Bjørn Økland

Research Professor

(+47) 916 28 225
bjorn.okland@nibio.no

Place
Ås H8

Visiting address
Høgskoleveien 8, 1433 Ås

Biography

My main research interests are:
  • Population dynamics of bark beetles
  • Determinants of insect diversity in forests
  • Climate change and insects
  • Invasive species

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Abstract

1 The European spruce bark beetle Ips typographus is a damaging pest on spruce in Europe. Beetle interactions with tree species originating outside the natural range of the beetle are largely unknown and may be unpredictable because trees without a co-evolutionary history with the beetle may lack effective defences. 2 The terpenoid composition and breeding suitability for I. typographus of the historic host Norway spruce Picea abies were compared with two evolutionary naïve spruces of North American origin that are extensively planted in North-West Europe: Sitka spruce Picea sitchensis and Lutz spruce Picea glauca x lutzii. 3 The bark of all three species had a similar chemical composition and similar levels of total constitutive terpenoids, although Norway spruce had higher total induced terpenoid levels. 4 Beetles tunnelling in the three spruce species produced similar amounts of aggregation pheromone. Controlled breeding experiments showed that I. typographus could produce offspring in all three species, with a similar offspring length and weight across species. However, total offspring production was much lower in Sitka and Lutz spruce. 5 Overall, the results of the present study suggest that I. typographus will be able to colonize Sitka and Lutz spruce in European plantations and in native spruce forests in North America if introduced there.

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Abstract

A warmer climate may potentially have a strong effect on the health status of European oak forests by weakening oak trees and facilitating mass reproduction of wood boring insects. We did a laboratory experiment in Slovakia to study the response of major pest beetles of oak and their parasitoids to different temperature regimes as background for predicting climatic effects and improving management tools of European oak forests. With higher temperatures the most important oak pest Scolytus intricatus emerged much earlier, which indicate that completion of a second generation and increased damage further north in European oak forests may be possible. Lower temperatures gave longer larval galleries and more offspring per parents but still lower beetle production due to semivoltine life cycle. For buprestids and longhorn beetles warmer temperatures resulted in more emerging offspring and a shift towards earlier emergence in the same season, but no emergence in the first season indicated that a change to univoltine populations is not likely. Reduced development success of parasitoids at the highest temperatures (25/30 °C) indicates a loss of population regulation for pest beetle populations. A warmer climate may lead to invasion of other population-regulating parasitoids, but also new serious pest may invade. With expected temperature increases it is recommended to use trap trees both in April and in June, and trap trees should be removed within 2 months instead 1 year as described in the current standard.

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Abstract

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.

Abstract

The Eurasian spruce bark beetle Ips typographus is a major forest pest in Europe, capable of mass-attacking and killing mature Norway spruces over extensive areas during outbreaks. Recurring outbreaks over the last few centuries have affected Central and Northern Europe. Outbreaks tend to be periodic and are in many cases triggered by large wind-felling events. For example in Scandinavia several large outbreaks have been triggered by storm disturbances in the last 50 years. In Europe I. typographus is widespread where spruce hosts are found except for in the British Isles. Here we review the identification and biology of this insect and present information about its invasiveness and the potential of success in early detection and control of outbreaks. There are indications that a warmer climate will increase the risk for outbreaks in the northern range of spruce in Europe, where outbreaks so far have been rare. More outbreaks are also expected at the southern margin of the spruce distribution in Europe, where lower than average precipitation seems to generally favor infestations. Establishments outside Eurasia have not been found despite frequent interceptions at ports of entry. Our experiments showed that North American spruce species may be suitable hosts and we conclude that it cannot be ruled out that future establishment can result from repeated imports. The most efficient control option of storm-triggered outbreaks is removal of wind-felled trees before the new generation beetles emerge.

