The European struggle to control Ips typographus - past, present and future

The achievement of the Scandinavian campaign is seen in the swift development and unorthodox transfer of an innovative trapping technique into the management of pest populations of the spruce engraver Ips typographus without the use of pesticides. The pheromone traps proved to be far more efficient in the prevention of engraver reproduction and the reduction of beetle numbers than the cumbersome trap trees they replaced. The wholesale application of pheromone traps as an integrated part of engraver pest management is justified alone by the empirically proven benefits that result from a "surveillance with the intent to control", although appraisal of the actual impact of trapping on the population density remains scientifically ambiguous for the lack of reliable methodology. Furthermore, the campaign opens a new era in bark beetle management by stipulating for the commercial availability of pheromones and traps as well as for research on basic and applied aspects. As the years to come may show, behaviour chemistry is likely to bear more promises to engraver management than just trapping techniques.     

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.     

云杉八齿小蠹的有效积温

采用以云杉八齿小蠹Ips typographus L.侵入云杉皮下为起点,至下一代成虫羽化外出为止的统计方法对其进行发育起点温度和有效积温的测定。结果表明,云杉八齿小蠹此段的发育起点温度为(18.3±0.2)℃,有效积温为(195.4±10)日.度。     

Computer simulated host selection in Ips typographus

Foraging behaviour by Ips typographus L. (Coleoptera; Scolytidae) was simulated to assess host finding and reproductive success of beetles following four different strategies: 1. random search, 2. upwind search, in which beetles follow a prevailing flight direction, 3. random search with short range olfactory attraction to susceptible trees (primary attraction), and 4. upwind search with primary attraction. For each strategy we varied three parameters in equidistant steps including number of susceptible trees per hectare (0.06–0.33), population size (200–1000 beetles), and flight capacity (500–3000 m). All possible combinations of these parameters were tested in 12 replicates with more than 400 000 imaginary beetles flown through the simulated forest. Reproductive success of strategy 4, which combined upwind flight with primary attraction, exceeded that of the other strategies (1–3) circa 27, 7, and 4 times, respectively. In random search, maintenance of the population required a flight capacity, population size and host tree abundance which are unlikely to occur at suboutbreak levels in nature.

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Zusammenfassung Die Wirtsbaumsuche des Buchdruckers wurde in einem Computermodell simuliert und Wirtsbaumfindung und Fortpflanzungserfolg in 4 verschiedenen Suchstrategien quantifiziert: 1. Zufallssuche, 2. Gegenwindsuche, in der die Käfer einer Hauptflugrichtung folgen, 3. Zufallssuche mit Wahrnehmung des bruttauglichen Wirtsbaumes aus kurzer Distanz (primäre Attraktivität), und 4. Gegenwindsuche mit primärer Attraktivität. Für jede Strategie wurden 3 Parameter in äquidistanten Schritten verändert: die Anzahl bruttauglicher Bäume pro Hektar (0.06–0.33), die Populationsdichte der Käfer (200–1000) und die Flugkapazität der Käfer (500–3000 m). Alle möglichen Kombinationen dieser Parameter wurden in 12 Wiederholungen getestet und daher mehr als 400 000 imaginäre Käfer durch den Modellwald geflogen. Der Fortpflanzungserfolg in Strategie 4, die Gegenwindsuche mit Wahrnehmung des Wirtsbaumes aus kurzer Distanz kombinierte, übertraf den in Strategie 1, 2 und 3 circa 27-, 7-, bzw 4-fach. Zur Erhaltung der Käferpopulation waren in der Zufallssuche eine Populationsdichte, Flugkapazität und Wirtsbaumdichte erforderlich, wie sie in der endemischen Phase wohl kaum anzutreffen sind.

     

云杉八齿小蠹信息素的研究进展

在复杂的生态环境中,来自于云杉八齿小蠹和植物的信息物质在云杉八齿小蠹的整个生活史中都发挥着重要的作用。雄性云杉八齿小蠹作为先锋虫以寄主植物的挥发物作为嗅觉信号搜寻和定位寄主,然后释放信息素吸引同性或异性云杉八齿小蠹前来聚集。云杉八齿小蠹成群攻击以降低寄主树木的抗性,寄主植物在受到云杉八齿小蠹攻击后能够利用其物理结构、分泌物及挥发性物质等抵御其危害。云杉八齿小蠹自身化学信息素的合成和对寄主树木次生代谢物质的转化能力,决定和影响着入侵寄主树木的速度和种群数量。本文从信息素组成、释放速率等方面、寄主和非寄主挥发物的作用及信息素诱捕器在监测和防治中的应用做一综述。     

