Can Norway spruce trees be ‘vaccinated’ against attack by Ips typographus?

1 A field experiment was carried out to test the hypothesis that treatment of Norway spruce trees with the Ips typographus-transmitted blue-stain fungus Ceratocystis polonica enhances tree resistance to later mass attack by this bark beetle. 2 Twenty-five mature trees were pretreated by inoculating a non-lethal dose of the fungus into the bark, while 18 trees served as untreated controls. Three and a half weeks after treatment a bark beetle attack was initiated by attaching dispensers with I. typographus pheromone to the tree trunks. 3 A significantly larger proportion (67%) of the control trees than of the pretreated trees (36%) were killed by the beetle attack. The result is discussed in relation to recent results regarding defence mechanisms in Norway spruce trees.     

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.     

Recent advance of white spruce (Picea glauca) in the coastal tundra of the eastern shore of Hudson Bay (Québec, Canada)

Aim The species‐specific response of tree‐line species to climatic forcing is a crucial topic in modelling climate‐driven ecosystem dynamics. In northern Québec, Canada, black spruce (Picea mariana) is the dominant species at the tree line, but white spruce (Picea glauca) also occurs along the maritime coast of Hudson Bay, and is expanding along the coast and on lands that have recently emerged because of isostatic uplift. Here we outline the present distribution, structure, dynamics and recent spread of white spruce from the tree line up to its northernmost position in the shrub tundra along the Hudson Bay coast. We aimed to obtain a minimum date of the arrival of the species in the area and to evaluate its dynamics relative to recent climate changes. Location White spruce populations and individuals were sampled along a latitudinal transect from the tree line to the northernmost individual in the shrub tundra along the Hudson Bay coast and in the Nastapoka archipelago in northern Québec and Nunavut, Canada (56°06′–56°32′ N). Methods White spruce populations were mapped, and the position, dimension, growth form and origin (seed or layering) of every individual recorded. Tree‐ring analyses of living and dead trees allowed an estimation of the population structure, past recruitment, growth trends and growth rate of the species. A macrofossil analysis was performed of the organic horizon of the northernmost white spruce stands and individuals. Radiocarbon dates of white spruce remains and organic matter were obtained. The rate of isostatic uplift was assessed by radiocarbon dating of drifted wood fragments. Results The first recorded establishment of white spruce was almost synchronous at all sites and occurred around ad 1660. Spruce recruitment was rather continuous at the tree line, while it showed a gap in the northern shrub tundra during the first decades of the 19th century. A vigorous, recent establishment of seedlings was observed in the shrub tundra; only wind‐exposed, low krummholz (stunted individuals) did not show any sexual regeneration. A period of suppressed growth occurred from the 1810s to the 1850s in most sites. A growth increase was evident from the second half of the 19th century and peaked in the 1880s and the 20th century. A shift from stunted to tree growth form has occurred since the mid‐19th century. No sample associated with white spruce remains gave a date older than 300 ¹⁴C years bp [calibrated age (cal.) ad 1430–1690]. Main conclusions White spruce probably arrived recently in the coastal tundra of Hudson Bay due to a delayed post‐glacial spread. The arrival of the species probably occurred during the Little Ice Age. The established individuals survived by layering during unfavourable periods, but acted as nuclei for sexual recruitment almost continuously, except in the northernmost and most exposed sites. Warmer periods were marked by strong seedling recruitment and a shift to tree growth form. Unlike white spruce, black spruce showed no evidence of an ongoing change in growth form and sexual recruitment. Ecological requirements and recent history of tree‐line species should be taken into account in order to understand the present dynamics of high‐latitude ecosystems.     

Recent climate warming forces contrasting growth responses of white spruce at treeline in Alaska through temperature thresholds

Northern and high‐latitude alpine treelines are generally thought to be limited by available warmth. Most studies of tree‐growth–climate interaction at treeline as well as climate reconstructions using dendrochronology report positive growth response of treeline trees to warmer temperatures. However, population‐wide responses of treeline trees to climate remain largely unexamined. We systematically sampled 1558 white spruce at 13 treeline sites in the Brooks Range and Alaska Range. Our findings of both positive and negative growth responses to climate warming at treeline challenge the widespread assumption that arctic treeline trees grow better with warming climate. High mean temperatures in July decreased the growth of 40% of white spruce at treeline areas in Alaska, whereas warm springs enhance growth of additional 36% of trees and 24% show no significant correlation with climate. Even though these opposing growth responses are present in all sampled sites, their relative proportion varies between sites and there is no overall clear relationship between growth response and landscape position within a site. Growth increases and decreases appear in our sample above specific temperature index values (temperature thresholds), which occurred more frequently in the late 20th century. Contrary to previous findings, temperature explained more variability in radial growth after 1950. Without accounting for these opposite responses and temperature thresholds, climate reconstructions based on ring width will miscalibrate past climate, and biogeochemical and dynamic vegetation models will overestimate carbon uptake and treeline advance under future warming scenarios.     