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Abstract

The purpose of this study is to increase the basic understanding of outbreak dynamics in order to improve the management of bark beetle outbreaks. The spruce bark beetle Ips typographus is a major disturbance agent of European forests and is the continent’s most economically and environmentally damaging bark beetle. Outbreaks of the spruce bark beetle are often triggered by large windfall episodes, and we have utilized a unique opportunity to study a Slovakian outbreak where little salvage logging was performed in some areas after a 2.5 million m3 storm-felling in 2004. Our analyses focused on the first five years after the windfall, and we used a combination of empirical data and simulation models to understand the spatial patterns of beetle-killed forest patches developing during the outbreak. The univoltine beetle population used an increasing proportion of the windfelled trees during the two first seasons after the storm, but from the third season onwards our comparisons of inter-patch distance distributions indicated a transition from beetle production largely in windfall areas to a self-sustaining outbreak with infestation patches developing independently of the windthrows. The size of new infestation patches formed after this transition was modeled as a function of beetle pressure, estimated by the proportion of a circle area surrounding new patches that was covered by infestation patches the previous year. Our model results of patch size distribution did not correspond well with the empirical data if patch formation was modeled as a pure dispersal–diffusion process. However, beetle aggregation on individual trees appears to be important for patch development, since good correspondence with empirical data was found when beetle aggregation was incorporated in the modeled dispersal process. The strength of correspondence between the beetle aggregation model and the empirical data varied with the density of aggregation trees in the modeled landscape, and reached a maximum of 83% for a density of three aggregation trees per infestation patch. Our results suggest that efficient removal of windfelled trees up until the start of the second summer after a major windfall is important to avoid a transition into a patch-driven bark beetle outbreak that is very difficult to manage. Our results also indicate that the outcome of a patch-driven outbreak is difficult to predict, since the development of new infestation patches is not a simple function of beetle pressure but is also affected by beetle behavior and local forest conditions.

Abstract

Increasing inter-continental trade with wood chips represents a challenge for phytosanitary authorities, as such trade may lead to pest introductions and invasions with huge impacts on forest ecosystems and economy. Predicting species invasions and their impacts in advance may be difficult, but improved information about potential invasive species ahead of any interceptions is an important precautionary step to reduce the probability of invasions. Here we identify bark- and wood-boring insects that have a potential to become invasive in northern Europe, and that may be introduced by import of deciduous wood chips from North America. The potentially most damaging species belong to the beetle genus Agrilus (Buprestidae), which includes the highly damaging emerald ash borer A. planipennis. We give a brief presentation of this and seven other Agrilus species or subspecies, and review factors of importance for the risk of establishment and potential economic and ecological impacts of these species. We also discuss one Scolytinae, Hylurgopinus rufipes. There are strong indications in the literature that some north European trees are highly susceptible to attack from the selected beetle species. We therefore conclude that because north European trees have not coevolved with these herbivores and thus may lack adequate defenses, most of the identified beetle species are likely to spread in “defense- and enemy-free space” if they are introduced to northern Europe, with considerable economic and ecological consequences.

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Abstract

Six trap models were compared with respect to their ability to capture European spruce bark beetles and technical details in handling and use. All trap models proved to be efficient and gave high captures of beetles during operation for one summer season (2013). Ranking the trap models in descending order of total capture of beetles gave this list: Lindgren trap, Theysohn, prototype-P, Ecotrap, BEKA, and prototype-K. However, there was much variation in trapping results between localities for all trap models, and the estimated means did in most cases not differ significantly between models. The estimated mean capture of the Lindgren trap was significantly higher than for prototype-K and BEKA, but their confidence intervals were wide and they were close to being insignificantly different. The retail price of the traps vary from about 10 € (Ecotrap) to 50-55 € (Lindgren), while prices are not available for the two prototypes, which are not in regular production. Advantages and disadvantages concerning weight, size, robustness, draining of rain water, ease of handling and mounting are commented on in the discussion, and technical details of the traps are presented in a separate appendix (6).

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Abstract

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Abstract

Introductions of the pine wood nematode (PWN), which causes Pine Wilt Disease (PWD), have devastating effects on pine forests in regions with susceptible host trees under suitable climate conditions. Norwegian authorities have proposed a contingency plan if PWN is detected in Norway. We compare the costs of implementing this plan with the costs of further spread and damage of PWN under two climate change scenarios: present and the most likely future climate. With the present climate, PWD will not occur in Norway. Under climatic change, the cost of PWD damage is approximately 0.078–0.157 million NOK (0.01–0.02 million Euros) estimated as net present value with 2 and 4% p.a. discount rate. In contrast, the corresponding costs of implementing the suggested contingency plan will be 1.7–2.2 billion NOK (0.2–0.25 billion Euros). These costs are caused by reduced income from industrial timber production and the costs of the eradication measures. Costs related to reduced recreation or biodiversity are expected to be very high, but are not included in the above estimates. Many of the factors in the analysis are burdened with high uncertainty, but sensitivity analyses indicate that the results are rather robust even for drastic changes in assumptions. The results suggest that there is a need to revise the current PWN contingency plan in Norway.

