Tree-killing Bark Beetles Research Articles

Spruce Protection against Ips typographus with Anti-Attractant Blend of Tree-Based Semiochemicals: From Small Experimental Plots to Stand Scales

Tree-killing bark beetles require rapid management, such as anti-attractants, to stop the enlargement of attack hot-spots. We tested two newer anti-attractant blends, both without verbenone and one with the addition of trans-4-thujanol, in traps against standard pheromone baits for inhibition of catch. Both blends provided effective catch reduction (>95%). We also tested these anti-attractant blends in tree protection experiments for two years. We had experimental plots with a center of an anti-attractant protected tree zone, with no traditional control area, but we followed tree kills in 10 m wide concentric rings to 100 m. In 2020, we had 12 plots, and 9 plots in 2021. Monitoring by low-strength pheromone traps followed beetle flight averaging 300/trap during the shorter period, August 2020, and 5000/trap during the longer period, May to August 2021. The blends of anti-attractants were 100% effective in avoiding tree mortality in both treated trees and their surroundings. There were no bark beetle attacks on any treated trees, and there was zero tree mortality up to 19 m in 2020, and up to 30 m in 2021, thus full protection to circa 20 m. The density of killed trees then increased from close to zero, over 20 to 50 m, reaching a level of ca 30 (trees/ha) then declined. The spatial pattern of tree mortality on our experimental plots was highly heterogeneous and individual 10m-ring data points on tree kill density could not be statistically separated. In contrast, a non-linear regression model showed a continuous increase of attacks over the distance from the center to a peak ca 60m, followed by a decline. This model agrees partly with the only similar study in the literature, but importantly, it does not give a peak of kill density at distances between the first and second rings close to treatment zones. Such patterns of close-quarter kills have been observed as a “switching” of attack in this and other scolytid systems manipulated by anti-attractants, but not in the present study, likely due to the elimination of verbenone from our blends.

Comparative transcriptomics reveals the conservation and divergence of reproductive genes across three sympatric Tomicus bark beetles

Three tree-killing bark beetles belonging to the genus Tomicus, Tomicus yunnanensis, Tomicus brevipilosus and Tomicus minor (Coleoptera; Curculionidae, Scolytinae), are serious wood-borers with larvae feeding on the phloem tissues of Pinus yunnanensis. The three Tomicus beetles, in some cases, coexist in a same habitat, providing a best system for exploring the conservation and divergence of reproductive genes. Here, we applied comparative transcriptomics and molecular biology approaches to characterize reproductive-related genes in three sympatric Tomicus species. Illumina sequencing of female and male reproductive systems and residual bodies generated a large number of clean reads, representing 185,920,232 sequences in T. yunnanensis, 169,153,404 in T. brevipilosus and 178,493,176 in T. minor that were assembled into 32,802, 56,912 and 33,670 unigenes, respectively. The majority of the genes had detectable expression in reproductive tissues (FPKM >1), particularly those genes in T. brevipilosus accounting for 76.61 % of the total genes. From the transcriptomes, totally 838 genes encoding 463 detoxification enzymes, 339 chemosensory membrane proteins and 36 ionotropic glutamate receptors (iGluRs) were identified, including 622 reproductive tissue-expressed genes. Of these, members of carboxylesterases (COEs), ionotropic receptors (IRs), sensory neuron membrane proteins (SNMPs) and iGluRs were highly conserved in gene numbers and sequence identities across three Tomicus species. Further, expression profiling analyses revealed a number of genes expressed in reproductive tissues and the diverse expression characteristics in these beetles. The results provide evidence for the conservation and differences of reproductive genes among three sympatric closely related beetles, helping understand their different reproductive strategies and the maximization of the reproductive success.

Monoterpene Composition of Phloem of Eastern Larch (Larix laricina (Du Roi) K. Koch) in the Great Lakes Region: With What Must the Eastern Larch Beetle (Dendroctonus simplex LeConte) Contend?

