Characterization of fungal communities associated with the bark beetle Ips typographus varies depending on detection method, location, and beetle population levels

The Eurasian spruce bark beetle Ips typographus and their fungal associates can cause severe damage to Norway spruce forests. In this paper, by using both molecular and cultural methods, we compared fungal assemblages on bark beetles from different locations, characterized by different beetle population levels. Ips typographus was trapped in the western Alps in two outbreak and in two control areas. Sequencing of clone libraries of Internal Transcribed Spacer (ITS) identified 31 fungal Operational Taxonomic Units (OTUs), while fungal isolations yielded 55 OTUs. Only three OTUs were detected by both molecular and cultural methods indicating that both methods are necessary to adequately describe fungal richness. Fungal assemblages on insects from these four and from an additional 12 study sites differed among stands in response to varying ecological conditions and to the limited spreading ability of I. typographus. Ophiostomatoid fungi showed higher diversity in outbreak areas; the pathogenic Ophiostoma polonicum was relatively uncommon, while O. bicolor was the most abundant species. This result was not unexpected, as insects were trapped not at the peak but at the end of the outbreaks and supports the hypothesis of a temporal succession among Ophiostoma species. Ubiquitous endophytes of trees or common airborne fungi were present both in outbreak and in control areas. Wood decaying basidiomycetes were almost never detected on beetles. Yeasts were detected only by molecular analysis, and the OTUs detected matched those reported elsewhere in Europe and in the world, suggesting a very long association between some yeasts and bark beetles.     

Conifer Stored Resources and Resistance to a Fungus Associated with the Spruce Bark Beetle Ips typographus

Bark beetles and associated fungi are among the greatest natural threats to conifers worldwide. Conifers have potent defenses, but resistance to beetles and fungal pathogens may be reduced if tree stored resources are consumed by fungi rather than used for tree defense. Here, we assessed the relationship between tree stored resources and resistance to Ceratocystis polonica, a phytopathogenic fungus vectored by the spruce bark beetle Ips typographus. We measured phloem and sapwood nitrogen, non-structural carbohydrates (NSC), and lipids before and after trees were attacked by I. typographus (vectoring C. polonica) or artificially inoculated with C. polonica alone. Tree resistance was assessed by measuring phloem lesions and the proportion of necrotic phloem around the tree''s circumference following attack or inoculation. While initial resource concentrations were unrelated to tree resistance to C. polonica, over time, phloem NSC and sapwood lipids declined in the trees inoculated with C. polonica. Greater resource declines correlated with less resistant trees (trees with larger lesions or more necrotic phloem), suggesting that resource depletion may be caused by fungal consumption rather than tree resistance. Ips typographus may then benefit indirectly from reduced tree defenses caused by fungal resource uptake. Our research on tree stored resources represents a novel way of understanding bark beetle-fungal-conifer interactions.     

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.     

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.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Relationships among wood‐boring beetles,fungi, and the decomposition of forest biomass

A prevailing paradigm in forest ecology is that wood‐boring beetles facilitate wood decay and carbon cycling, but empirical tests have yielded mixed results. We experimentally determined the effects of wood borers on fungal community assembly and wood decay within pine trunks in the southeastern United States. Pine trunks were made either beetle‐accessible or inaccessible. Fungal communities were compared using culturing and high‐throughput amplicon sequencing (HTAS) of DNA and RNA. Prior to beetle infestation, living pines had diverse fungal endophyte communities. Endophytes were displaced by beetle‐associated fungi in beetle‐accessible trees, whereas some endophytes persisted as saprotrophs in beetle‐excluded trees. Beetles increased fungal diversity several fold. Over forty taxa of Ascomycota were significantly associated with beetles, but beetles were not consistently associated with any known wood‐decaying fungi. Instead, increasing ambrosia beetle infestations caused reduced decay, consistent with previous in vitro experiments that showed beetle‐associated fungi reduce decay rates by competing with decay fungi. No effect of bark‐inhabiting beetles on decay was detected. Platypodines carried significantly more fungal taxa than scolytines. Molecular results were validated by synthetic and biological mock communities and were consistent across methodologies. RNA sequencing confirmed that beetle‐associated fungi were biologically active in the wood. Metabarcode sequencing of the LSU/28S marker recovered important fungal symbionts that were missed by ITS2, though community‐level effects were similar between markers. In contrast to the current paradigm, our results indicate ambrosia beetles introduce diverse fungal communities that do not extensively decay wood, but instead reduce decay rates by competing with wood decay fungi.     