Reemergence and second brood in the bark beetle Ips typographus

The knowledge about reemergence of parent spruce bark beetles Ips typographus . their dispersal, and production of a second brood is reviewed. A majority of the beetles reemerge after their first brood. The process is mainly determined by temperature but high breeding density decreases the average time spent in the tree. The difference between males and females in reemergence seems to be small. In the field, a positive relationship between residence time and fat content at reemergence seems to exist, whereas the opposite tendency is found under laboratory conditions. In the laboratory, the survival of beetles reemerging late is on the average lower than that for early reemerging beetles. The time of reemergence and size of the first brood do not, however, seem to influence the production of a second brood. A large proportion of the reemerging beetles are able to establish a new brood in the laboratory, but calculations based on the number of attacked trees and estimated reemergence in forests in South Norway suggest that only about one third of the beetles actually produce a second brood. In northern Europe, successful colonization of new trees by the reemerging beetles seems to require a period of several warm days during the main reemergence period.     

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.     

Resource dynamic plays a key role in regional fluctuations of the spruce bark beetles Ips typographus

Abstract 1 We analysed time series of spruce bark beetles (N_t) caught in pheromone traps from 1979 to 2000 in approximately 100 localities throughout south‐east Norway. 2 The analysis was performed at two spatial scales (whole area and 12 subregions), and included additional analyses of the time‐series sequences before and after a large windfelling (1987) that appeared to induce a shift in the dynamics. 3 Regression of R_t (logarithmic per‐capita growth rate) on N_{t ? d} indicated that the endogenous dynamics were dominated by lag 1 density dependence. This was particularly so in the sequence after the large windfelling (R² = 0.79), probably due to intraspecific competition for breeding substrate. 4 The sequence before the windfelling (1979–1987) was declining without significant density dependence. We suggest this to be a transition period, reflecting a drop in carrying capacity due to depletion of susceptible trees during the preceding outbreak period (1970s) and a drop in beetle number to below the density required to kill trees. 5 Windfelling (W_{t ? 1}) appears to be an important predictor of the dynamics; however, it was difficult to evaluate the statistical significance of this factor due to only one large windfall event. 6 The weak influence of drought stress (D_{t ? 1}) could be due to the absence of severe drought periods within the time series.     

Exogenous application of methyl jasmonate elicits defenses in Norway spruce (Picea abies) and reduces host colonization by the bark beetle Ips typographus

The terpenoid and phenolic constituents of conifers have been implicated in protecting trees from infestation by bark beetles and phytopathogenic fungi, but it has been difficult to prove these defensive roles under natural conditions. We used methyl jasmonate, a well-known inducer of plant defense responses, to manipulate the biochemistry and anatomy of mature Picea abies (Norway spruce) trees and to test their resistance to attack by Ips typographus (the spruce bark beetle). Bark sections of P. abies treated with methyl jasmonate had significantly less I. typographus colonization than bark sections in the controls and exhibited shorter parental galleries and fewer eggs had been deposited. The numbers of beetles that emerged and mean dry weight per beetle were also significantly lower in methyl jasmonate-treated bark. In addition, fewer beetles were attracted to conspecifics tunneling in methyl jasmonate-treated bark. Stem sections of P. abies treated with methyl jasmonate had an increased number of traumatic resin ducts and a higher concentration of terpenes than untreated sections, whereas the concentration of soluble phenolics did not differ between treatments. The increased amount of terpenoid resin present in methyl jasmonate-treated bark could be directly responsible for the observed decrease in I. typographus colonization and reproduction.     

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.     

Northward shift in temperature sum isoclines may favour Ips typographus outbreaks in European Russia

1. As the development of the eight‐toothed spruce bark beetle Ips typographus is temperature‐dependent, climate change may encourage development of its additional generations per year and facilitate mass outbreaks further north than previously known.
2. The aim of the study was to analyse historical changes in effective temperature sums (ETSs) and early season swarming weather for I. typographus in different forest zones of European Russia between 1960 and 2016. The difference in ETSs was analysed with linear regression using daily temperature data from the 30 meteorological stations. Historical data regarding the location of I. typographus outbreaks were examined and changes in their distribution during the entire study period were analysed.
3. There was a substantial increase in ETSs, especially in the latter half of the study period. Increased ETSs coincided with more favourable conditions for swarming of I. typographus. Areas with favourable ETSs for the complete development of bivoltine populations of I. typographus (>1500 DD) shifted northwards on average 450 km during the entire study period.
4. The northward shift of ETSs may enhance the transition from univoltine to bivoltine life cycles of I. typographus in the south and middle taiga and from bivoltine to trivoltine life cycles in conifer‐broadleaf forests.

     

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.     