The eastern larch beetle (Dendroctonus simplex LeConte) is the only tree-killing bark beetle that colonizes tamarack, or eastern larch, (Larix laricina (Du Roi) K. Koch) in the Great Lakes region. Historically, outbreaks have been intermittent and of short duration, frequently following predisposing factors such as drought or defoliation. However, over the past two decades, this insect has been in a perpetual state of outbreak in parts of the U.S. Great Lakes region, a deviation from historic norms. From 2001–2021, the insect impacted 300,000 ha, or 60% of the tamarack forests in Minnesota. This activity has prompted renewed interest in the beetle’s chemical ecology, including aspects of host semiochemistry. While foliar chemistry has been well documented in L. laricina, characterization of the monoterpene composition of the phloem has been lacking. We collected phloem samples from 56 tamarack trees across 14 locations in Wisconsin and Minnesota and assessed the relative abundances of the major monoterpenes present using gas chromatography-flame ionization detector (GC-FID). Individual terpenoid components identified included α-pinene (39.4%) and ∆-3-carene (30.0%) followed by several other components in small (<8%) amounts. This knowledge provides a basis for future testing of monoterpene synergists or antagonists in pheromone lures targeting eastern larch beetle and/or its natural enemies.

Linoleic Acid Promotes Emission of Bark Beetle Semiochemicals by Fungal Symbionts

Tree-killing bark beetles in conifer forests vector symbiotic fungi that are thought to help the beetles kill trees. Fungal symbionts emit diverse volatile blends that include bark beetle semiochemicals involved in mating and host localization. In this study, all 12 tested fungal isolates emitted beetle semiochemicals when growing in medium amended with linoleic acid. These semiochemicals included the spiroacetals chalcogran, trans-conophthorin and exo-brevicomin, as well as 2-methyl-3-buten-1-ol, the main aggregation pheromone component of the spruce bark beetle Ips typographus. The emission of these compounds was affected by the type of fatty acid present (linoleic vs. oleic acid). Accumulating evidence shows that the fatty acid composition in conifer bark can facilitate colonization by bark beetles and symbiotic fungi, whereas the fatty acid composition of non-host trees can be detrimental for beetle larvae or fungi. We hypothesize that beetles probe the fatty acid composition of potential host trees to test their suitability for beetle development and release of semiochemicals by symbiotic fungi.

Prevalence and drivers of a tree-killing bark beetle, Ips typographus (Coleoptera, Scolytinae), in international invasion pathways into the USA

Prevalence and drivers of a tree-killing bark beetle, Ips typographus (Coleoptera, Scolytinae), in international invasion pathways into the USA

Number of gallery arms of the tree-killing bark beetle Polygraphus proximus (Coleoptera: Curculionidae: Scolytinae)

• Gallery length increased as the number of gallery arms in a gallery system increased. • Polygraphus proximus laid more eggs in two-armed galleries than in one- and three-armed ones. • Galleries with more than two arms did not have more eggs compared to two-armed ones. • Gallery length with no oviposition increased as the number of gallery arms increased. • The optimal number of arms (in terms of the number of eggs oviposited) might be two. Polygraphus proximus Blandford (Coleoptera: Curculionidae: Scolytinae) is a non-aggressive monogynous bark beetle that has caused mass mortality in fir ( Abies spp.) forests in the last decade. Although gallery structures made by polygynous beetles may influence their reproductive success, the gallery structure and the number of eggs laid by the monogynous tree-killing bark beetle P . proximus has not been investigated in detail in the natural setting of their native range. We, therefore, investigated the length of mother galleries of P . proximus and the number of eggs oviposited by the beetles in relation to gallery systems with different numbers of arms. The number of eggs oviposited in two-armed galleries was significantly greater than that observed in one- and three-armed galleries. Additionally, the length of one- or two-armed galleries with no oviposition was significantly shorter than that in galleries consisting of more than two arms. Our data may suggest that the optimal number of arms in a gallery system (in terms of the number of eggs oviposited) was two.

