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.

     

Spatio‐temporal dynamics of an Ips acuminatus outbreak and implications for management

• 1 Understanding spatio‐temporal processes of bark beetle infestations is crucial for predicting beetle behaviour and aiding management decisions aiming to prevent or mitigate tree mortality. We recorded the spatial and temporal distribution of killed trees during the 5‐year period of an Ips acuminatus outbreak.
• 2 Killed trees were always grouped in well‐defined patches (infestation spots). In years of high population density, infestation spots were large and aggregated, whereas, in years of low density, infestation spots were small and weakly aggregated or randomly distributed within the study area.
• 3 Most trees were killed in the spring by beetles that had hibernated but, in some years, trees were also killed in the summer by new‐generation beetles originating from spring attacks. Spring‐killed trees always formed new infestation spots at new locations (i.e. spot proliferation). By contrast, summer‐killed trees always occurred at the edge of active spots established in the spring, thus resulting in spot growth.
• 4 With regard to management strategies, the results obtained in the present study suggest that areas located in close proximity to infestations of the previous year should be prioritized for risk assessment. Because large spots account for most of the observed tree mortality, the cut‐and‐remove method should be focused on these spots as soon as crown discoloration appears in the summer. If applied timely, this strategy will remove the new‐generation beetles originating from the spring attacks before they emerge and also reduce the risk of spot growth.

     

Diapause and overwintering of two spruce bark beetle species

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce‐dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi‐ or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.     

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.     

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

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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.

     

Climate drivers of bark beetle outbreak dynamics in Norway spruce forests

Bark beetles are among the most devastating biotic agents affecting forests globally and several species are expected to be favored by climate change. Given the potential interactions of insect outbreaks with other biotic and abiotic disturbances, and the potentially strong impact of changing disturbance regimes on forest resources, investigating climatic drivers of destructive bark beetle outbreaks is of paramount importance. We analyzed 17 time‐series of the amount of wood damaged by Ips typographus, the most destructive pest of Norway spruce forests, collected across 8 European countries in the last three decades. We aimed to quantify the relative importance of key climate drivers in explaining timber loss dynamics, also testing for possible synergistic effects. Local outbreaks shared the same drivers, including increasing summer rainfall deficit and warm temperatures. Large availability of storm‐felled trees in the previous year was also strongly related to an increase in timber loss, likely by providing an alternative source of breeding material. We did not find any positive synergy among outbreak drivers. On the contrary, the occurrence of large storms reduced the positive effect of warming temperatures and rainfall deficit. The large surplus of breeding material likely boosted I. typographus population size above the density threshold required to colonize and kill healthy trees irrespective of other climate triggers. Importantly, we found strong negative density dependence in I. typographus that may provide a mechanism for population decline after population eruptions. Generality in the effects of complex climatic events across different geographical areas suggests that the large‐scale drivers can be used as early warning indicators of increasing local outbreak probability.     

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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Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

SHORT-TERM DEMOGRAPHIC RESPONSE OF THE RED-BACKED VOLE TO SPRUCE BEETLE INFESTATIONS IN ALASKA

Abstract: How small mammals are affected by habitat changes caused by forest insect epidemics is largely unknown. Our objective was to determine the influence of spruce beetle (Dendroctonus rufipennis) epidemics on the dynamics of northern red-backed vole (Clethrionomys rutilus) populations approximately 10 years post-infestation. We conducted a mark-recapture study on northern red-backed voles for 2 field seasons in the Copper River Basin, Alaska, USA, where recent beetle infestations were widespread. Using the robust sampling design, we produced estimates of vole abundance, survival, and recruitment in 3 locations that varied in their degree of beetle-induced spruce mortality. Vole abundance inversely related to the level of spruce mortality. Vole recruitment showed a larger contribution from both immigration and in situ reproduction in the low infestation site than in the medium and heavy infestation sites. No differences in vole survival were detectable across the 3 locations with varied beetle-induced spruce mortality levels. Measured vole food resources and protective vegetative cover did not vary greatly across infestation levels. Abundance and recruitment parameters indicate a negative change induced by spruce beetle infestations. However, the effect of beetles was not large enough to cause the variation in vole survival. Spruce mortality levels may need to be over 50% before greatly influencing the habitat and the demographics of northern red-backed voles.     