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Abstract

Increasing inter-continental trade of wood chips for biofuel represents a significant risk of introducing invasive pest species that can cause biome-scale impacts on forest ecosystems. Some potentially invasive species have the capacity to cause high tree mortality on the Eurasian continent and could cause significant impacts on biodiversity and ecosystem functions. Because eradication of established species is difficult, there is a need for scientific studies that can evaluate the reliability of current import control practices to ensure lowest possible risk of establishment of potentially harmful species. We used a stochastic simulation model and sensitivity analyses to evaluate the chance of detecting harmful pests in imported wood chips by sampling according to the current use of internationally accepted standards. As an example, we focused on the North American beetle Agrilus anxius (bronze birch borer) that can cause 100% mortality of European and Asian birch species in North America. We simulated the process from logging in North America to sampling the wood chips upon arrival in Europe. The probability of pest detection for current sampling protocols used by port inspectors was very low (<0.00005), while a 90% chance of detection may require sampling 27 million litres of wood chips per shipload.

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Abstract

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.

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Abstract

Frequent bark beetle outbreaks cause biome-scale impacts in boreal and temperate forests worldwide. Despite frequent interceptions at ports of entry, the most aggressive bark beetle species of Ips and Dendroctonus in North America and Eurasia have failed to establish outside their original home continents. Our experiments showed that Ips typographus can breed in six North American spruce species: Engelmann spruce, white spruce¸ Sitka spruce, Lutz spruce, black spruce and red spruce. This suggests that differences between the Eurasian historical host and North American spruce species are not an insurmountable barrier to establishment of this tree-killing species in North America. However, slightly diminished quality of offspring beetles emerged from the North American spruces could reduce the chance of establishment through an Allee effect. The probabilistic nature of invasion dynamics suggests that successful establishments can occur when the import practice allows frequent arrivals of non-indigenous bark beetles (increased propagule load). Model simulations of hypothetical interactions of Dendroctonus rufipennis and I. typographus indicated that inter-species facilitations could result in more frequent and severe outbreaks than those caused by I. typographus alone. The potential effects of such new dynamics on coniferous ecosystems may be dramatic and extensive, including major shifts in forest structure and species composition, increased carbon emissions and stream flow, direct and indirect impacts on wildlife and invertebrate communities, and loss of biodiversity.

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Abstract

The Eurasian spruce bark beetle, Ips typographus, is one of the major forest insect pests in Europe, capable of mass-attacking and killing mature Norway spruce trees. The initiation and development of a new generation are strongly controlled by temperature and a warmer climate may affect the number of generations that is produced per year and hence the outbreak dynamics. Experimental knowledge regarding reproductive diapause adaptations is, however, too sparse for largescale assessments of future trends. We developed a model description of diapause induction, and used gridded observational temperature data to evaluate multiple combinations of day length and temperature thresholds to find the model parameterisation most coherent with I. typographus monitoring data from Scandinavia. The selected model parameterisation is supported by European literature data, though further experimental studies are required to analyse population specific adaptations and capacity for adjustments to changing climate conditions. Implementing the model description of reproductive diapause in a temperature driven model of bark beetle phenology (swarming activity and development from egg to mature bark beetle), enabled us to assess the length of the late summer swarming period that is a critical determinant of the risk of forest damage. By using regional climate model data we show that higher temperatures can result in increased frequency and length of late summer swarming events, producing a second generation in southern Scandinavia and a third generation in lowland parts of central Europe. Reproductive diapause will not prevent the occurrence of an additional generation per year, but the day length cues may restrict the length of the late summer swarming period.

Abstract

Bark beetles (Coleoptera: Curculionidae, Scolytinae) feed and breed in dead or severely weakened host trees. When their population densities are high, some species aggregate on healthy host trees so that their defences may be exhausted and the inner bark successfully colonized, killing the tree in the process. Here we investigate under what conditions participating with unrelated conspecifics in risky mass attacks on living trees is an adaptive strategy, and what this can tell us about bark beetle outbreak dynamics. We find that the outcome of individual host selection may deviate from the ideal free distribution in a way that facilitates the emergence of tree-killing (aggressive) behavior, and that any heritability on traits governing aggressiveness seems likely to exist in a state of flux or cycles consistent with variability observed in natural populations. This may have implications for how economically and ecologically important species respond to environmental changes in climate and landscape (forest) structure. The population dynamics emerging from individual behavior are complex, capable of switching between `endemic` and `epidemic`’ regimes spontaneously or following changes in host availability or resistance. Model predictions are compared to empirical observations, and we identify some factors determining the occurrence and self-limitation of epidemics.