Great Basin bristlecone pine mortality: Causal factors and management implications

High-elevation five-needle pines are foundational species and iconic components of subalpine forests across western North America. Because they often grow at environmental extremes, high-elevation pines are vulnerable to changing climate conditions. In addition to the direct effects of recent climatic changes, these species are increasingly threatened by biotic disturbances that thrive in the warming and drying conditions now occurring at higher elevations. Among the high-elevation pines, Great Basin bristlecone pine (Pinus longaeva) is revered for its extreme longevity and has been considered an icon of stability during periods of change. Life history strategies of Great Basin bristlecone pine that contribute to its longevity include physiological traits that enhance survival in harsh and dry habitats, and defensive traits that make it less vulnerable than other high-elevation pines to tree-killing bark beetles. Recent increases in growing degree days with no associated increase in precipitation is causing temperature-amplified tree drought stress, while warming temperatures positively influence bark beetle population growth. We report on preliminary investigations into recent and unexpected Great Basin bristlecone pine mortality at two sites, including the potential roles of weather-induced stress and bark beetles. At both sites climatic water deficit (CWD), a cumulative measure of moisture stress, and mean annual temperature increased during the 2010 decade and CWD was the highest in 2020 relative to any time during the past 40 years. Although Great Basin bristlecone pine mortality has not previously been attributed to bark beetles, we observed recent (i.e., 2013 to 2020) bark beetle-attacked trees at both sites, coincident with the timing of increasing temperature and CWD. Few adult beetles were produced, however, and our results support previous research that Great Basin bristlecone pine is a population sink for bark beetles. Because bark beetles are likely not self-sustaining in Great Basin bristlecone pine, bark beetle-caused mortality of this iconic species will most likely occur when it grows mixed with or near other pine species that support bark beetle population growth. We found Ips confusus and Dendroctonus ponderosae attacking Great Basin bristlecone pine in areas where their host trees, P. monophylla and P. flexilis, were also growing. These results suggest that the presence of these infested conifers likely contributed to Great Basin bristlecone pine mortality. We highlight several factors that may be used for prioritizing future research and monitoring to facilitate development of management strategies for protecting this iconic species.

Scientific response to intensifying bark beetle outbreaks in Europe and North America

Tree-killing bark beetles are globally the most destructive forest pests and their impacts have increased in recent decades. Such an increase has been consistently reported from Europe and North America, and it is, with high confidence, driven by climate change. We investigated how the scientific community in both continents responded to this situation by conducting a comprehensive search of the Scopus database from 1970 to 2020. Studies that investigated interactions between climate change and two prominent bark beetles in Europe and North America, the European spruce bark beetle Ips typographus (ESBB) and the mountain pine beetle Dendroctonus ponderosae (MPB), were identified. We used several hierarchical search criteria, starting from general aspects of pest – climate change interactions, to studies with clear implications for management and policies.We found that authors investigating the two bark beetle species mentioned climate change in publications beginning in 1998, and have constituted 8.9 and 13.8 % of all studies on ESBB (N = 987) and MPB (N = 1479) recorded in Scopus. However, only part of these studies addressed climate change as a fundamental or integral part of their research design (59.1 % in ESBB and 38.7 % in MPB). We identified 30 studies on ESBB and 50 studies on MPB which informed efforts towards improving bark beetle management strategies to address climate change-affected ecosystem dynamics. Publications on both insects consistently highlighted the importance of vegetation management aiming to reduce the risk and severity of outbreaks and prevent large-scale population expansion. Only a minor portion of studies placed their findings into the context of relevant policies and legislation, and this connection was particularly lacking in studies on MPB.We conclude that research on bark beetle management under climate change has received inadequate attention and it lags behind observed and foreseen global-scale impacts. We suggest that focused and applied research with clear management implications is needed to develop new climate-adapted and evidence-based management strategies.