Population dynamics of the spruce bark beetle: a long‐term study

Bark beetle population dynamics is thought to be primarily driven by bottom‐up forces affecting insect performance and host tree resistance. Although there are theoretical predictions and empirical evidences that predation and parasitism may play an important role in driving bark beetle population fluctuations, long‐term studies testing the role of both biotic and abiotic controls on population dynamics are still rare. The aim of the study was to quantify the relative importance of predation, negative density feedback and abiotic factors in driving Ips typographus population dynamics. We analyzed a unique time series of population density of I. typographus and its main predator Thanasimus formicarius over almost two decades in four regions across Sweden. We used a discrete population model and a multi‐model inference approach to evaluate the importance of both bottom up and top down factors. We found that availability of breeding substrates in the form of storm‐felled trees was the main outbreak trigger, while strong intra‐specific competition for host trees was the main endogenous regulating factor. Although temperature‐related metrics are known to have strong individual effect on I. typographus development and number of generations, they did not emerge as important drivers of population dynamics. A positive effect of low summer rainfall was evident only in the region located in the southernmost and warmest part of the spruce distribution range in Sweden. Predator density did not emerge as an important prey regulating factor. As the reported damage from storms seems to have increased across whole Europe, spruce forests are expected to be increasingly susceptible to large outbreaks of I. typographus with important economic and ecological consequences for boreal ecosystems. However, the observed negative density feedback seems to be a natural regulating mechanism that impedes a strong long‐term propagation of the outbreaks.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Negative Feedbacks on Bark Beetle Outbreaks: Widespread and Severe Spruce Beetle Infestation Restricts Subsequent Infestation

Understanding disturbance interactions and their ecological consequences remains a major challenge for research on the response of forests to a changing climate. When, where, and how one disturbance may alter the severity, extent, or occurrence probability of a subsequent disturbance is encapsulated by the concept of linked disturbances. Here, we evaluated 1) how climate and forest habitat variables, including disturbance history, interact to drive 2000s spruce beetle (Dendroctonus rufipennis) infestation of Engelmann spruce (Picea engelmannii) across the Southern Rocky Mountains; and 2) how previous spruce beetle infestation affects subsequent infestation across the Flat Tops Wilderness in northwestern Colorado, which experienced a severe landscape-scale spruce beetle infestation in the 1940s. We hypothesized that drought and warm temperatures would promote infestation, whereas small diameter and non-host trees, which may reflect past disturbance by spruce beetles, would inhibit infestation. Across the Southern Rocky Mountains, we found that climate and forest structure interacted to drive the 2000s infestation. Within the Flat Tops study area we found that stands infested in the 1940s were composed of higher proportions of small diameter and non-host trees ca. 60 years later. In this area, the 2000s infestation was constrained by a paucity of large diameter host trees (> 23 cm at diameter breast height), not climate. This suggests that there has not been sufficient time for trees to grow large enough to become susceptible to infestation. Concordantly, we found no overlap between areas affected by the 1940s infestation and the current infestation. These results show a severe spruce beetle infestation, which results in the depletion of susceptible hosts, can create a landscape template reducing the potential for future infestations.     

Climate change induces multiple risks to boreal forests and forestry in Finland: A literature review

Climate change induces multiple abiotic and biotic risks to forests and forestry. Risks in different spatial and temporal scales must be considered to ensure preconditions for sustainable multifunctional management of forests for different ecosystem services. For this purpose, the present review article summarizes the most recent findings on major abiotic and biotic risks to boreal forests in Finland under the current and changing climate, with the focus on windstorms, heavy snow loading, drought and forest fires and major insect pests and pathogens of trees. In general, the forest growth is projected to increase mainly in northern Finland. In the south, the growing conditions may become suboptimal, particularly for Norway spruce. Although the wind climate does not change remarkably, wind damage risk will increase especially in the south, because of the shortening of the soil frost period. The risk of snow damage is anticipated to increase in the north and decrease in the south. Increasing drought in summer will boost the risk of large‐scale forest fires. Also, the warmer climate increases the risk of bark beetle outbreaks and the wood decay by Heterobasidion root rot in coniferous forests. The probability of detrimental cascading events, such as those caused by a large‐scale wind damage followed by a widespread bark beetle outbreak, will increase remarkably in the future. Therefore, the simultaneous consideration of the biotic and abiotic risks is essential.     

Gene pool state and degree of infestation by bark beetle (<Emphasis Type="Italic">Ips tipographus</Emphasis> L.) of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> L. Karst.) natural populations and planted stands in Moscow region

A comparative analysis of the gene pool state in natural populations and planted stands of Norway spruce and the degree of their infestation by the bark beetle in the Moscow region was conducted taking into account the dynamic state of communities (4 populations, 148 samples, 24 isoenzyme loci). The degree of infestation by the bark beetle of conditionally native communities is 0%; for planted stands, it is 90–100%; and for a short-term community, it is 15–20%. The comparison of “healthy” populations and those infested with bark beetle by average values of observed heterozygosity (H _O) detected no significant differences. However, the test on allelic frequency heterogeneity demonstrated the difference of planted stands from conditionally native populations both by three loci (Fe-2, Idh-1, Mdh-3) and by the totality of 18 polymorphic isoenzyme loci; the short-term population differs from conditionally native population only by two loci. The value of the inbreeding coefficient by the Idh-1 locus is significantly higher in both populations infested with the bark beetle than in “healthy” populations. The results of conducted studies demonstrate the necessity of continuation of the study on the gene pool state in Norway spruce populations owing to the degree of their infestation by the bark beetle along with the study on the dynamic state of the communities; this can provide a key to solving the problem of the forest preservation from pests.     