To document

Abstract

Bark beetles (Coleoptera: Curculionidae, Scolytinae) feed and breed in dead or severely weakened host trees. When their population densities are high, some species aggregate on healthy host trees so that their defences may be exhausted and the inner bark successfully colonized, killing the tree in the process. Here we investigate under what conditions participating with unrelated conspecifics in risky mass attacks on living trees is an adaptive strategy, and what this can tell us about bark beetle outbreak dynamics. We find that the outcome of individual host selection may deviate from the ideal free distribution in a way that facilitates the emergence of tree-killing (aggressive) behavior, and that any heritability on traits governing aggressiveness seems likely to exist in a state of flux or cycles consistent with variability observed in natural populations. This may have implications for how economically and ecologically important species respond to environmental changes in climate and landscape (forest) structure. The population dynamics emerging from individual behavior are complex, capable of switching between “endemic” and “epidemic” regimes spontaneously or following changes in host availability or resistance. Model predictions are compared to empirical observations, and we identify some factors determining the occurrence and self-limitation of epidemics.

Abstract

Today the spruce bark beetle Ips typographus is always univoltine in Northern Europe including Norway and completes development from egg to adult between May and August. Further south in Europe, development is bivoltine with the completion of two generations in most years. A temperature-driven developmental model suggests that by 2070-2100 the voltinism of I. typographus will change dramatically in Norway. If summers become only 2.5°C warmer than today bivoltinism can be expected every single year in the major spruce growing areas in S-Norway. This is likely to have dramatic effects on forestry since two generations per year will give two, instead of one, attack periods each summer. In addition to increasing the number of attacked trees the effect of the attacks may also be more severe, as Norway spruce is more susceptible to beetle attacks later in the summer. However, climate change will probably also change the phenology of Norway spruce and thus its susceptibility to attack by I. typographus and its phytopathogenic fungal associates. We are currently modelling how tree resistance varies with temperature and tree phenology in order to provide more well-founded advice to forest managers on the interaction between bark beetles and tree in a future climate.

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Abstract

The pinewood nematode (PWN) is one of the worst tree-killing exotic pests in East-Asian countries. The first European record of establishment in Portugal in 1999 triggered extensive surveys and contingency plans for eradication in European countries, including immediate removal of large areas of conifer host trees. Using Norway as an example, we applied a simulation model to evaluate the chance of successful eradication of a hypothetical introduction by the current contingency plan in a northern area where wilting symptoms are not expected to occur. Despite a highly variable spread of nematode infestations in space and time, the probability of successful eradication in 20 years was consistently low (mean 0.035, SE 0.02). The low success did not change significantly by varying the biological parameters in sensitivity analyses (SA), probably due to the late detection of infestations by the survey (mean 14.3 years). SA revealed a strong influence of management parameters. However, a high probability of eradication required unrealistic measures: achieving an eradication probability of 0.99 in 20 years required 10,000 survey samples per year and a host tree removal radius of 8,000 m around each detection point.

Abstract

In recent years several forest insects have been recorded as newcomers or as more abundant than before towards northern latitudes and higher elevations in Norway. Such observations are from different groups of forest insect, including herbivorous geometrids, phloeophagous bark beetles and insects in cone seed .......

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Abstract

The current report is a background for a Pest Risk Assessment (PRA) of the bark beetle species Ips amitinus in the PRA area of Norway, following the International Standards for Phytosanitary Measures ISPM 11 (FAO 2004). The report is based on updated information about distribution, pathogenicity and ecological information, which add to a previous PRA for this species. It is concluded that the risk of establishment is high with the current import and management practices. I. amitinus may potentially cause significant tree damage alone or in interaction with Ips typographus; however, the outcome is very much dependent on scenarios of climate development and population dynamics.