Bark Beetle Outbreaks in Europe: State of Knowledge and Ways Forward for Management

Outbreaks of tree-killing bark beetles have reached unprecedented levels in conifer forests in the northern hemisphere and are expected to further intensify due to climate change. In parts of Europe, bark beetle outbreaks and efforts to manage them have even triggered social unrests and political instability. These events have increasingly challenged traditional responses to outbreaks, and highlight the need for a more comprehensive management framework. Several synthesis papers on different aspects of bark beetle ecology and management exist. However, our understanding of outbreak drivers and impacts, principles of ecosystem management, governance, and the role of climate change in the dynamics of ecological and social systems has rapidly advanced in recent years. These advances are suggesting a reconsideration of previous management strategies. We synthesize the state of knowledge on drivers and impacts of bark beetle outbreaks in Europe and propose a comprehensive context-dependent framework for their management. We illustrate our ideas for two contrasting societal objectives that represent the end-members of a continuum of forest management goals: wood and biomass production and the conservation of biodiversity and natural processes. For production forests, we propose a management approach addressing economic, social, ecological, infrastructural, and legislative aspects of bark beetle disturbances. In conservation forests, where non-intervention is the default option, we elaborate under which circumstances an active intervention is necessary, and whether such an intervention is in conflict with the objective to conserve biodiversity. Our approach revises the current management response to bark beetles in Europe and promotes an interdisciplinary social-ecological approach to dealing with disturbances.

The Signature of Endemic Populations in the Spread of Mountain Pine Beetle Outbreaks.

The mountain pine beetle (MPB) is among the most destructive eruptive forest pests in North America. A recent increase in the frequency and severity of outbreaks, combined with an eastward range expansion towards untouched boreal pine forests, has spurred a great interest by government, industry and academia into the population ecology of this tree-killing bark beetle. Modern approaches to studying the spread of the MPB often involve the analysis of large-scale, high-resolution datasets on landscape-level damage to pine forests. This creates a need for new modelling tools to handle the unique challenges associated with large sample sizes and spatial effects. In two companion papers (Koch et al. in Environ Ecol Stat. https://doi.org/10.1007/s10651-020-00456-2 , 2020a; J R Soc Interface 17(170):20200434, 2020b), we explain how the computational challenges of dispersal and spatial autocorrelation can be addressed using separable kernels. In this paper, we use these ideas to capture nonstationary patterns in the dispersal flights of MPB. This facilitates a landscape-level inference of subtle properties of MPB attack behaviour based on aerial surveys of killed pine. Using this model, we estimate the size of the cryptic endemic MPB population, which formerly has been measurable only by means of costly and time-intensive ground surveys.

Distribution pattern of entry holes of the tree-killing bark beetle Polygraphus proximus.

Bark beetles attack their hosts at uniform intervals to avoid intraspecific competition in the phloem. Bark texture and phloem thickness also affect bark beetle attacks, and the bark characteristics are not spatially homogeneous; therefore, the distribution patterns of entry holes can demonstrate an aggregated distribution. Polygraphus proximus Blandford (Coleoptera: Scolytinae) is a non-aggressive phloephagous bark beetle that feeds on Far Eastern firs. They have caused mass mortality in Russia and Japan. However, the distribution pattern of entry holes of P. proximus and spatial relationships with bark characteristics have not been studied. Thus, we investigated the distribution pattern of entry holes of P. proximus. The distribution of entry holes was significantly uniform in most cases. As the attack density increased, an aggregated distribution pattern within a short distance (< 4.0 cm) was observed. The rough bark had a significantly higher number of entry holes than the remaining bark. The distribution pattern of entry holes demonstrated a significantly aggregated spatial association with rough bark. Finally, rough bark around knots had significantly thicker phloem than the remaining barks. These suggest that P. proximus may preferentially attack rough bark to reproduce in the thicker phloem under a rough bark surface.

Interspecific variation in spruce constitutive and induced defenses in response to a bark beetle-fungal symbiont provides insight into traits associated with resistance.

