Kairomonal responses of Tomicus destruens (Col., Scolytidae) to host volatiles α-pinene and ethanol

The Mediterranean pine shoot beetle Tomicus destruens is one of the most damaging bark beetles attacking Mediterranean pine forests in southern Europe and north Africa. We studied the attractiveness of the host volatiles α-pinene and ethanol at a range of release rates, alone or in combination, to T. destruens , in order to develop an attractive lure for the management of this beetle. T. destruens was attracted slightly to the host volatile α-pinene, but a strong synergistic effect was found in the attraction towards monoterpene when ethanol was added to the bait. The highest catches of T. destruens were obtained by the optimal blend releasing 300 mg/day of α-pinene and 900 mg/day of ethanol. In contrast to data reported for the related species T. piniperda , Mediterranean pine shoot beetles were clearly attracted to baits releasing ethanol alone (1350 mg/day). trans -Verbenol, which was also added to host volatiles in some tests, did not affect the response. The use of the attractive blend proposed would have a low impact on the natural enemy population of Thanasimus formicarius because of asynchronies in flight periods. Other non-target insects, such as the facultative predator or competitor Oxipleurus nodieri , were also significantly attracted. These results allow the development of an operative lure for the effective monitoring of T. destruens , although improvements by the addition of other host volatiles should be studied.     

Development of an improved attractive lure for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

Abstract 1 The pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), is an exotic pest of pine, Pinus spp., and was first discovered in North America in 1992. 2 Although primary attraction to host volatiles has been clearly demonstrated for T. piniperda, the existence and role of secondary attraction to insect‐produced pheromones have been widely debated. 3 Currently, commercial lures for T. piniperda include only the host volatiles α‐pinene in North America and α‐pinene, terpinolene and (+)‐3‐carene in Europe. Several potential pheromone candidates have been identified for T. piniperda. 4 We tested various combinations of host volatiles and pheromone candidates in Michigan, U.S.A., and Ontario, Canada, to determine an optimal blend. 5 Attraction of T. piniperda was significantly increased when trans‐verbenol (95% pure, 3.2%cis‐verbenol content) was added with or without myrtenol to α‐pinene or to blends of α‐pinene and other kairomones and pheromone candidates. 6 Our results, together with other research demonstrating that trans‐verbenol is produced by T. piniperda, support the designation of trans‐verbenol as a pheromone for T. piniperda. A simple operational lure consisting of α‐pinene and trans‐verbenol is recommended for optimal attraction of T. piniperda.     

Characterization of five microsatellite loci in the pine shoot beetle Tomicus piniperda (Coleoptera: Scolytidae)

Tomicus piniperda is a bark beetle that causes damage to various pine species across a wide geographical range. We developed five microsatellite polymorphic markers using an enrichment protocol. All loci could be successfully amplified with no evidence of null alleles and will be useful for population genetic studies. Cross‐species amplifications show that at least some of the markers could be useful in four other Tomicus species.     

Pine shoot beetle, Tomicus piniperda (Col., Scolytidae), responses to common green leaf volatiles

We tested the hypothesis that green leaf volatiles (GLVs) disrupt the response of overwintered pine shoot beetles, Tomicus piniperda (L.) to multiple-funnel traps baited with the attractive host volatile α-pinene. A combination of four GLV alcohols, 1-hexanol ( E )-2-hexen-1-ol ( Z )-2-hexen-1-ol, and ( Z )-3-hexen-1-ol, caused 54 and 36% reduction in the number of pine shoot beetles captured in two separate trapping experiments. Similarly, a combination of the four alcohols plus two GLV aldehydes, hexanal and ( E )-2-hexenal, caused 38% reduction in the number of pine shoot beetles captured compared with α-pinene alone. A blend of the two GLV aldehydes was not disruptive. None of the four GLV alcohols nor the two GLV aldehydes were disruptive when tested individually. The finding that the blend of four GLV alcohols reduced attraction of T. piniperda supports the general hypothesis that GLVs common to nonhost angiosperms are disruptive to conifer-attacking bark beetles (Scolytidae).     

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.     