Primary succession of subarctic vegetation and soil on the fast‐rising coast of eastern Hudson Bay,Canada

Aim The objectives of the study are: (1) to evaluate the dynamics of the maritime tree line and forest limit of white spruce, Picea glauca, within the dual framework of primary succession induced by the rapid post‐glacial land emergence on the eastern coast of Hudson Bay and the impacts of recent and past climate changes; and (2) to determine the time lapse between land emergence and seedling, tree, and forest establishment in the context of the primary chronosequence occurring on rising, well‐drained sandy beaches and terraces. Location The study area was located on the eastern coast of Hudson Bay (56°20′ N, 76°32′ W) in northern Québec, Canada. Methods We evaluated the colonization dynamics of white spruce as seedlings, tree‐line trees and primary‐forest trees at eight sites distributed along a 200‐km latitudinal gradient based on a mean land emergence rate of 1.2 m century^?1. A 30‐m wide by 140–300‐m long quadrat was positioned at random at the centre of each site. The elevation above sea level, position and age of all individuals of spruce present in the quadrat areas were determined, and the soils of each chronosequence were described. Results The main stages of primary succession along the emerging coast were common to all the sites, regardless of latitude, but occurred at different elevations above sea level (a.s.l.). White spruce seedlings colonized near‐shore beaches 2 m a.s.l., whereas the tree line and forest limit tended to form only at about 3–4 m and 4–8 m a.s.l., corresponding approximately to 180–825 years and 310–1615 years after land emersion, respectively. White spruce establishment at the tree line occurred about 50 years ago. Climatic conditions at this time were probably more favourable to tree colonization than when the species established at the forest limit. Soil formation was influenced primarily by distance from the seashore and elevation above sea level, with podzolization being accelerated by white spruce cover. Main conclusions The current tree‐line and forest‐limit positions on the rising coast of eastern Hudson Bay correspond to ecological limits established during the course of primary succession within a context of changing climatic conditions. The recent establishment of trees at the tree line and forest limit at relatively old coastal sites is associated with warmer conditions over the last 100 years. Although white spruce was present nearby, coastal sites were devoid of trees before the 20th century.     

Fungal associates of the tree-killing bark beetle,Ips typographus,vary in virulence,ability to degrade conifer phenolics and influence bark beetle tunneling behavior

The bark beetle Ips typographus carries numerous fungi that could be assisting the beetle in colonizing live Norway spruce (Picea abies) trees. Phenolic defenses in spruce phloem are degraded by the beetle's major tree-killing fungus Endoconidiophora polonica, but it is unknown if other beetle associates can also catabolize these compounds. We compared the ability of five fungi commonly associated with I. typographus to degrade phenolic compounds in Norway spruce phloem. Grosmannia penicillata and Grosmannia europhioides were able to degrade stilbenes and flavonoids faster than E. polonica and grow on minimal growth medium with spruce bark constituents as the only nutrients. Furthermore, beetles avoided medium amended with phenolics but marginally preferred medium colonized by fungi. Taken together our results show that different bark beetle-associated fungi have complementary roles in degrading host metabolites and thus might improve this insect's persistence in well defended host tissues.     

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.     