Differences in defensive traits of tree species may predict why some conifers are susceptible to bark beetle-fungal complexes and others are not. A symbiotic fungus (Leptographium abietinum (Peck) M.J. Wingf.) associated with the tree-killing bark beetle (Dendroctonus rufipennis Kirby) is phytopathogenic to host trees and may hasten tree decline during colonization by beetles, but defense responses of mature trees to the fungus have not been experimentally examined. To test the hypothesis that interspecific variation in spruce resistance is explained by defense traits we compared constitutive (bark thickness and constitutive resin ducts) and induced defenses (resin flow, monoterpene composition, concentration, phloem lesion formation and traumatic resin ducts) between two sympatric spruces: Engelmann spruce (Picea engelmannii Parry ex Engelm.-a susceptible host) and blue spruce (Picea pungens Engelm.-a resistant host) in response to fungal inoculation. Four central findings emerged: (i) blue spruce has thicker outer bark and thinner phloem than Engelmann spruce, which may restrict fungal access to phloem and result in less beetle-available resource overall; (ii) both spruce species induce monoterpenes in response to inoculation but blue spruce has higher constitutive monoterpene levels, induces monoterpenes more rapidly, and induces higher concentrations over a period of time consistent with spruce beetle attack duration; (iii) Engelmann and blue spruce differed in the monoterpenes they upregulated in response to fungal inoculation: blue spruce upregulated α-pinene, terpinolene and γ-terpinene, but Engelmann spruce upregulated 3-carene and linalool; and (iv) blue spruce has a higher frequency of constitutive resin ducts and produces more traumatic resin ducts in annual growth increments than Engelmann spruce, though Engelmann spruce produces more resin following aseptic wounding or fungal inoculation. These findings suggest that higher constitutive resin duct densities and monoterpene concentrations, as well as the ability to rapidly induce specific monoterpenes in response to L. abietinum inoculation, are phenotypic traits associated with hosts resistant to spruce beetle colonization.

Cross-scale interaction of host tree size and climatic water deficit governs bark beetle-induced tree mortality

The recent Californian hot drought (2012–2016) precipitated unprecedented ponderosa pine (Pinus ponderosa) mortality, largely attributable to the western pine beetle (Dendroctonus brevicomis; WPB). Broad-scale climate conditions can directly shape tree mortality patterns, but mortality rates respond non-linearly to climate when local-scale forest characteristics influence the behavior of tree-killing bark beetles (e.g., WPB). To test for these cross-scale interactions, we conduct aerial drone surveys at 32 sites along a gradient of climatic water deficit (CWD) spanning 350 km of latitude and 1000 m of elevation in WPB-impacted Sierra Nevada forests. We map, measure, and classify over 450,000 trees within 9 km2, validating measurements with coincident field plots. We find greater size, proportion, and density of ponderosa pine (the WPB host) increase host mortality rates, as does greater CWD. Critically, we find a CWD/host size interaction such that larger trees amplify host mortality rates in hot/dry sites. Management strategies for climate change adaptation should consider how bark beetle disturbances can depend on cross-scale interactions, which challenge our ability to predict and understand patterns of tree mortality.

Mating Systems of the Tree-Killing Bark Beetle Polygraphus proximus (Coleoptera: Curculionidae: Scolytinae).

Polygraphus proximus Blandford (Coleoptera: Curculionidae: Scolytinae) has caused mass mortality of fir (Abies spp. (Pinaceae)) forests across large areas of Russia in the past decade. More recently, mass mortality of A. veitchii Lindl. due to P. proximus infestation has been reported in Japan. This bark beetle species traditionally has been considered to be polygynous because their galleries have multiple gallery arms, and because harem-polygyny is common in the tribe Polygraphini. Although the mating system(s) potentially could have a marked effect on their reproductive success and population dynamics, the reproductive behavior of the tree-killing bark beetle P. proximus has not been investigated in detail in a natural setting in Japan. We, therefore, investigated the number of males and females in a gallery and the number of gallery arms in Abies species in Japan. None of the galleries examined contained more than one male, and 57.2% of the galleries had multiple gallery arms, even though only 2.8% of the galleries contained two females. The findings showed that the typical mating system employed by P. proximus is monogyny and that this species constructs multiple gallery arms in each gallery. In addition, 70.4% of galleries in which the sex of adult beetles could be determined contained no males, and 26.6% contained no females, suggesting that P. proximus males and females re-emerge.