Host and non-host odour signals governing host selection by the pine shoot beetle, Tomicus piniperda and the spruce bark beetle, Hylurgops palliatus (Col., Scolytidae)

Abstract:  Tomicus piniperda and Hylurgops palliatus colonize susceptible host trees by responding to host-specific odour signals as well as by avoiding volatiles emanating from non-host conifers. In the field, the pine shoot beetle, T. piniperda , responded in high numbers to natural odour sources provided by their host tree, Pinus sylvestris , while the non-host conifers Larix decidua , Picea abies , or Pseudotsuga menziesii were significantly less attractive. In contrast, the spruce bark beetle, Hylurgops palliatus , preferentially responded to its main host, P. abies . Furthermore, T. piniperda attacks on P. sylvestris bolts decreased in presence of bark and wood particles from the non-host P. abies , whereas particles from P. menziesii appeared not to affect T. piniperda attacks. Apparently, tree-specific volatiles act at close range as specific signals that lead to the successful discrimination and colonization of the respective host tree species.     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Shoot feeding and overwintering in the lesser pine shoot beetle Tomicus minor (Col., Scolytidae) in north-west Spain

During 1991, 1992, &, 1993 a study on shoot feeding, overwintering and the presence of Tomicus minor inside pine shoots was carried out in the province of León (NW Spain). This paper deals with the duration of these processes, overwintering sites and pine shoot feeding characteristics: age, diameter, length, occurrence of exit holes, multiple attacks and length of the burrowed tunnels. It also refers to the degree of shoot occupation, sex-ratio and detected adult mortality.     

Threats to pine stands by the pine shoot beetles Tomicus piniperda (L.) and Tomicus minor (Hart.) (Col., Scolytidae) around a sawmill in southern Poland

Forest stands growing around sawmills are especially exposed to damage due to the feeding by the pine shoot beetles Tomicus piniperda (L.) and Tomicus minor (Hart.) (Col., Scolytidae) in the pine shoots, causing a considerable loss of increment. The purpose of this study was to determine the relation of the distance zone of a stand from a sawmill to the density of the pine shoot fall, width of annual rings, and pine stand quality. The investigations were carried out between 1992 and 1996 in a Pinus silvestris – Abies alba stand adjacent to a sawmill in the Zagnańsk Forest District. In the investigated stand, eight zones situated at different distances from the sawmill were marked out, in which the fallen pine shoots were collected and dendrometric measurements taken. The study results showed that the pine shoot fall in the area up to 300 m away from the sawmill was significantly greater than the fall in zones situated further away. In addition, the mean current annual and 10-year radial increments at breast height were significantly smaller, and the pine stand quality class was lower within that area. The radial increment at breast height in the area up to 300 m away from the sawmill was almost twice as low as that in the control area, and about 40% lower than the increment in the area situated 300–900 m away. The increment pattern of pine in this area differed from that of fir, suggesting that the incremental loss and lower pine stand quality are the result of long-term feeding of the pine shoot beetles in the pine shoots. These results indicate that other causes, such as differences in site class, climatic factors or air pollution, can be excluded.     

Suitability of some southern and western pines as hosts for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

The pine shoot beetle, Tomicus piniperda (L.), is an exotic pest that has become established in North America. Discovered in Ohio in 1992, it has since been found in at least 13 states and parts of Canada. The beetle can cause significant growth loss in pines, and it represents a potential threat to trees in areas where it has not yet become established. To evaluate this threat to native pines, field and laboratory tests were conducted on several common and important southern and western species to determine whether they are acceptable hosts for T. piniperda. Comparisons with Pinus sylvestris L., Scots pine, a preferred natural host for the beetle, were made where possible. Measurements of beetle attack success on southern pine billets showed that Pinus taeda L., Pinus echinata Miller, Pinus elliottii var. elliottii Engelmann, Pinus palustris Miller, and Pinus virginiana Miller (loblolly, shortleaf, slash, longleaf, and Virginia pine, respectively) and two western pines, Pinus ponderosa Lawson and Pinus contorta Douglas (ponderosa and lodgepole pine, respectively), were acceptable for breeding material, but brood production was highly variable. Among the southern pines, P. taeda and P. echinata were susceptible to shoot feeding by T. piniperda, whereas P. elliottii was highly resistant and P. palustris seemed to be virtually immune. Shoot feeding tests on the western pines were conducted only in the laboratory, but there was moderate-to-good survival of adults feeding on both species. It seems that if T. piniperda is introduced into the south and west it will likely establish and may cause some damage to native pines. P. taeda may be affected more than other southern pines because it is the most abundant species, it is readily attacked for brood production, which can result in moderately large broods, and the beetle survives well during maturation feeding on P. taeda shoots.     