Bark beetles as lidar targets and prospects of photonic surveillance

Forestry is raising concern about the outbreaks of European spruce bark beetle, Ips typographus, causing extensive damage to the spruce forest and timber values. Precise monitoring of these beetles is a necessary step towards preventing outbreaks. Current commercial monitoring methods are catch-based and lack in both temporal and spatial resolution. In this work, light scattering from beetles is characterized, and the feasibility of entomological lidar as a tool for long-term monitoring of bark beetles is explored. Laboratory optical properties, wing thickness, and wingbeat frequency of bark beetles are reported, and these parameters can infer target identity in lidar data. Lidar results from a Swedish forest with controlled bark beetle release event are presented. The capability of lidar to simultaneously monitor both insects and a pheromone plume mixed with chemical smoke governing the dispersal of many insects is demonstrated. In conclusion, entomological lidar is a promising tool for monitoring bark beetles.     

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.     

The influence of Ceratocystis polonica inoculation and methyl jasmonate application on terpene chemistry of Norway spruce,Picea abies

Constitutive and inducible terpene production is involved in conifer resistance against bark beetles and their associated fungi. In this study 72 Norway spruce (Picea abies) were randomly assigned to methyl jasmonate (MJ) application, inoculation with the bluestain fungus Ceratocystis polonica, or no-treatment control. We investigated terpene levels in the stem bark of the trees before treatment, 30 days and one year after treatment using GC–MS and two-dimensional GC (2D-GC) with a chiral column, and monitored landing and attack rates of the spruce bark beetle, Ips typographus, on the trees by sticky traps and visual inspection. Thirty days after fungal inoculation the absolute amount and relative proportion of (+)-3-carene, sabinene, and terpinolene increased and (+)-α-pinene decreased. Spraying the stems with MJ tended to generally increase the concentration of most major terpenes with minor alteration to their relative proportions, but significant increases were only observed for (?)-β-pinene and (?)-limonene. Fungal inoculation significantly increased the enantiomeric ratio of (?)-α-pinene and (?)-limonene 1 month after treatment, whereas MJ only increased that of (?)-limonene. One year after treatment, both MJ and fungal inoculation increased the concentration of most terpenes relative to undisturbed control trees, with significant changes in (?)-β-pinene, (?)-β-phellandrene and some other compounds. Terpene levels did not change in untreated stem sections after treatment, and chemical induction by MJ and C. polonica thus seemed to be restricted to the treated stem section. The enantiomeric ratio of (?)-α-pinene was significantly higher and the relative proportions of (?)-limonene were significantly lower in trees that were attractive to bark beetles compared to unattractive trees. One month after fungal inoculation, the total amount of diterpenes was significantly higher in putative resistant trees with shorter lesion lengths than in putative susceptible trees with longer lesions. Thus, terpene composition in the stem bark may be related to resistance of Norway spruce against I. typographus and C. polonica.     

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.     

Specialization of the olfactory receptor cells in the bark beetleIps typographus and its predatorThanasimus formicarius to bark beetle pheromones and host tree volatiles

Summary Olfactory receptor cells of the spruce bark beetle,Ips typographus, and its predator, the clerid beetleThanasimus formicarius, were studied using electrophysiological techniques. Recordings were made of nerve impulses from single cells and of the summated receptor potential (electroantennogram).Information from bark beetle pheromones and host volatiles is detected by separate olfactory receptor cells inI. typographus. Those which detected bark beetle pheromones responded to only one key substance. Some receptor cells which responded to spruce bark volatiles were strongly activated by one of the synthetic host compounds tested. However, too few host compounds were tested to reach definite conclusions about the specialization of host odour cells. T. formicarius has evolved olfactory receptor cells for bark beetle pheromones. These have similar specificities (specialist types) to those of the bark beetles. Furthermore, the predator has olfactory receptor cells for many bark beetle pheromones. This indicates thatT. formicarius is able to detect and discriminate between many bark beetle species. No significant differences were found between prey and predator cells which responded to host volatiles.     

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.     

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.     