Spruce beetle outbreak was not driven by drought stress: Evidence from a tree‐ring iso‐demographic approach indicates temperatures were more important

Climate change has amplified eruptive bark beetle outbreaks over recent decades, including spruce beetle (Dendroctonus rufipennis). However, for projecting future bark beetle dynamics there is a critical lack of evidence to differentiate how outbreaks have been promoted by direct effects of warmer temperatures on beetle life cycles versus indirect effects of drought on host susceptibility. To diagnose whether drought‐induced host‐weakening was important to beetle attack success we used an iso‐demographic approach in Engelmann spruce (Picea engelmannii) forests that experienced widespread mortality caused by spruce beetle outbreaks in the 1990s, during a prolonged drought across the central and southern Rocky Mountain region. We determined tree death date demography during this outbreak to differentiate early‐ and late‐dying trees in stands distributed across a landscape within this larger regional mortality event. To directly test for a role of drought stress during outbreak initiation we determined whether early‐dying trees had greater sensitivity of tree‐ring carbon isotope discrimination (?¹³C) to drought compared to late‐dying trees. Rather, evidence indicated the abundance and size of host trees may have modified ?¹³C responses to drought. ?¹³C sensitivity to drought did not differ among early‐ versus late‐dying trees, which runs contrary to previously proposed links between spruce beetle outbreaks and drought. Overall, our results provide strong support for the view that irruptive spruce beetle outbreaks across North America have primarily been driven by warming‐amplified beetle life cycles whereas drought‐weakened host defenses appear to have been a distant secondary driver of these major disturbance events.     

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.     

Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Resistance of fast- and slow-growing subalpine fir to pheromone-induced attack by western balsam bark beetle (Coleoptera: Scolytinae)

Abstract 1 We investigated the resistance of fast‐ and slow‐growing subalpine fir to pheromone‐induced attack by western balsam bark beetle at two sites in the interior of British Columbia, Canada. 2 Attack success by the beetle and subsequent tree mortality were higher in slow‐growing trees than in fast‐growing trees. 3 Fast‐growing trees were more likely to produce secondary resin, and in greater quantities, than slow‐growing trees after attack. 4 Host vigour (indicated by recent radial growth) was positively related to the induced defense response and resistance of subalpine fir to bark beetle attack. These results are discussed in the context of plant defense and plant–herbivore interaction hypotheses. 5 Given the preference of western balsam bark beetle for weakened trees, as well as the reduced defenses and increased mortality rates in less vigorous trees, effective management tactics for this beetle may include strategies that increase the growth and vigour of its subalpine fir host.     

Bacteria in oral secretions of an endophytic insect inhibit antagonistic fungi

Abstract.  1. Colonisation of host trees by an endophytic herbivore, the spruce beetle, Dendroctonus rufipennis , is accompanied by invasion of its galleries by a number of fungal species. Four of these associated species were identified as Leptographium abietinum , Aspergillus fumigatus , Aspergillus nomius , and Trichoderma harzianum .
2.  Trichoderma and Aspergillus significantly reduced spruce beetle survival and reproduction in controlled assays.
3. A previously undescribed behaviour was observed, in which spruce beetle adults exuded oral secretions, especially within fungus-pervaded galleries.
4. These oral secretions inhibited the growth of fungi except A. nomius , and disrupted the morphology of the latter. Administration of these secretions indicated a dose-dependent inhibitory effect.
5. Oral secretions cultured on microbiological media yielded substantial bacterial growth.
6. Filter-sterilised secretions failed to inhibit fungal growth, evidence that the bacteria are responsible for the antifungal activity.
7. Nine bacterial isolates belonging to the Actinobacteria, Firmicutes, Gammaproteobacteria, and Betaproteobacteria taxa were obtained from the secretions.
8. Bacterial isolates showed species-specific inhibitory activity against the four fungi antagonistic to spruce beetle. The bacterium with the strongest fungal inhibition activity was the actinomycete Micrococcus luteus .
9. The production of bark beetle secretions containing bacteria that inhibit fungal growth is a novel finding. This suggests an additional level of complexity to ecological associations among bark beetles, conifers, and microorganisms, and an important adaptation for colonising subcortical tissue.     

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.     

Predicting Ecosystem Resilience to Fire from Tree Ring Analysis in Black Spruce Forests

Climate change has increased the occurrence, severity, and impact of disturbances on forested ecosystems worldwide, resulting in a need to identify factors that contribute to an ecosystem’s resilience or capacity to recover from disturbance. Forest resilience to disturbance may decline with climate change if mature trees are able to persist under stressful environmental conditions that do not permit successful recruitment and survival after a disturbance. In this study, we used the change in proportional representation of black spruce pre- to post-fire as a surrogate for resilience. We explored links between patterns of resilience and tree ring signals of drought stress across topographic moisture gradients within the boreal forest. We sampled 72 recently (2004) burned stands of black spruce in interior Alaska (USA); the relative dominance of black spruce after fire ranged from almost no change (high resilience) to a 90% decrease (low resilience). Variance partitioning analysis indicated that resilience was related to site environmental characteristics and climate–growth responses, with no unique contribution of pre-fire stand composition. The largest shifts in post-fire species composition occurred in sites that experienced the compounding effects of pre-fire drought stress and shallow post-fire organic layer thickness. These sites were generally located at warmer and drier landscape positions, suggesting they are less resilient to disturbance than sites in cool and moist locations. Climate–growth responses can provide an estimate of stand environmental stress to climate change and as such are a valuable tool for predicting landscape variations in forest ecosystem resilience.     