三种绿叶挥发性物质对云南切梢小蠹寄主定位行为的干扰作用

为了研究环境中非寄主阔叶植物释放出的绿叶挥发性物质（GLVs）对针叶树蛀干害虫云南切梢小蠹Tomicus yunnanesis的影响, 选取了(E)-2-己烯醛、 (E)-2-己烯醇和(Z)-3-己烯醇3种释放量较大的绿叶挥发性物质, 通过室内松梢取食试验测试了单组分及两两混合后对云南切梢小蠹寄主定位行为的干扰作用。结果表明： 源于阔叶植物的3种绿叶挥发性物质及其混合物能够不同程度干扰云南切梢小蠹的寄主定位行为。当虫放入广口瓶12 h后, 3个单组分绿叶挥发性物质处理组［A： (E)-2-己烯醛, P＜0.01; B： (E)-2-己烯醇, P＜0.01; C： (Z)-3-己烯醇, P＜0.01］及2个混合组分［D： (E)-2-己烯醛+(E)-2-己烯醇, P＜0.01）; E： (E)-2-己烯醛+(Z)-3-己烯醇, P＜0.01］中滞留在松梢外部的虫数与对照组相比都有显著性差异, 绿叶挥发性物质的存在显著降低了云南切梢小蠹侵害云南松松梢的概率。但是, 24 h后只有D组（P＜0.01）和E组（P＜0.01）滞留在松梢外部的虫数与对照组相比具有显著性差异, 在48 h后只有D组（P＜0.01）与对照相比仍具有显著性差异。本研究为利用非寄主植物的次生代谢产物防治云南切梢小蠹进行了有益的探索。     

Attraction of the bark beetle Tomicus piniperda and some other bark- and wood-living beetles to the host volatiles α-pinene and ethanol

The attraction of some bark- and ambrosia beetles as well as associated beetles to the host volatiles -pinene and ethanol was studied in field tests with flight barrier traps. Tomicus piniperda (L.) (Scolytidae), Thanasimus formicarius (L.) (Cleridae), and Rhizophagus ferrugineus (Payk.) (Rhizophagidae) were attracted by -pinene, while Hylurgops palliatus (Gyll.) and Trypodendron lineatum (Oliv.) (Scolytidae) were attracted by ethanol and Epuraea spp. (Nitidulidae) by both -pinene and ethanol. Combinations of -pinene and ethanol attracted high numbers of H. palliatus, T. lineatum, R. ferrugineus, Epuraea spp., and Glischrochilus spp. (Nitidulidae) and the catches increased with increasing release rates of ethanol. By contrast, lower numbers of T. piniperda were caught in traps baited with combinations of -pinene and ethanol than in traps baited with -pinene alone, and the catches of this species decreased with increasing release rates of ethanol. Traps baited with a combination of -pinene and ethanol or with -pinene alone caught similar numbers of T. formicarius. The results are discussed on the basis of species differences in preference for breeding substrate.

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Zusammenfassung Die Anlockung mehrerer Borkenkäfer und assoziierter Käferarten zu den flüchtigen Wirtsstoffen -Pinen und Äthanol wurde in Feldversuchen mit Flugbarrierenfallen studiert. Tomicus piniperda (L.) (Scolytidae), Thanasimus formicarius (L.) (Cleridae) und Rhizophagus ferrugineus (Payk.) (Rhizophagidae) wurden durch -Pinen angelockt, die Borkenkäfer Hylurgops palliatus (Gyll.) und Trypodendron lineatum (Oliv.) durch Äthanol und die Epuraea-Arten (Nitidulidae) durch sowohl -Pinen als auch Äthanol.Kombinationen von -Pinen und Äthanol lockten viele Individen von H. palliatus, T. lineatum, R. ferrugineus, Epuraea spp. und Glischrochilus spp. (Nitidulidae) an, und die Fänge nahmen mit zunehmender Äthanol-Abgabe zu. Umgekehrt wurden weniger T. piniperda in Fallen mit Kombinationen von -Pinen und Äthanol gefangen als in Fallen mit -Pinen allein, und die Waldgärtner-Fänge nahmen mit zunehmender Äthanol-Abgabe ab. Die Fänge von T. formicarius in Fallen mit einer Kombination von -Pinen und Äthanol unterschieden sich nicht von denen in Fallen mit nur -Pinen.Die Ergebnisse werden auf der Grundlage der Unterschiede zwischen den Arten in der Wahl des Brut-substrats besprochen.

     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.     

