Host tree resistance influencing pheromone production in Ips typographus (Coleoptera: Scolytidae)

Analyses of volatiles in hindguts of Ips typgraphus males from different spruce trees and attack phases are reviewed. The composition of monoterpenes, and the chirality of α-pinene, have been determined in phloem samples. Relationships between compounds emanating from spruce trees and bark beetles, respectively, have been studied. Male beetles depend on their host tree for the production of pinene alcohols. The ratio between the pinene alcohols is almost constant in males boring in the same tree but can vary widely between males from different spruce trees. Very good correlations were found between some host tree monoterpene hydrocarbons and bark beetle produced pinene alcohols. The production of the essential pheromone component 2-methyl-3-buten-2-ol, was not correlated with the monoterpene content in the host trees, while the production of the other essential pheromone component, cis-verbenol, depends on the amount of the precursor, (−)-α-pinene in the phloem. Male beetles boring in a resistant spruce tree will continue to produce the pinene alcohols, including cis -verbenol, as long as the tree defends itself with resin.     

Behavioural sequence in the attraction of the bark beetle Ips typographus to pheromone sources

ABSTRACT. The functions of the two synergistic pheromone components, (4S)- cis -verbenol (cV) and 2-methyl-3-buten-2-01 (MB), and the role of ipsdienol in the attraction of Ips typographus (L.) (Scolytidae) to pheromone sources were studied in the field. Absolute and relative beetle catches were compared between several traps placed at and nearby a central pheromone source: a pipe trap containing the source, a surrounding sticky trap, a nearby window trap, and four distant window traps. A higher catch in the outer down-wind distant traps indicated an up-wind anemotaxis to the source. Increased MB release, with cV constant, increased the proportion caught in the central pipe trap, indicating MB as a landing stimulus. Release of MB alone gave a very small catch. Ipsdienol could not substitute for cV in the synergism with MB. An increase of cV, with MB constant, increased the number of beetles caught, but not the proportion caught in the pipe trap. The sex ratio was equal in the window traps, but fell to 30% males in both sticky and pipe traps, showing that a large proportion of the males attracted to the source did not land. The proportion of males in the pipe trap was reduced at the highest cV dose. The results support the idea of each pheromone component having a different relative importance in releasing different steps in the behavioural chain.     

Behavior of male and female spruce bark beetles,Ips typographus,on the bark of host trees during mass attack

The behavior of 118 spruce bark beetles, Ips typographus,was observed on trees under colonization. Most individuals were followed from when they landed until they entered or left the tree. Both males and females spent most time inspecting crevices and searching for a place to start boring or for a hole to enter. These behaviors accounted for 87 and 70% of all behavioral acts recorded for males and females, respectively. Females entered galleries with males only after a period of pushing at the gallery entrance. Males spent on average 3 min and females 4 min on the bark before entering or leaving the tree. Thirty-three percent of the beetles eventually entered the tree, 31% flew away, 35% dropped from the host, and one beetle was eaten by a predator. The results are discussed in relation to the question of mate choice in bark beetles and to studies on attack dynamics of spruce bark beetle populations.     

Ips typographus (L.) pheromone trapping in south Alps: spring catches determine damage thresholds

Abstract: Ips typographus is one of the major forest pests in the Italian Alps. From 1996, populations of I. typographus in Friuli‐Venezia Giulia region (NE Italy) have been permanently monitored in areas of outbreak by using pheromone traps. At the same time, damage caused by the insect was estimated annually. Preliminary analysis of the data (1996–2002) reveals some interesting information: (i) the flight activity of I. typographus is very extended and occurs over a period of about 4 months (May to August); (ii) there is a high correlation between mean captures per trap and annual damage; (iii) there is a high correlation between spring captures (May to mid‐June) and total captures (May to August), and between spring captures (May to mid‐June) and annual damage. The correlation between spring captures and damage allows the determination of a reliable risk‐damage threshold (about 5000 beetles/trap, in spring), reducing at the same time, the monitoring period and the general costs.     