Sub-cortical Coleoptera in dead pines and spruces: is primeval species composition maintained in managed forests?

The sub-cortical beetle fauna of dead Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) trunks was compared in primeval forests and managed forests in central Finland. The numbers of both individuals and species were higher in the managed forest in spite of the smaller trunk surface area examined. The proportion of rare species was higher in the primeval forest. Although most species occurred both in primeval and managed forests, there were striking differences in the abundance relations: there was only one species (Pytho depressus) in common among the ten most abundant species. The proportion of bark beetles (Scolytidae) was more than 50% in the managed forests, and less than 5% in the primeval forests. The number of species per site was associated with observation date, occurrence of snails and trunk position (standing or lying) in the primeval forest, and with trunk diameter in the managed forests. The occurrence of rare beetle species in dead conifer trunks was related to man's effects on the forest. Although many sub-cortical species can live in managed forests, the fauna differs drastically from that of primeval forests.     

Synchronous rise and fall of bark beetle and parasitoid populations in windthrow areas

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Climate change alters elevational phenology patterns of the European spruce bark beetle (Ips typographus)

The European spruce bark beetle Ips typographus is the most important insect pest in Central European forests. Under climate change, its phenology is presumed to be changing and mass infestations becoming more likely. While several studies have investigated climate effects across a latitudinal gradient, it remains an open question how phenology will change depending on elevation and topology. Knowing how an altered climate is likely to affect bark beetle populations, particularly across diverse topographies and elevations, is essential for adaptive management. We developed a time‐varying distributed delay model to predict the phenology of I. typographus. This approach has the particular advantage of capturing the variability within populations and thus representing its stage structure at any time. The model is applied for three regional climate change scenarios, A1B, A2 and RCP3PD, to the diverse topography of Switzerland, covering a large range of elevations, aspects and slopes. We found a strong negative relationship between voltinism and elevation. Under climate change, the model predicts an increasing number of generations over the whole elevational gradient, which will be more pronounced at low elevations. In contrast, the pre‐shift in spring swarming is expected to be greater at higher elevations. In comparison, the general trend of faster beetle development on steep southern slopes is only of minor importance. Overall, the maximum elevation allowing a complete yearly generation will move upwards. Generally, the predicted increase in number of generations, earlier spring swarming, more aggregated swarming, together with a projected increase in drought and storm events, will result in a higher risk of mass infestations. This will increase the pressure on spruce stands particularly in the lowlands and require intensified management efforts. It calls for adapted long‐term silvicultural strategies to mitigate the loss of ecosystem services such as timber production protection against rockfall and avalanches and carbon storage.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Semiochemical diversity diverts bark beetle attacks from Norway spruce edges

The semiochemical diversity hypothesis (SDH) states that interference with host‐selection from non‐host volatiles (NHV) is an important mechanism for associational resistance. Inhibition of bark beetle attraction to point sources by non‐host volatiles (NHV) is well established and might be a signal serving in host‐selection also at the habitat scale. In forests dominated by Norway spruce in middle and northern Europe (N Slovakia 2006 & 2007, SE Sweden 2007), we applied a blend of NHV and verbenone, released from dispensers fixed at 2 and 6 m height at forest edges with high Ips typographus populations. In Slovakia, three different doses (0.2–0.7 dispensers/m forest edge) were tested in 20‐tree zones of spruce stand edges. The Swedish experiments used only the middle dose. In Slovakia, there was high tree mortality but dispensers with the anti‐attractants reduced killed trees in a dose‐dependent manner. The reduction in tree killing ranged from 35 to 76% compared to untreated zones. Regression analysis of relative tree kill on log dispenser density was highly significant ( = 0.34, corresponding effect size d≈ 0.98). In Sweden, with lower beetle populations, most attacks (99%) were found outside the experimental areas, with high attack rates (15 trees/ha) in a range of 15–30 m from treated groups, indicating an active inhibitory radius exceeding the previous estimates. The SDH as a functional aspect of biodiversity was tested by converting spruce monocultures into an artificial semiochemically mixed forests. The use of NHV provides the only non‐insecticidal method of direct protection of conifer forests. The demonstrated principle of protection is still too expensive for area‐wide use, but viable for high‐value areas (nature reserves). Further development of push–pull strategies or area‐wide applications may prove more cost efficient. In the long‐term, the only sustainable approach is a forest landscape of mixed habitats.