Abstract

Pine Wood Nematode (PWN, Bursaphelenchus xylophilus) is the causal organism of Pine Wilt Disease (PWD), the worst forest pest of Japan. In Europe PWN is known to exist in Portugal. The Norwegian Food Safety Authority (Mattilsynet) is concerned about the plant health risks and the consequences to the society if PWN should establish in Norway. Mattilsynet needs a scientific assessment of the proposed measures in a contingency plan for PWN. Mattilsynet also needs the risks connected with recent spread of PWN in Portugal to be evaluated before possible changes can be made in the current phytosanitary policy of Norway. On this background Mattilsynet requested a pest risk assessment of PWN from the Norwegian Scientific Committee for Food Safety (Vitenskapskomiteen for mattrygghet, VKM). To answer the request, VKM commissioned a draft pest risk assessment report from the Norwegian Institute for Agricultural Sciences and Environmental Research (Bioforsk). A working group appointed by VKM’s Panel on Plant Health (Panel 9) has been involved during Bioforsk’s work on the report. VKM’s Panel 9 has used the report as a basis for VKM’s opinion. The current document answers Part 1 of Mattilsynet’s request, and was adopted by Panel 9 on a meeting 3rd September 2008. VKM’s Panel 9 gives the following main conclusions of the risk assessment: The PRA area of this assessment is Norway. PWN is not known to occur in Norway. With present trade pattern the probability of entry of PWN into Norway is expected to be high. The most probable pathway for entry of PWN into Norway would be wood packaging material (WPM). The probability that PWN will establish and spread in Norway is considered as high. With regard to the so-called Pest Free Areas (PFAs) of Portugal, the criteria given in ISPM No. 4 (FAO 1995) for establishing and maintaining PFAs have not been met, and the data available is not sufficient to confirm the existence of PFAs. Acceptance of untreated conifer wood from all parts of Portugal will result in a very high probability of entry and a high probability of establishment and spread of PWN and its vector to Norway. Uncertainty factors: To the best of our knowledge PWN is absent from the PRA area. The beetle M. sutor is regarded as a potential vector or PWN, but this has so far not been demonstrated in nature. The currently low vector densities may retard establishment of the PWN and PWD, but it will probably not stop establishment in a longer perspective. Lack of information on the dynamics of PWN populations in cool climates complicates estimates of the spread of the nematode and PWD. Custom routines may fail in their detection of PWN. Import of a seemingly harmless material might therefore pose an unknown risk. WPM follows consignments of all kinds and is a good example of a hazardous material, which often escapes plant health inspections. Detailed assessments of economic consequences of a possible establishment and spread of PWN in Norway, the effects of global warming and other climate changes on the probability for PWD outbreaks, and the effect of possible phytosanitary measures, will be given in Part 2.

Abstract

Extensive timber imports represent potential introduction pathways for exotic bark beetles (Col: Scolytidae) that may pose ecological hazards and economical risks to native forests. One such species, Ips amitinus Eichh., has been intercepted several times at Norwegian ports of entry in the years since 2002, the year of the first Scandinavian record. Detection of overwintering individuals of I. amitinus at the timber storage site of import timber and preliminary results of a stepwise import model may suggest a high risk of establishment and spread in Norway spruce forests in Scandinavia.Using various modeling approaches, our goal is to reduce the risk of introduction, establishment, and spread of introduced bark beetles. Our objectives are to:model the processes of dispersal and establishment of arriving bark beetlesexplore to what extent an introduced species interacting with native Ips typographus L., the most dominant species in Norway spruce, will lead to stronger and more frequent outbreaks of I. typographusassess potential patterns of spread of newly established bark beetle species and the spatiotemporal outbreak dynamics resulting from interactions between native and introduced species; andadvise on the implications for forest industry and management.Here we present current efforts to model dispersal (objective 1). Dispersal patterns, and hence rates of establishment and spread, may vary considerably depending on dispersal behaviors of insects, such as directionality of movement and aggregation propensity. To assess underlying assumptions of dispersal models, we are using an individual-based model where traits governing dispersal are inherited with random mutations. Individual reproductive success is determined by resource availability and density-dependence in a simulated landscape governed by external forces (e.g., windfellings) and beetle activity (consumption of resources). Evolvable traits include straight line vs. random-walk flight paths and aggregation propensity.Model simulations show that the chance of successful reproduction is greatest for intermediate to high levels of directionality, and that directionality increases over time up to a certain point determined by the landscape features as well as other traits of the species. Assuming limited (local) information in a stochastic landscape, intermediate to high degree of flight directionality is selected for.

Abstract

The detection in 1999 of the pine wood nematode (PWN), Bursaphelenchus xylophilus, in Portugal triggered survey activities in many European countries. With the assumption that PWN would reach frequency 10 times lower than the native B. mucronatus and the requirement of a 95 % confidence limit suggested 2 995 samples to be required for a safe statement on the absence of PWN from the territory surveyed. Samples were taken from 10 circular areas with 50 km diameter erected from a point of import of risk materials. In the period 2000-2006 3 165 wood samples, 2 880 from Pinus sylvestris, 279 from Picea abies and 6 from unknown wood, were collected from 446 logging sites, in 84 municipalities and 13 counties. Of the total material 85 % of the samples came from cutting wastes, timber or lying trees. Wood showing signs of insect activity (incl. Monochamus) formed 73 % of the total material. Nematodes were recorded in 85 % of the samples. The order Rhabditida was most frequent, followed by the orders Aphelenchida, Tylenchida and Dorylaimida. Rhabditid nematodes were equally frequent in pine and spruce, while Aphelenchida (Aphelenchus, Aphelenchoides, Cryptaphelenchus, Seinura and Bursaphelenchus) and Tylenchida (Filenchus, Lelenchus, Ditylenchus, Deladenus and Nothotylenchus) tended to be more common in pinewood. Aphelenchoides was the most common genus. The genus Bursaphelenchus occurred in 1 % of the samples. B. mucronatus was detected in 0,3 % of the samples and most often in cutting waste of pine. The pine wood nematode (PWN), B. xylophilus, was not detected in this survey. The unexpected low natural occurrence of B. mucronatus indicates that the number of potential niches for PWN also is lower than expected, and hence it will be necessary to continue this surveillance program to reach 10 000 samples. The present zone sites in central and south-eastern Norway will be supplemented with 1-2 zone sites in southwestern region of the country. In the future these zone sites will function as permanent observation areas. Care will also be taken to collect all samples from detached wood with signs of Monochamus activity.