Mass attack of trees by Ips typographus induced by sex-specific pheromone: a model of attack dynamics

A model of attack dynamics on a single tree under mass attack by the spruce bark beetle Ips typographus is developed from earlier studies of pheromone production and response of the flying population to the pheromone signal. The model is explored for different numbers and sex-ratios of pioneer beetles and responding beetles. Both numbers of pioneers and responders were important for the dynamics, while the sex-ratio of pioneers was less important. A decreased proportion of males among the responding beetles was important for the result of the model. In the early stage of the attack the model predictions of both numbers and sex-ratio of beetles landing were similar to examples from naturally mass-attacked trees, although the attack proceeded more slowly than in the field. Several aspects of the attack dynamics, such as density-regulation of the number of attacks on the tree, as observed in the field, were not predicted by the model. Gaps in the knowledge of the chemical ecology of the tree-insect relation, such as qualitative change in odours from the tree, spacing mechanisms on the bark surface, and mortality due to host resistence, are pointed out with help of the model.     

Seasonal flight patterns of Ips typographus in southern Sweden and thermal sums required for emergence

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Pathogen's level and parasitism rate in Ips typographus at high population densities: importance of time

We compared the levels of pathogen infection in parental beetles, parasitism of the offspring, abundance of predators and breeding performance success of univoltine populations of Ips typographus in plots characterized by short‐term (2–3 years) outbreaks vs. those with long‐term (>10 years) outbreaks on two localities at ca. 1100 m altitude in the ?umava Mts. The numbers of I. typographus were high in all plots, whether the plots were characterized by long‐term outbreaks or short‐term outbreaks. The numbers of maternal galleries in the sample areas ranged from 300 to 400 per m². The density of parental beetle galleries, abundance of surviving specimens of I. typographus, and length of maternal galleries did not differ between plots. The short‐term outbreaks had only fewer ectoparasitoids of I. typographus and a lower percentage of parasitism and infection level of Mattesia schwenkei than the long‐term outbreaks even though the maternal gallery densities and beetle production were the same. The most mortality appeared to be caused by intraspecific larval competition, and the identical reproductive success in plots with short‐term and long‐term outbreaks indicates that the negative feedback resulting from parasitoids and entomopathogens does not substantially reduce beetle numbers. Although entomopathogenic fungi as Beauveria bassiana occur naturally in the galleries of spruce bark beetles, there was no evidence of its presence in the studied population. The low levels of predation and/or parasitism in both kinds of plots indicate that natural enemies did not play a significant role in reducing outbreak numbers of I. typographus.     

Flight propensity and diapause incidence in five populations of the bark beetle Ips typographus in Scandinavia

Emergence from brood logs, take-off activity and the duration of tethered flight was recorded in the first laboratory generation of the bark beetle Ips typographus L. (Coleoptera: Scolytidae) from five locations (ca 300 km apart) in Sweden and Denmark. Beetles of northerly origin emerged later from brood logs. This pattern was associated with an increasing proportion on non-fliers towards the north. After overwintering, both the rate of development of flight activity and the proportion of non-fliers were the same among populations. Flight duration of fliers was similar among populations and appeared unaffected by outbreak conditions.     

Characterization of microsatellite loci in the spruce bark beetle Ips typographus (Coleoptera: Scolytinae)

Ips typographus is an economically important pest of Norway spruce stands. We developed five polymorphic microsatellite markers using a biotin enrichment protocol. The number of alleles ranged from three to 11 per locus and no strong evidence for null alleles was found. Heterozygosity ranged from 0.3 to 0.97. These markers could be useful tools to study the population structure and genetic consequences of I. typographus population outbreaks.     

The role of sulphur compounds for breeding success of Ips typographus L. (Col., Scolytidae) on Norway Spruce (Picea abies [L.] Karst.)

Bark beetles, especially Ips typographus L. represent a severe biotic threat for spruce ( Picea abies [L.] Karst.) at low altitudes in Europe. We compared sulphur (total S, SO₄²⁻, glutathione, cysteine, methionine), nitrogen (total N, NO₃⁻, total protein, free amino acids), carbon, total phosphorus and PO₄³⁻, tree vigour index (TVI) and water content of the phloem after felling, and their dependent changes (tdc) with the breeding success of I. typographus . Twenty trees were classified according to age (34/90 years) and crown density (high/intermediate/low). Water content was higher in young trees than in old trees, higher in the crown than at breast height, and decreased significantly within the 8-week study period. In old trees, breeding success, length of mother galleries and SO₄²⁻ were significantly higher, while total protein, NO₃⁻ and water content were significantly lower than in young trees. Trees with intermediate crown density provided the best breeding success for I. typographus and had significantly higher arginine content and C/N ratio as well as low amounts of phosphate and glutamine. During the period of bark beetle breeding, total sulphur, glutathione, protein, NO₃⁻, aspartate, glutamine, glutamate, arginine and γ-aminobutyrate decreased significantly. The results support previous investigations that I. typographus develops best in physiologically weakened trees.