevicomis and D. frontalis in Arizona. However, geographical variation in attractiveness to lures is evident even within this region of the beetles’ distributions. Differential attraction of Dendroctonus and their predators to these lures suggests potential use in field trapping and control programmes.     

Can chemical communication be cryptic? Adaptations by herbivores to natural enemies exploiting prey semiochemistry

Predators and parasites commonly use chemical cues associated with herbivore feeding and reproduction to locate prey. However, we currently know little about mechanisms by which herbivores may avoid such natural enemies. Pheromones are crucial to many aspects of herbivore life history, so radical alterations of these compounds could be disadvantageous despite their exploitation by predators. Instead, minor modifications in pheromone chemistry may facilitate partial escape while maintaining intraspecific functionality. We tested this hypothesis using Ips pini, an endophytic beetle that develops in the phloem tissue of pine trees. Its predominant predators in the Great Lakes region of North America are Thanasimus dubius and Platysoma cylindrica, both of which are highly attracted to I. pini’s pheromones. However, there are significant disparities between prey and predator behaviors that relate to nuances of pheromone chemistry. Thanasimus dubius is most attracted to the (+) stereoisomer of ipsdienol, and P. cylindrica is most attracted to the (−) form; Ips pini prefers racemic mixtures intermediate between each predator’s preferences. Further, a component that is inactive by itself, lanierone, greatly synergizes the attraction of I. pini to ipsdienol, but has a weak or no effect on its predators. A temporal component adds to this behavioral disparity: lanierone is most important in the communication of I. pini during periods when its predators are most abundant. The difficulties involved in tracking prey are further compounded by spatial and temporal variation in prey signaling on a local scale. For example, the preferences of I. pini vary significantly among sites only 50 km apart. This chemical crypsis is analogous to morphological forms of camouflage, such as color and mimicry, that are widely recognized as evasive adaptations against visually searching predators. Presumably these relationships are dynamic, with predators and prey shifting responses in microevolutionary time. However, several factors may delay predator counter adaptations. The most important appears to be the availability of alternate prey, specifically I. grandicollis, whose pheromone ipsenol is highly attractive to the above predators but not cross-attractive with I. pini. Consistent with this view, the specialist parasitoid, Tomicobia tibialis, has behavioral preferences for pheromone components that closely correspond with those of I. pini. These results are discussed in terms of population dynamics and coevolutionary theory. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Kairomonal Responses of Natural Enemies and Associates of the Southern Ips (Coleoptera: Curculionidae: Scolytinae) to Ipsdienol, Ipsenol and Cis-Verbenol

Bark beetle infested pines are an ephemeral habitat utilized by a diverse assemblage of insects. Although many bark beetle insect associates have little or no measurable impact on bark beetle brood production, some reduce brood production by either competing with brood for the limited phloem tissue or by feeding on brood. Several studies have observed synchrony between the colonization of hosts by bark beetles and the arrival of insect associates. Some insect associates mediate synchrony with bark beetle mass attacks with kairomonal responses to bark beetle aggregation pheromones. The objectives of this study were to document the community of Coleoptera associated with the southern Ips (Ips avulsus, Ips calligraphus and Ips grandicollis) and to test the hypothesis that synchrony of insect associates with the southern Ips is mediated by kairomonal responses to aggregation pheromones. A large community of Coleoptera (109 species) was recorded from traps baited with southern Ips pheromones. A significant treatment effect was observed for the guilds of meristem feeders, natural enemies and woodborers. The southern Ips pheromone ipsenol was broadly attractive to meristem feeders, natural enemies and woodborers and in general blends were more attractive than individual compounds. These results demonstrate that a diverse community of Coleoptera is associated with the southern Ips and that several members of this community facilitate synchrony with kairomonal responses to southern Ips aggregation pheromones.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Chiral escape of bark beetles from predators responding to a bark beetle pheromone

Summary Two species of predatory beetles that locate their prey, Ips pini, by responding to its aggregation pheromone have different chiral preferences to ispdienol than does the herbivore. This suggests that chiral disparity may provide some escape for bark beetles from predation, and that geographic variation in herbivore communication systems may be partially due to predator — imposed selection pressures. These results also suggest ways in which the semiochemical and biological control of North America's most damaging group of forest insects can be improved.     

Primary attraction and kairomonal host discrimination in three species of Dendroctonus (Coleoptera: Scolytidae)

Abstract 1 Synthetic blends of bole and foliage volatiles of four sympatric species of conifers were released from pheromone‐baited multiple‐funnel traps to determine if three species of tree‐killing bark beetles (Coleoptera: Scolytidae): (i) exhibited primary attraction to volatiles of their hosts and (ii) discriminated among volatiles of four sympatric species of host and nonhost conifers. 2 Bole and foliage volatiles from Douglas‐fir, Pseudotsuga menziesii (Mirb.) Franco, increased the attraction of coastal and interior Douglas‐fir beetles, Dendroctonus pseudotsugae Hopkins, to pheromone‐baited traps. Primary attraction to bole volatiles was observed in interior D. pseudotsugae. Beetles were significantly less attracted to the pheromone bait when it was combined with volatiles of lodgepole pine, Pinus contorta var. latifolia Engelm. or interior fir, Abies lasiocarpa × bifolia. 3 The monoterpene myrcene synergized attraction of mountain pine beetles, Dendroctonus ponderosae Hopkins, to their aggregation pheromones, but there was no evidence of primary attraction to host volatiles or discrimination among volatiles from the four conifers. 4 There was significant primary attraction of the spruce beetle, Dendroctonus rufipennis Kirby, to bole and foliage volatiles of interior spruce, Picea engelmannii × glauca, but beetles did not discriminate among volatiles of four sympatric conifers when they were combined with pheromone baits. 5 Our results indicate that host volatiles act as kairomones to aid pioneer Douglas‐fir beetles and spruce beetles in host location by primary attraction, and that their role as synergists to aggregation pheromones is significant. For the mountain pine beetle, we conclude that random landing and close range acceptance or rejection of potential hosts would occur in the absence of aggregation pheromones emanating from a tree under attack.     

Estimating the most effective and economical pheromone for monitoring the European spruce bark beetle

In recent decades, there have been an increasing number of outbreaks of Ips typographus in Europe. A large amount of sanitary felling has taken place, with significant economic and ecological consequences. In order to anticipate such large-scale outbreaks, an effective monitoring system should be set up. One important aspect of monitoring is deciding which pheromone to use. Therefore, we decided to test five different commercially available pheromone lures under different disturbance conditions: Pheroprax®, IT Ecolure Extra®, Ipstyp®, Ipsowit® and Typosan®. We investigated the ability of the pheromones to distinguish between disturbed and undisturbed locations, their cost-efficiency ratio, and side effects such as bycatch abundance and composition. We set 50 traps in two areas with sites that were disturbed and undisturbed by windstorms. We collected the catch from traps every 1–2 weeks from the end of March until the end of September in 2019. We found that IT Ecolure Extra®, Ipsowit® and ® Pheroprax® caught the most I. typographus and best showed changes in the trap catch of I. typographus throughout the whole season. There was a low amount of bycatch (<6% of the total catch) and a low number of predators (a few specimens), but some groups seem to prefer certain pheromones. The cost of the pheromones increased with their effectiveness. However, pheromone costs are low relative to the personnel costs involved in setting traps and collecting the catch. Based on all of the gathered data, we created an index which helps to assess the cost-efficiency of the five chosen commercially available pheromones. We also present guidelines on how to make such an index to assist other researchers in choosing the right pheromone for monitoring populations of I. typographus or other bark beetle species.