Abstract

Model simulations show that an increased frequency in storms and drought periods may result in more frequent and shorter outbreaks of bark beetles. Warmer summers can result in two bark beetle generations per summer instead of one, giving bark beetles the opportunity to attack forests twice in a single year.

Abstract

Studies of defense mechanisms in Norway spruce Picea abies against the spruce bark beetle Ips typographus show that inducible defenses are triggered by unsuccessful beetle attacks. Moreover, spruce trees can be vaccinated against bark beetle attacks through stem bark inoculation with Ceratocystis polonica, an aggressive fungal associate of Ips typographus. Artificial inoculation with certain other fungi has a similar effect, making treated trees a tougher mach for attacking beetles. A similar enhancement of resistance can be achieved by stem surface treatment with methyl jasmonate. Enhanced resistance is associated with the formation of traumatic resin ducts, and mobilization of phenolic substances in living parenchyma cells. The phenolics appear to mix with the traumatic resin. Similar mechanisms are operating in other conifers. We hypothesize that unsuccessful beetle attacks produce a vaccination effect in the trees, enhancing their resistance to later attack. Adding the effect of induced resistance as a model factor shows an immediate effect on the population dynamics of Ips typographus.

Abstract

Detailed analyses of thresholded ecological interactions can improve our understanding of the transition from aperiodic to periodic dynamics. We develop a threshold model of the population dynamics of outbreaking bark beetle populations that alternate between non-epidemic and epidemic behavior. The model involves accumulation of resources during low-density periods and depletion during outbreaks. The transition between the two regimes is caused by disturbance events in the form of major tree felling by wind. The model is analyzed with particular reference to the population dynamics of the spruce bark beetle (Ips typographus) in Scandinavia for which a comprehensive literature allows full parameterization. The fairly constant outbreak lengths and the highly variable waiting time between outbreaks that are seen in the historical records of this species agree well with the predictions of the model. The thresholded resource-depletion dynamics result in substantial variation in the degree of periodicity between stochastic realizations. The completely aperiodic tree colonizations are partly predictable when the timing of the irregular windfall events are known. However, the predictability of inter-outbreak periods is low due to the large variation of cases falling most frequently in the middle between the extremes of purely nonperiodic (erratic) and periodic (cyclic) fluctuations.

Abstract

Pulpwood has been imported to Norway since the beginning of the 20th century. However, exotic plant species hitchhiking with pulpwood were not observed before c.1985. Prior to this the imported timber was debarked, and the chances that diaspores would have attached to a debarked log (compared to a log with bark) are very small.A greenhouse germination experiment based on 385 dm2 of sifted bark from the holds of nine Estonian pulpwood ships yielded 3187 seeds of 201 species of vascular plants, few of which were typical of coniferous forests. Approximately 39% of the hitchhiking seeds did not germinate until after a period of cold treatment.Most of the species germinated in low densities, and those occurring in greater numbers are common and widespread in Norway. Six species were new to Norway: Agrostis clavata, Androsace filiformis, Bidens radiata, Carex montana, Melica picta, and Ranunculus cassubicus.During the last 20 years, pulpwood has been imported to Norway from many countries around the world, e.g. Russia, Scotland, Canada, and Zaire. While the present data do not indicate any immediate threat from aggressively invasive exotics, in order to avoid the introduction of non-native species and reduce the potential for biological invasion, timber should ideally be debarked prior to importation.