Bark beetle diversity at different spatial scales

To determine how the scale of observation affects ecological patterns we studied bark beetle (Coleoptera, Scolytidae) diversity in southern Finland. A block covering 160 × 160 m of a forest was delimited in four stands of different site types. Each block was divided into 256 squares (10 × 10 m) in which the occurrence of bark beetle species was recorded. In addition, environmental variables describing site type, trees, and breeding material appropriate for bark beetles were measured. The species presence/absence data were combined at different scales of resolution (10 × 10 m. 20 × 20 m, 40 × 40 m, 80 × 80 m, 160 × 160 m). At the finest scale a recently thinned pine stand showed relatively high diversity compared to other study stands due to a few evenly distributed and abundant species. However, the species diversity increased faster toward larger scales in mature spruce stands with several sporadically distributed species. According to logistic regression analyses, breeding material and site characteristics explained the occurrence of most beetle species. However, these variables did not explain the occurrence of the six most frequent species, probably because the factors regulating their distribution and occurrence operate at larger scales.     

Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.

     

Inducibility of chemical defenses in Norway spruce bark is correlated with unsuccessful mass attacks by the spruce bark beetle

Secondary attraction to aggregation pheromones plays a central role in the host colonization behavior of the European spruce bark beetle Ips typographus. However, it is largely unknown how the beetles pioneering an attack locate suitable host trees, and eventually accept or reject them. To find possible biomarkers for host choice by I. typographus, we analyzed the chemistry of 58 Norway spruce (Picea abies) trees that were subsequently either (1) successfully attacked and killed, (2) unsuccessfully attacked, or (3) left unattacked. The trees were sampled before the main beetle flight in a natural Norway spruce-dominated forest. No pheromones were used to attract beetles to the experimental trees. To test the trees' defense potential, each tree was treated in a local area with the defense hormone methyl jasmonate (MeJ), and treated and untreated bark were analyzed for 66 different compounds, including terpenes, phenolics and alkaloids. The chemistry of MeJ-treated bark correlated strongly with the success of I. typographus attack, revealing major chemical differences between killed trees and unsuccessfully attacked trees. Surviving trees produced significantly higher amounts of most of the 39 analyzed mono-, sesqui-, and diterpenes and of 4 of 20 phenolics. Alkaloids showed no clear pattern. Differences in untreated bark were less pronounced, where only 1,8-cineole and (-)-limonene were significantly higher in unsuccessfully attacked trees. Our results show that the potential of individual P. abies trees for inducing defense compounds upon I. typographus attack may partly determine tree resistance to this bark beetle by inhibiting its mass attack.     

ALLOZYME AND MORPHOLOGICAL DIFFERENTIATION OF MOUNTAIN PINE BEETLES DENDROCTONUS PONDEROSAE HOPKINS (COLEOPTERA: SCOLYTIDAE) ASSOCIATED WITH HOST TREE

The purpose of this study was to determine whether mountain pine beetles utilizing different host species were differentiated for either morphological or protein variation. Genetic differentiation among host species has been reported for the southern pine beetle, the Douglas-fir beetle, the jeffrey pine beetle, and the mountain pine beetle. However, in these studies, the host trees were sampled at separate sites, and hence geographic variation and variation due to host tree were confounded. The mountain pine beetle occasionally utilizes three host trees (ponderosa pine, lodgepole pine, and limber pine) at single sites in Colorado. Five polymorphic enzyme loci and six morphological characters were used to describe beetles resident in different hosts. Differentiation within a site among host trees was detected at two of five polymorphic proteins, and for both size and morphological shape. The magnitude of genetic differentiation among hosts within a site was approximately equivalent to the magnitude of differentiation among sites. These data suggest that the species of host tree may be an important biotic factor associated with the genetic structure of bark beetle communities. The results are discussed in terms of their potential role in the process of speciation by host race formation.     

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.     

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.