Pathogens of Ips amitinus: new species and comparison with Ips typographus

Ips amitinus and I. typographus are two serious pests of spruce in Europe, have similar bionomics and are likely to occur and meet on the same host trees. We therefore hypothesized that the two species support similar levels of similar pathogens. To test this hypothesis, we collected mature beetles from three trap trees at each of eight study sites and determined beetle numbers and pathogen infection levels. In total, 938 mature I. amitinus beetles and 3435 of I. typographus were dissected; five pathogens, as well as intestinal nematodes and endoparasitoids, were detected. The neogregarine Mattesia schwenkei is reported here for the first time as a new pathogen in 9.4% of I. amitinus individuals at one site. Average infection levels of most pathogens (Chytridiopsis typographi, Gregarina typographi, Mattesia schwenkei and parasitoids) were significantly higher in I. typographus than in I. amitinus. Metschnikowia typographi was confirmed only in Ips amitinus, while the microsporidium of Nosema typographi occurred only in I. typographus. Within‐season increases in G. typographi infection levels were documented in Ips amitinus.     

Biosynthesis of 2-methyl-3-buten-2-ol,a pheromone component of Ips typographus (Coleoptera: Scolytidae)

Exposure of beetles to ¹⁴C-labelled mevalonate by injection, resulted in significant incorporation of radioactivity in 2-methyl-3-buten-2-ol. Radiolabelled 2-methyl-3-buten-2-ol was obtained both by organic solvent extraction of beetle hindguts and by entrainment of volatiles in air surrounding logs with boring beetles. It is suggested that 2-methyl-3-buten-2-ol, an essential component of the aggregation pheromone of Ips typographus, can be synthesized from mevalonate.     

Behavioural and electrophysiological responses of the bark beetle, Ips typographus, to potential pheromone components

ABSTRACT. Field tests showed ( S )-(-)- cis -verbenol and ( R )-(+)- trans -verbenol in combination with a second pheromonal component, 2-methyl-3-buten-2-ol, more attractive than the combination with their optical antipodes. Inhibition of response to the attractant component, ( S )-(-)- cis -verbenol, by high concentrations of its optical antipode did not occur. No significant differences were noted for response to the attractant, ( S )-(-)- cis -verbenol and 2-methyl-3-buten-2-ol, with the addition of either ipsdienol enantiomer or a racemic mixture of ipsdienol enantiomers. Electroantennogram (EAG) studies correlated well with the behavioural studies. EAGs recorded from male and female beetles revealed both sexes to have a lower threshold for the pheromone, ( S )-(-)- cis -verbenol, than its host terpene precursor, (-)- alpha -pinene. EAGs showed a greater number of acceptors for (-)- alpha-pinene in males than in females. EAGs at acceptor saturation to the enantiomers of alpha -pinene and the verbenol isomers showed males more responsive to (-)- alpha -pinene, (±)- cis -verbenol, and ( R )-(+)- trans -verbenol. Significantly greater EAGs were elicited in females than in males to (-)- alpha -pinene, and (±)- and ( S )-(-)- cis -verbenol. No significant differences in EAGs of females to the enantiomers of trans -verbenol were noted. EAGs showed similar thresholds in males and females to the pheromone component, 2-methyl-3-buten-2-ol; however, female response at threshold was significantly greater than male response. The results are discussed with regard to olfactory acceptor evolution.     