Abstract

Ips typographus is considered the most destructive of the bark beetles in the coniferous forests of the Palaearctic region. At low population densities this species breeds in fresh windfalls or dying trees of Norway spruce. At high densities, in contrast, the beetles may show tree-killing outbreaks at regional scales. Spatio-temporal analyses based on pheromone trap data indicated that large windfall events may be a major instigator and synchronizer of beetle outbreaks in areas subjected to regionalized weather systems. The general dominance of lag 1 density dependence of the time series indicated that the beetle populations are constrained by intraspecific competition for breeding substrates. The time series sequence before the large windfelling of 1987 was however declining without significant density dependence. We suggest this to be a transition period, reflecting a drop in carrying capacity due to depletion of susceptible trees during the preceding outbreak period (1970s) and a drop in beetle number to below the density required to kill trees. A variety of organisms exhibit episodes of explosive population growth, triggered by disturbance events. A new general model is constructed to capture the coupling of disturbance events with resource accumulation / depletion. The model is refined and parameterized using the dynamics of the Ips typographus in Scandinavia as a model system. The waiting times between outbreaks of simulated time series were generally longer and more variable than the outbreak lengths, which agree well with the historical records in the last 250 years. Spectral analysis and logspline density plots of waiting times suggest that the transition from aperiodic to periodic population dynamics should be regarded as a continuum.

Abstract

The spruce bark beetle is the most serious pest of mature spruce trees in Eurasia. At low population densities it breeds in weakened or newly dead trees, but at high densities it becomes a tree killer. The exact factors triggering outbreaks are not fully understood, but climatic variables are important candidates. Populations in SE Norway have been monitored since 1979. Various time series methods are used to explore the role of climate in outbreaks of the spruce bark beetle in space and time, and to estimate the distribution of bivoltism under different scenarios of climate change. Large windfall events appear to be a major synchronizer of beetle outbreaks in areas subjected to regionalized weather systems, and the northern border of bivoltism may be moved towards north as a function of increasing temperature. Preliminary models of the population dynamics emphasize the frequency of large windfall events and phenological changes due to temperature increase. Final aims are to estimate the regional risks of forest damage under different scenarios of climate change, and to describe practices that may reduce the impact for forest managers.

Abstract

At least three mechanisms are known to cause synchrony among spatially separated insect populations: 1) dispersal among populations, 2) synchronous stochastic effects, often referred to as the Moran effect, and 3) trophic interactions with other species that are either themselves synchronized or mobile.The present study brings in the role of insect taxa for spatial synchrony. The spatial synchrony observed in several North American and Eurasian epidemic bark beetles was compared with patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect taxa.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 hence more synchronous spatial dynamics in the Lepidoptera. Among the epidemic bark beetles tested, spatial synchrony of outbreaks in the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other species.

Abstract

Spatio-temporal analyses of non-epidemic bark beetle populations may provide insight in dynamics predisposing for outbreaks. The present article presents a spatio-temporal analysis of the population dynamics of Ips typographus based on pheromone trap data from southeast and mid-Norway in the post-epidemic period 19792002. The analyses include regression analyses, hierarchical cluster analysis, and analysis of spatial synchrony of beetle time series and climatic data by means of nonparametric spatial covariance functions. The mean abundance of beetles declined linearly with latitude. In addition, the time series means were higher in areas with high forest productivity and rocky soils predisposed to drought. The time series patterns differed significantly between northern and southern study areas. The regional synchrony of the time series was fairly high (0.38), indicating that some large-scale climatic factor may influence the dynamics. Windfelling was the external variable showing the most parallel pattern of correlation to the beetle dynamics. We thus posit that large windfall events may be a major instigator and synchronizer of beetle outbreaks in areas subjected to regionalized weather systems.

Abstract

Several studies conclude that sun-exposed habitats with sufficient supply of dead wood are essential for many saproxylic and mycetophagous beetle species in forests. This has induced Scandinavian foresters to leave artificial tall stumps on clear-cuts in spruce forests.In the present study of Norway spruce forest, habitat preferences during flight were studied for saproxylic and mycetophagous beetles recorded as sporocarp-visitors of Fomitopsis pinicola and Fomes fomentarius, using 690 randomly placed window traps in 69 sites covering three forest types: clear-cuts, young and oldgrowth stands.Many of the sporocarp-visitors showed a strong preference for oldgrowth forest, avoiding clear-cuts and young replantations. By stepwise linear multiple regression, presence of canopy cover appeared as a major factor while variables associated with sporocarps and dead wood were most often secondary in importance. The results indicate that sufficient canopy cover is important for many sporocarp-visiting beetles in Norway spruce forests

Abstract

Insect species in 20 families of Diptera were collected by an equal number of interception traps in spruce forests of different management practice:young regrowth 5-10 years after logging and replantingmanaged old-growth forestgap opening of old-growth forest with plenty of even-aged dead wooda shady remnant biotope of uncultivated old-growth forest with plenty of dead wood in various levels of decay. Forest cultivation seems to favour saprophagous generalists on behalf of the specialised fungal fauna. The species in mycorrhizal fungi were clearly reduced in young regrowth, while species in wood-inhabiting fungi were most diverse in the remnant biotope with a heterogeneous composition of dead wood.The abundance of saprophagous generalists was significantly increased in the managed forest variants (young regrowth and managed old-growth forest). More studies are necessary to evaluate how the shift in decomposer fauna influence nutrient cycling and other ecosystem processes in forests.