Unravelling the gut bacteriome of Ips (Coleoptera: Curculionidae: Scolytinae): identifying core bacterial assemblage and their ecological relevance

Bark beetles often serve as forest damaging agents, causing landscape-level mortality. Understanding the biology and ecology of beetles are important for both, gathering knowledge about important forest insects and forest protection. Knowledge about the bark beetle gut-associated bacteria is one of the crucial yet surprisingly neglected areas of research with European tree-killing bark beetles. Hence, in this study, we survey the gut bacteriome from five Ips and one non-Ips bark beetles from Scolytinae. Results reveal 69 core bacterial genera among five Ips beetles that may perform conserved functions within the bark beetle holobiont. The most abundant bacterial genera from different bark beetle gut include Erwinia, Sodalis, Serratia, Tyzzerella, Raoultella, Rahnella, Wolbachia, Spiroplasma, Vibrio, and Pseudoxanthomonas. Notable differences in gut-associated bacterial community richness and diversity among the beetle species are observed. Furthermore, the impact of sampling location on the overall bark beetle gut bacterial community assemblage is also documented, which warrants further investigations. Nevertheless, our data expanded the current knowledge about core gut bacterial communities in Ips bark beetles and their putative function such as cellulose degradation, nitrogen fixation, detoxification of defensive plant compounds, and inhibition of pathogens, which could serve as a basis for further metatranscriptomics and metaproteomics investigations.

Flight of the Mountain Pine Beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae: Scolytinae), in Suburban Cheyenne, Wyoming, USA during Summer 2011

The mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae: Scolytinae), is a major tree-killing bark beetle that attacks primarily ponderosa pine, Pinus ponderosa Lawson & C. Lawson, and lodgepole pine, Pinus contorta Douglas ex Loudon (Pinaceae), among other pines, across the western coniferous forests of North America (Negron and Fettig 2014). A continental-scale outbreak of MPB occurred from the late 1990s through the mid- to late 2010s, ranging from the southern Rockies north to British Columbia (Jarvis and Kulakowski 2015; Negron and Fettig 2014). In northern Colorado and southern Wyoming, most of the mortality occurred among lodgepole pine but mortality also occurred among ponderosa pine, whitebark pine, Pinus albicaulis Engelm., and limber pine, Pinus flexilis James. Elevated populations of mountain pine beetles across Wyoming reached their highest levels from 2007-2010. Peak mortality in 2009 affected approximately 500,000 ha. A decade later, populations had markedly declined, with tree mortality occurring on < 200 ha by 2017 (DeSantis 2017).

Molecular underpinnings of methyl jasmonate-induced resistance in Norway spruce.

In response to various stimuli, plants acquire resistance against pests and/or pathogens. Such acquired or induced resistance allows plants to rapidly adapt to their environment. Spraying the bark of mature Norway spruce (Picea abies) trees with the phytohormone methyl jasmonate (MeJA) enhances resistance to tree-killing bark beetles and their associated phytopathogenic fungi. Analysis of spruce chemical defenses and beetle colonization success suggests that MeJA treatment both directly induces immune responses and primes inducible defenses for a faster and stronger response to subsequent beetle attack. We used metabolite and transcriptome profiling to explore the mechanisms underlying MeJA-induced resistance in Norway spruce. We demonstrated that MeJA treatment caused substantial changes in the bark transcriptional response to a triggering stress (mechanical wounding). Profiling of mRNA expression showed a suite of spruce inducible defenses are primed following MeJA treatment. Although monoterpenes and diterpene resin acids increased more rapidly after wounding in MeJA-treated than control bark, expression of their biosynthesis genes did not. We suggest that priming of inducible defenses is part of a complex mixture of defense responses that underpins the increased resistance against bark beetle colonization observed in Norway spruce. This study provides the most detailed insights yet into the mechanisms underlying induced resistance in a long-lived gymnosperm.