A host monoterpene influences Ips typographus (Coleoptera: Curculionidae,Scolytinae) responses to its aggregation pheromone

1 Host tree terpenes can influence attraction of conifer‐infesting bark beetles to their aggregation pheromones, and both synergistic and inhibitory effects have been reported. 2 We tested a gradient of ratios of (–)‐α‐pinene, the predominant monoterpene in Norway spruce, to the pheromone of Ips typographus, a major pest of Norway spruce. 3 Attraction of I. typographus increased as the release rate of (–)‐α‐pinene increased. The two highest (–)‐α‐pinene : pheromone ratios (526 : 1 and 2595 : 1) attracted twice as many I. typographus as pheromone alone, whereas low to intermediate ratios (56 : 1, 274 : 1) did not differ from pheromone alone. 4 Our results are in agreement with a proposed model, which suggests that bark beetles display unique response profiles to host terpenes depending on the physiological condition of the host trees that they typically colonize. Ips typographus, which is an aggressive species capable of colonizing and killing healthy trees, showed an increased attraction to monoterpene : pheromone ratios, and this may be high enough to inhibit attraction of less aggressive beetle species typically colonizing dead, dying or stressed trees. 5 Attraction of associates of I. typographus was also modified by (–)‐α‐pinene. Ips duplicatus, a competitor of I. typographus, showed increased attraction to the pheromone of I. typographus across all concentrations of (–)‐α‐pinene.     

Interruption of aggregation pheromone in Ips typographus (L.) (Col. Scolytidae) by non-host bark volatiles

Abstract 1 The antennally active nonhost bark volatiles (NHVs): trans‐conophthorin (tC), C₆‐alcohols (green leaf volatiles; GLVs) and C₈‐alcohols, were tested for their ability to reduce attraction of the spruce bark beetle Ips typographus (L) (Col. Scolytidae) to its pheromone sources in both laboratory walking bioassy and field trapping experiments. 2 In the walking bioassay with I. typographus females, individual NHVs such as tC, 3‐octanol and 1‐octen‐3‐ol, and the unsuitable host signal, verbenone (Vn), were inactive at the doses tested. However, the blend of C₆‐alcohols (3GLVs) and all the binary, ternary, or quarternary blends significantly reduced the female attraction to the pheromone sources. 3 In the field trapping experiments, individual NHV signals (tC, C6‐alcohols and C₈‐alcohols) all reduced catch of I. typographus in pheromone‐baited traps, with their inhibitory effects similar to that of the known inhibitor, Vn. The binary, ternary or quarternary combinations of these NHV signals or Vn, all caused significantly stronger reductions in trap catches than the individual signals. The blends showed similar levels of interruption, except the binary blend of C₈‐alcohols (2C8OH) and Vn. 4 Difference in trapping mechanism between pipe traps (attraction and landing) and Lindgren funnel traps (attraction) did not affect the pattern of inhibition of these active NHV signals and Vn. 5 These behaviourally active nonhost volatiles and Vn might be used effectively to protect spruce trees or stands against attacks by I. typographus.     

Tree killing by Ips typographus (Coleoptera: Scolytidae) at stand edges with and without colonized felled spruce trees

Abstract 1 To maintain biodiversity in managed spruce forests in Sweden more wind‐felled trees must be retained. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col. Scolytidae). 2 To simulate wind‐felled trees, living spruce trees were cut at spruce stand edges bordering fresh clear‐cuttings. Treatments comprised edges with zero, one or five cut trees colonized by I. typographus. Edges with naturally wind‐felled trees colonized by I. typographus were also included in the analyses. 3 During the two following summers, the number of trees killed by I. typographus did not differ between edges with and without felled trees, or between edges with one or five felled trees. 4 Within edges with felled trees, there were more killed trees close to the felled trees than at other parts of the edges. Thus, felled trees provided focal points for attacks within edges. 5 It is concluded that small numbers of wind‐felled trees colonized by I. typographus may be left near spruce stand edges without increasing the risk of beetle‐induced tree mortality.     

Quantification of different yeasts associated with the bark beetle,Ips typographus,during its attack on a spruce tree

There were different amounts and types of yeasts associated with individuals ofIps typographus spruce bark beetles during different phases of their attack on a healthy spruce tree. The yeasts were isolated on Sabouraud agar medium in order to identify them and estimate their numbers.Hansenula holstii andCandida diddensii type yeasts were most frequently isolated. The increase in number of these two yeast types probably accounted for most of the total yeast increase found during the later attack phases of the bark beetles. Lesser amounts ofHansenula capsulata, Pichia pinus, Candida nitratophila, and twoCryptococcus type yeasts were also found.     