Abstract

Information about larval ecology is fundamental in entomological research; however, in many insect species the larval habitat is still unknown. In the present project, Diptera insects were reared from various microhabitats and substrates of coniferous and deciduous forests of southern Norway. The material included 54 species that have not been reared earlier and 213 new species-microhabitat relationships. Many new records were found in dead wood of common tree species, such as Picea abies, Populus tremula and Fraxinus excelsior. Microhabitats associated with the root zone of windfelled trees showed the highest number of new species-microhabitat relationships.

Abstract

A short survey shows a lack of continuity in research on the dipteran family Milichiidae in Norway. One common species has several records from the last century, while little else is found before recent time and the new material presented in this article. The number of Norwegian species in Milichiidae is hereby raised to five, and in addition one possible new species. Comparisons with species numbers in neighbouring countries give an indication that still new species for Norway may be found. Rearing habitats for Neophyllomyza acyglossa (Villeneuve, 1920) and the possible new species Leptometopa sp. are presented for the first time.

Abstract

The genus Bryomyia comprises altogether eight species in the Palearctic region, including one new species Bryomyia amurensis Mamaev et Økland described in the present article. A revised key to the species of Bryomyia in the Holarctic region is presented.

Abstract

The subfamily ground midges (Lestremiinae) has been poorly studied in Fennoscandia. Until quite recently (1986) only eight species of this group was known from this area. However, the number of species recorded has increased considerably the few last years. This article surveys new and old records of ground midge from Fennoscandia. The survey is based on a new material collected in 49 localities in various parts of Fennoscandia, and on records in previous publications. A list of altogether 73 species of ground midge is presented, including 25 species which are new to Finland, Norway or Sweden.

Abstract

A high trapping efficiency was found both for window traps and trunk-window traps, while the efficiency of extraction cylinders was low. Trunk-window traps are suitable for comparison of different substrates within the same forest environment, while comparative studies of different forest environments are more difficult, due to baiting effects. Window trap captures are more suitable for comparing different forest environments, but are related to ecological conditions over wide areas, and are almost unaffected by substrate conditions in the near surroundings of the traps

Abstract

There is a growing awareness to preserve the biodiversity in the forest ecosystems. A first step in biodiversity research is to find out what species really are found in the forest, using the scientific tool named taxonomy. Still there are species-rich groups of forest insects which have been poorly researched, and which include several unclearities concerning the species identities. One such group is the free-developing gall midges connected with litter, dead wood and fungi in the ground layer of the forest. This papers intends to improve the taxonomy of one genus in this group - Anaretella (Enderlein 1911), including the description of two new species and a key to the current species within the Holarctic region

Abstract

The relationship between the mycetophilid fauna and a set of environmental variables was studied in spruce forests of southeastern Norway. At the stand level, the continuous presence of wood in all decay stages combined with a tree cover appeared to be a major factor for increasing the species richness of mycetophilids. However, the spatial representation of suitable stands in the landscape seems to be particularly important, since the % area of oldgrowth in the surrounding 100 km showed the strongest influence on both species richness and abundance within individual species. The present findings indicate that conservation of a species-rich mycetophilid fauna requires networks of stands within dispersal distances. A reserve network in southern Norway should give special priority to little disturbed forests in the more remote sub-montane areas, since they appear to be very species-rich, and since their strong populations may provide a long-term viability of many mycetophilid species.

Abstract

The diversity in different groups of obligate saproxylic beetles was related to ecological variables at three levels of spatial scale in mature spruce-dominated forest. The variables were connected to: decaying woodwood-inhabiting fungilevel of disturbancelandscape ecologyvegetational structure. Several strong relationships were found at medium (1 km2) and large scales (4 km2), while only weak relationships were found at a small scale (0.16 ha; 1ha=104m2). This may be explained by the local variations in habitat parameters and the high mobilities of many beetle species. Factors connected to decaying wood and wood-inhabiting fungi were clearly the most important factors at all scale levels. In particular, the variables diversity of dead tree parts, number of dead trees of large diameter and number of polypore fungi species increased the species richness of many groups and increased the abundance of many species. Eight species were not recorded below a certain density of decaying wood per 1 or 4 km2. Former extensive cutting was a negative factor at large scale, probably because of decreasing recolonization with increasing distance to source habitats. Thinning reduced the diversity of species associated with birch. The development of guidelines favouring the diversity of saproxylic beetles are discussed below