Performance of the tree‐killing bark beetles Ips typographus and Pityogenes chalcographus in non‐indigenous lodgepole pine and their historical host Norway spruce

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When the forest dies: the response of forest soil fungi to a bark beetle-induced tree dieback

Coniferous forests cover extensive areas of the boreal and temperate zones. Owing to their primary production and C storage, they have an important role in the global carbon balance. Forest disturbances such as forest fires, windthrows or insect pest outbreaks have a substantial effect on the functioning of these ecosystems. Recent decades have seen an increase in the areas affected by disturbances in both North America and Europe, with indications that this increase is due to both local human activity and global climate change. Here we examine the structural and functional response of the litter and soil microbial community in a Picea abies forest to tree dieback following an invasion of the bark beetle Ips typographus, with a specific focus on the fungal community. The insect-induced disturbance rapidly and profoundly changed vegetation and nutrient availability by killing spruce trees so that the readily available root exudates were replaced by more recalcitrant, polymeric plant biomass components. Owing to the dramatic decrease in photosynthesis, the rate of decomposition processes in the ecosystem decreased as soon as the one-time litter input had been processed. The fungal community showed profound changes, including a decrease in biomass (2.5-fold in the litter and 12-fold in the soil) together with the disappearance of fungi symbiotic with tree roots and a relative increase in saprotrophic taxa. Within the latter group, successive changes reflected the changing availability of needle litter and woody debris. Bacterial biomass appeared to be either unaffected or increased after the disturbance, resulting in a substantial increase in the bacterial/fungal biomass ratio.     

Colonization of storm gaps by the spruce bark beetle: influence of gap and landscape characteristics

• 1 After storm disturbances, there is a risk for degradation of the quality of fallen trees, and for subsequent tree mortality caused by the spruce bark beetle Ips typographus (L.) (Coleoptera: Curculionidae). Models assessing the risk for bark beetle colonization of different kinds of storm gaps would be a valuable tool for management decisions.
• 2 The present study aimed to determine which gap and landscape characteristics are correlated with the probability of colonization of wind‐felled Norway spruce trees by I. typographus.
• 3 The study included 36 storm gaps, varying in size from three to 1168 wind‐felled spruces, created by the storm Gudrun in southern Sweden in January 2005.
• 4 In the first summer, on average, 5% of the wind‐felled spruces were colonized by I. typographus. The percentage of colonized wind‐felled trees per gap was negatively correlated with the total area of storm gaps within 2000 m in the surrounding forest landscape.
• 5 In the second summer, the proportion of colonized trees increased to 50%. Both gap (mean diameter of wind‐felled trees and basal area of living spruce trees) and landscape variables (amount of spruce forest) were significantly correlated with colonization percentage and explained almost 50% of the variation between gaps.
• 6 There was no relationship between gap area and colonization percentage. This implies that landscapes with many large storm gaps, where logging resources will be most effectively used, should be salvaged first.

     

Reducing spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) emergence for hibernation in central British Columbia by felling infested trees

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Successful reproduction and pheromone production by the spruce bark beetle in evolutionary naïve spruce hosts with familiar terpenoid defences

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Induced responses in stilbenes and terpenes in fertilized Norway spruce after inoculation with blue-stain fungus, Ceratocystis polonica

In conifers, attacks by bark beetles and associated pathogenic fungi cause an induced wound response, which is characterized by accumulation of antifungal compounds and morphological changes that aid wound healing. In this article the stilbene and terpene concentrations of Norway spruce phloem were monitored as symptoms of induced wound responses in relation to changed nutrient conditions caused by fertilization. Plots of mature Norway spruce were fertilized with N, P or NPK. One year after fertilization the trees were artificially infected with Ceratocystis polonica, a pathogenic fungus associated with the bark beetle Ips typographus. The response of stilbenes to fungal inoculation was mainly qualitative. The concentration of stilbene glycosides in the phloem decreased, and in the immediate vicinity of the site of fungal inoculation, stilbene glycosides were less frequent than in mechanically wounded or unwounded phloem. Corresponding stilbene aglycones were most frequent inside the reaction lesion. The concentration of total stilbene aglycones near the inoculation site was significantly lower in N-fertilized trees than in unfertilized trees. Fungal inoculation caused a strong quantitative response in terpenes. The total terpene concentration of the phloem increased significantly, to almost 100 times greater near the inoculation site compared to the constitutive values. N fertilization significantly reduced the total terpene and total stilbene aglycone concentrations near the inoculation sites. Thus, N fertilization may reduce the ability of Norway spruce to defend itself against fungal pathogens.