Use of pityol to reduce damage by the white pine cone beetle, Conophthorus coniperda (Col., Scolytidae) in seed orchards

Abstract:  The white pine cone beetle, Conophthorus coniperda (Schwarz), is a common and destructive pest of eastern white pine cones. The potential of mating disruption as a pest management tool against C. coniperda was tested during this experiment. The 5.5 ha white pine seed orchard used in this study was separated in three equal sections and different blocks were treated with pityol over 2-year experiments. Ten and 20 pityol dispensers (bubble caps) were hung per treated block in 2001 (east block) and 2002 (centre block), respectively, to evaluate their impact on cone protection. During both seasons, the percentage of damaged cones at permanent stations was evaluated at monthly intervals in the middle of the treated block and at different places in the control area. For both years, the mean reduction in cone damage was 63.7% in the pheromone-treated blocks. With this study, we have been able to demonstrate that mating disruption could be used as a biological pest management tool for the control of C. coniperda in white pine seed orchards.     

Influence of the proportion of (−) α-pinene in pheromone bait on Ips typographus (Col., Scolytidae) catch in pheromone trap barriers and in single traps

Abstract:  Further possibilities of increasing catch efficiency of Ips typographus L. in pheromone trap barriers with two different levels of release of (4S)- cis -verbenol (cV) and in single traps were tested. Pheromone mixtures containing cV, 2-methyl-3-buten-2-ol (MB) and 0, 1, 10, 35, 60 and 90% (−) α -pinene were prepared. Basically, MB was successively replaced by (−) α -pinene in the mixtures. Tests were performed on the edge of a spruce forest when outbreaks of I. typographus occurred. The substitution of MB with (−) α -pinene had different effects in different types of pheromone trap barriers and in single traps. It is not possible to increase the attractiveness of the bait having a higher percentage of cV by supplementing a certain percentage of MB with (−) α -pinene. The catch of I. typographus decreased with an increase in the percentage of (−) α -pinene in the bait. It is possible to increase the attractiveness of the bait having a lower percentage of cV by supplementing certain percentages of MB with (−) α -pinene. A slight synergistic effect of (−) α -pinene with MB/cV mixture is visible in this type of bait. In the case of single traps, supplementing MB with (−) α -pinene possibly causes a slight synergistic effect. An increase in the percentage of (−) α -pinene increases the percentage of males in the catch.     

The role of beetle and host volatiles in host colonization in the European oak bark beetle, Scolytus intricatus (Ratzeburg) (Col., Scolytidae)

Abstract:  To understand the role of chemical signals involved in Scolytus intricatus (Ratzeburg) (Col., Scolytidae) host colonization, the attractiveness of intact and beetle colonized host material was investigated in the field and in the laboratory. In the field, chemical signals operating at long range were investigated by means of sticky traps. In the laboratory, close-range chemical interactions were investigated with an arena olfactometer. Field experiments showed no differences in the attractiveness between infested and non-infested host material. On the contrary, laboratory experiments revealed higher attractiveness of infested host and beetle-produced frass in comparison with intact host and mechanically produced sawdust respectively. Laboratory data also disclosed the attractiveness of beetle extracts of both sexes. Our data show that: (1) host kairomones play an important role during host colonization, and that (2) S. intricatus does not use a sex-specific secondary attractant system. Differences between results of field and laboratory trials are discussed.     

N-Linked Glycosylation of the α-Amino-3-Hydroxy-5-Methylisoxazole-4-Propionate(AMPA)-Selective Glutamate Receptor Channel α2 Subunit Is Essential for the Acquisition of Ligand-Binding Activity

Abstract: The N-linked glycosylation of the α2 subunit of the mouse α-amino-3-hydroxy-5-methylisoxazole-4-propionate(AMPA)-selective glutamate receptor (GluR) channel was characterized. The receptor subunit protein has five putative N -glycosylation sites. The recombinant receptor proteins were identified by [³⁵S]methionine/[³⁵S]cysteine metabolic labeling, western blot analysis, immunocytochemical detection, and [³H]AMPA binding experiments when expressed in insect Spodoptera frugiperda cells using a baculovirus system. The effect of tunicamycin on the metabolic labeling and immunoblots suggested that the two products, a major protein species of ∼102 kDa and a minor species of ∼98 kDa, correspond to glycosylated and unglycosylated forms, respectively, which was also supported by the enzymic deglycosylation experiments. Immunofluorescence staining of tunicamycin-treated cells expressing only the unglycosylated form differed little from that of tunicamycin-nontreated cells expressing both glycosylated and unglycosylated forms. The lack of AMPA-binding activity of the unglycosylated form expressed in the presence of tunicamycin suggested that N -glycosylation is required, directly or indirectly, for functional expression in insect cells for ligand binding. These results demonstrate that occupancy of at least one N -glycosylation site is required for the formation and maintenance of the GluRα2 subunit protein in an active conformation for ligand binding. Possible roles of N -glycosylation of GluRα2 subunit protein are discussed.