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.

     

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.     

A predictive framework to assess spatio‐temporal variability of infestations by the European spruce bark beetle

Natural disturbances are key factors for the development of forest ecosystems. In forests of central Europe and Scandinavia, the European spruce bark beetle Ips typographus is the most devastating biotic disturbance agent in Norway spruce Picea abies, but our understanding of the factors determining its spatio‐temporal dynamics is still quite limited. To quantify the drivers of bark beetle dynamics, we analyzed a survey dataset with annual resolution that covers 17 yr and 469 forest districts (10 860 km² of forest area) all over Switzerland. We used Poisson log‐normal models in a Bayesian framework to analyze the spatio‐temporal dynamics of bark beetle infestation spots at the forest district level. Bark beetle infestations increased with increasing heat sum (> 8.3°C), volume of standing Norway spruce stock, and the number of infestation spots of the previous year. Precipitation tended to slightly affect the risk of bark beetle infestations. Two major storm events further increased the spatio‐temporal variability of bark beetle infestations. Spruce abundance, storm damage and temperature are known to be important factors influencing the population dynamics of the European spruce bark beetle. Our study is the first to quantify the combined effects of spruce abundance and heat sum, whereby the heat sum turned out to be the most important and consistent predictor. Because our study area encompasses large ecological and climatological gradients, our model is likely to be applicable to Norway spruce forests in other regions of central Europe and Scandinavia.     

Spatio-temporal impact of climate change on the activity and voltinism of the spruce bark beetle, Ips typographus

The spruce bark beetle Ips typographus is one of the major insect pests of mature Norway spruce forests. In this study, a model describing the temperature-dependent thresholds for swarming activity and temperature requirement for development from egg to adult was driven by transient regional climate scenario data for Sweden, covering the period of 1961–2100 for three future climate change scenarios (SRES A2, A1B and B2). During the 20th century, the weather supported the production of one bark beetle generation per year, except in the north-western mountainous parts of Sweden where the climate conditions were too harsh. A warmer climate may sustain a viable population also in the mountainous part; however, the distributional range of I. typographus may be restricted by the migration speed of Norway spruce. Modelling suggests that an earlier timing of spring swarming and fulfilled development of the first generation will significantly increase the frequency of summer swarming. Model calculations suggest that the spruce bark beetle will be able to initiate a second generation in South Sweden during 50% of the years around the mid century. By the end of the century, when temperatures during the bark beetle activity period are projected to have increased by 2.4–3.8 °C, a second generation will be initiated in South Sweden in 63–81% of the years. The corresponding figures are 16–33% for Mid Sweden, and 1–6% for North Sweden. During the next decades, one to two generations per year are predicted in response to temperature, and the northern distribution limit for the second generation will vary. Our study addresses questions applicable to sustainable forest management, suggesting that adequate countermeasures require monitoring of regional differences in timing of swarming and development of I. typographus , and planning of control operations during summer periods with large populations of bark beetles.     

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

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

     

Effects of heat on the dispersal performance of Ips typographus

Under present climate conditions, Ips typographus (L.) is Europe's most critical disturbance agent for mature Norway spruce (Picea abies). With ongoing climate change, the bark beetle will most probably become more prominent as a pest. The aim of this study was to analyse the dispersal performance of I. typographus under various weather conditions, especially hot days with a maximum air temperature above 30°C. In a field study, marked bark beetles were released from breeding logs and could be retraced in traps distributed across the survey area. With daily collections of the trapped beetles, it was possible to analyse the flight activity and the average flight distance of the bark beetles during hot, moderate and cool days. The numbers of daily catches and the average flight distance during the hottest days (air temperature maximum ≥30°C) did not significantly differ from the moderate days (air temperature maximum ≥22°C and <30°C). The numbers of daily catches and the average flight distance during the cool days (air temperature maximum <22°C) were significantly lower than during hot and moderate days. The results give an insight on the dispersal capacity of I. typographus under climate change driven future conditions. Increased air temperatures do not seem to impair the flight performance of I. typographus. A small proportion of cool days during the swarming period even seems to favour dispersal of I. typographus.     

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

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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.     

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.

     

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.     

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.     

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.     

Distribution of Norway spruce bark and wood‐boring beetles along Alpine elevational gradients

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Best environmental predictors of breeding phenology differ with elevation in a common woodland bird species

Temperatures in mountain areas are increasing at a higher rate than the Northern Hemisphere land average, but how fauna may respond, in particular in terms of phenology, remains poorly understood. The aim of this study was to assess how elevation could modify the relationships between climate variability (air temperature and snow melt‐out date), the timing of plant phenology and egg‐laying date of the coal tit (Periparus ater). We collected 9 years (2011–2019) of data on egg‐laying date, spring air temperature, snow melt‐out date, and larch budburst date at two elevations (~1,300 m and ~1,900 m asl) on a slope located in the Mont‐Blanc Massif in the French Alps. We found that at low elevation, larch budburst date had a direct influence on egg‐laying date, while at high‐altitude snow melt‐out date was the limiting factor. At both elevations, air temperature had a similar effect on egg‐laying date, but was a poorer predictor than larch budburst or snowmelt date. Our results shed light on proximate drivers of breeding phenology responses to interannual climate variability in mountain areas and suggest that factors directly influencing species phenology vary at different elevations. Predicting the future responses of species in a climate change context will require testing the transferability of models and accounting for nonstationary relationships between environmental predictors and the timing of phenological events.     

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.

     

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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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.     

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.     

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.     

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.