Attacks on living spruce trees by the bark beetle Ips typographus (Col. Scolytidae) following a storm-felling: a comparison between stands with and without removal of wind-felled trees

Abstract 1 To maintain biodiversity in forests more wind‐felled trees must be left where they fall. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col.: Scolytidae). 2 In the 5 years following a major storm disturbance the number of standing spruces killed by I. typographus was determined in a total of 53 stands. In five of the stands all wind‐thrown trees were left (unmanaged stands) and in 48 of the stands, which were situated at distances of 1.4–10.0 km from each focal unmanaged stand, the wind‐felled trees were removed directly after the storm (managed stands). In the winter preceding the fifth summer new storm‐fellings occurred in the study area. 3 In the 4‐year period between the first and second storm‐fellings, 50–322 standing trees were killed by I. typographus per unmanaged stand. There was a direct linear relationship between the number of storm‐felled spruces colonized by I. typographus and the number of trees subsequently killed in the unmanaged stands. 4 Tree mortality caused by I. typographus in the unmanaged stands was almost nil in the first year, peaked in the second or third year, and decreased markedly to a low level in the fourth and fifth years. 5 In the 4‐year period between the first and second storm‐fellings twice as many trees were killed per ha in the unmanaged stands than in the managed stands: the average difference being 6.2 killed trees per ha, equivalent to 19% of the number of spruce trees felled by the first storm in the unmanaged stands. 6 Much higher numbers of trees were killed per ha in the stand edges than in the interiors of both the unmanaged and the managed stands.     

Characteristics of Norway spruce trees (Picea abies) surviving a spruce bark beetle (Ips typographus L.) outbreak

The characteristics of spruce individuals, which survived a massive bark beetle outbreak, were compared with the characteristics of neighbouring attacked trees in Šumava National Park (Czech Republic). Selected parameters related to crown geometry, stand conditions and distances between trees were measured or estimated. Significant differences were found between the surviving trees and the neighbouring trees attacked by I. typographus. Trees with a higher level of stem shading (longer crown length) tended to survive. The attacked trees were usually located in areas with larger basal area, especially southwards from them. A shorter distance to a previously attacked tree increased the probability of additional attack. Spruce trees with more progressive crown structure transformation (primary structure defoliation) were significantly more frequently attacked by spruce bark beetle. Superior and taller trees had a clearly longer life expectancy than dominant ones. These results show that the attack of trees by bark beetle can be predicted to a certain degree, which can be used in management of endangered spruce forests.     

Volatiles from spruce trap-trees detected by Ips typographus bark beetles: chemical and electrophysiological analyses

In the search for compounds that contribute to the host or habitat discrimination, antennae of Ips typographus were screened for sensitivity to volatiles released by spruce trap-trees using gas chromatography linked to electroantennography. The antennally active compounds were determined using comprehensive two-dimensional gas chromatography with time-of-flight mass spectrometric detection. Data show that I. typographus antennae respond to compounds emitted by the host. In total, 18 of antennally active compounds were detected: α-pinene, camphene, sabinene, β-pinene, myrcene, Δ-3-carene, p-cymene, limonene, β-phellandrene, 1,8-cineole, γ-terpinene, terpinolene, nonanal, camphor, trans-pinocamphone, cis-pinocamphone, terpinen-4-ol, and verbenone. Unequivocal identification of all active minor compounds is provided and confirmed using synthetic standards. Compounds in minor quantities like 1,8-cineole, β-phellandrene, camphor, cis-pinocamphone, and trans-pinocamphone were more active than major spruce monoterpenes. We hypothesize that the minor spruce compounds may play so far unrecognized role in conveying information about host suitability for I. typographus.     