Responses of Engelmann spruce to inoculation with Leptographium abietinum, a symbiotic fungus of the North American spruce beetle

Symbiotic fungi associated with tree-killing bark beetles can alter host-tree physiology with consequences for tree survival, and symbiont genetic variation and environmental variability may impact these interactions. Here, we test whether multiple genetically distinct isolates of a symbiotic fungus (Leptographium abietinum (Peck) M.J. Wingf.) associated with North American spruce beetle (Dendroctonus rufipennis (Kirby, 1837)) vary in their ability to manifest defensive responses consistent with disease symptoms in seedlings of Engelmann spruce (Picea engelmannii Parry ex Engelm.), a primary host-tree species for the beetle–fungus complex in North America. Our experiments incorporate variation in both host-tree water availability and host defenses (phloem monoterpene concentration). Three central findings emerged: (i) isolates varied considerably in their effects on host trees — inoculation with L. abietinum isolates from Colorado caused significantly larger phloem lesions than isolates from Wyoming, though all isolates caused phloem oxidation; (ii) neither water availability nor spruce phloem monoterpene concentrations impacted lesion formation; and (iii) both inoculation with L. abietinum and water deficit inhibited the formation of callus tissue at wound sites. We conclude that L. abietinum isolates vary in their virulence and that inoculation is not lethal but may benefit beetles by altering tree defensive responses.

Late Glacial and Holocene records of tree-killing conifer bark beetles in Europe and North America: Implications for forest disturbance dynamics

Outbreaks of conifer bark beetles in Europe and North America have increased in scale and severity in recent decades. In this study, we identify existing fossil records containing bark beetle remains from the end of the Last Glacial Maximum (~14,000 cal. yr BP) to present day using the online databases Neotoma and BugsCEP and literature searches, and compare these data with modern distribution data of selected tree-killing species. Modern-day observational data from the Global Biodiversity Information Facility (GBIF) database was used to map recorded distributions from AD 1750 to present day. A total of 53 fossil sites containing bark beetle remains, from both geological and archeological sites, were found during our searches. Fossil sites were fewer in Europe ( n = 21) than North America ( n = 32). In Europe, 29% of the samples in which remains were found were younger than 1000 cal. yr BP, while in North America, remains were mainly identified from late Glacial (~14,000–11,500 cal. yr BP) sites. In total, the fossil records contained only 8 of 20 species we consider important tree-killing bark beetles in Europe and North America based on their impacts during the last 100 years. In Europe, Ips sexdentatus was absent from the fossil record. In North America, Dendroctonus adjunctus, Dendroctonus frontalis, Dendroctonus jeffreyi, Dendroctonus pseudotsugae, Dryocoetes confusus, Ips calligraphus, Ips confusus, Ips grandicollis, Ips lecontei, Ips paraconfusus, and Scolytus ventralis were absent. Overall, preserved remains of tree-killing bark beetles are rare in the fossil record. However, by retrieving bulk material from new and existing sites and combining data from identified bark beetle remains with pollen, charcoal, tree rings, and geochemistry, the occurrence and dominance of bark beetles, their outbreaks, and other disturbance events can be reconstructed.

Priming of inducible defenses protects Norway spruce against tree-killing bark beetles.

Plants can form an immunological memory known as defense priming, whereby exposure to a priming stimulus enables quicker or stronger response to subsequent attack by pests and pathogens. Such priming of inducible defenses provides increased protection and reduces allocation costs of defense. Defense priming has been widely studied for short-lived model plants such as Arabidopsis, but little is known about this phenomenon in long-lived plants like spruce. We compared the effects of pretreatment with sublethal fungal inoculations or application of the phytohormone methyl jasmonate (MeJA) on the resistance of 48-year-old Norway spruce (Picea abies) trees to mass attack by a tree-killing bark beetle beginning 35days later. Bark beetles heavily infested and killed untreated trees but largely avoided fungus-inoculated trees and MeJA-treated trees. Quantification of defensive terpenes at the time of bark beetle attack showed fungal inoculation induced 91-fold higher terpene concentrations compared with untreated trees, whereas application of MeJA did not significantly increase terpenes. These results indicate that resistance in fungus-inoculated trees is a result of direct induction of defenses, whereas resistance in MeJA-treated trees is due to defense priming. This work extends our knowledge of defense priming from model plants to an ecologically important tree species.