Redundancy, synergism, and active inhibitory range of non-host volatiles in reducing pheromone attraction in European spruce bark beetle Ips typographus

Brandt's voles ( Microtus brandti ) are small native mammals that inhabit the grasslands of Inner Mongolia, China. The species is considered a pest, particularly during population outbreaks, which have increased in frequency since 1970 from 1 every 7 years to 1 every 3 to 5 years. Using historical records taken between 1948 and 1998, we found that there was significant correlation between years for which the monthly averages of the Southern Oscillation Index (SOI) were consistently high, and years in which outbreaks occurred. Also there was a tendency for outbreaks not to occur in years with precipitation above or below average. For some years at some sites, annual or biannual estimates are available for the density of Brandt's vole populations. We found that the seasonal rate of increase over the non-breeding season from autumn to spring was negatively correlated with the density in autumn. The rate of increase over the breeding season from spring to autumn was independent of the density in spring but instead, reflects the species' preference for habitat with short, sparse grass; populations do not persist in highly degraded grasslands or in tall, dense grassland. The link between outbreaks and climatic indices, and the numerical response of Brandt's voles to particular habitat characteristics, suggest that current grazing intensity tends to maintain grass at low height in years with average precipitation. The substantial increase in livestock numbers over the last 50 years appears to have increased the number of years when the balance of grazing and plant growth favours high rates of increase in Brandt's vole populations, and hence, there has been an increase in the frequency of outbreaks.     

Statistical evaluation of Ips typographus population density: a useful tool in protected areas and conservation-oriented forestry

Ips typographus (Col., Curculionidae, Scolytinae) occurring on Picea abies stems is a species characterised by large fluctuations in population numbers and causing frequent outbreaks. In protected areas, I. typographus is regarded as a sensitive bioindicator responsive to changes in forest health and vitality. In conservation-oriented forestry, attention is being paid to the ecological value of I. typographus beetles as ecosystem engineers and keystone species, driving forest natural regeneration and conversion. Despite many publications devoted to I. typographus, no accurate method for estimating the population density of this species has been developed. The objective of this study was to develop a statistical method for estimating I. typographus population density that enables calculation of estimation errors. The proposed method consists of two parts: tree-level analyses and stand-level analyses. Part one allows calculation of the total density of I. typographus infestation of each of P. abies selected stem (after selecting sample windfalls), part two allows estimation of the mean total infestation density of the stem in the area investigated. Linear regression functions were applied to part one and survey sampling to part two. The method is only marginally invasive because it involves debarking of one 0.5 m-long stem section on maximum 50 P. abies windfalls (trap trees can optionally be used). The developed method was employed to assess I. typographus population density in the ?wi?tokrzyskie Mountains in Central Poland in an area of ca. 4,000 ha. In 2010, in the area investigated, the mean total I. typographus infestation density of the P. abies stem was 440.6 maternal galleries/m² (from 358.7 to 522.6 maternal galleries/m² with ?? = 0.05; the relative error of estimation was 18.6%). The examined I. typographus population was in a progradation phase. The proposed method can be used in nature reserves, national parks and managed forests, mainly for scientific purposes.     

Influence of proportion of (4S)-cis-verbenol in pheromone bait on Ips typographus (Col., Scolytidae) catch in pheromone trap barrier and in single traps

Abstract:   The possibilities of increasing catching efficiency of male Ips typographus L in pheromone traps barriers and in single traps were tested. Pheromone dispensers with different percentages of (4S)- cis -verbenol (cV) in pheromone mixtures were used. The results show that the different male catching efficiencies of pheromone trap barriers and single traps were related to the different percentages of cV in the pheromone bait. The proportions of males were higher in pheromone trap barriers than in single traps. Thus increasing the catch of males in pheromone trap barriers is possible by using a combination of pheromone traps containing pheromones giving a higher release of cV with others containing pheromones giving a lower release of cV .