Interactions among Norway spruce, the bark beetle Ips typographus and its fungal symbionts in times of drought

Resilience and functionality of European Norway spruce forests are increasingly threatened by mass outbreaks of the bark beetle Ips typographus promoted by heat, wind throw and drought. Here, we review current knowledge on Norway spruce and I. typographus interactions from the perspective of drought-stressed trees, host selection, colonisation behaviour of beetles, with multi-level effects of symbiotic ophiostomatoid fungi. By including chemo-ecological, molecular and behavioural perspectives, we provide a comprehensive picture on this complex, multitrophic system in the light of climate change. Trees invest carbon into specialised metabolism to produce defence compounds against biotic invaders; processes that are strongly affected by physiological stress such as drought. Spruce bark contains numerous terpenoid and phenolic substances, which are important for bark beetle aggregation and attack success. Abiotic stressors such as increased temperatures and drought affect composition, amounts and emission rates of volatile compounds. Thus, drought events may influence olfactory responses of I. typographus, and further the pheromone communication enabling mass attack. In addition, I. typographus is associated with numerous ophiostomatoid fungal symbionts with multiple effects on beetle life history. Symbiotic fungi degrade spruce toxins, help to exhaust tree defences, produce beetle semiochemicals, and possibly provide nutrition. As the various fungal associates have different temperature optima, they can influence the performance of I. typographus differently under changing environmental conditions. Finally, we discuss why effects of drought on tree-killing by bark beetles are still poorly understood and provide an outlook on future research on this eruptive species using both, field and laboratory experiments

Modelling natural disturbances in forest ecosystems: a review

Natural disturbances play a key role in ecosystem dynamics and are important factors for sustainable forest ecosystem management. Quantitative models are frequently employed to tackle the complexities associated with disturbance processes. Here we review the wide variety of approaches to modelling natural disturbances in forest ecosystems, addressing the full spectrum of disturbance modelling from single events to integrated disturbance regimes. We applied a general, process-based framework founded in disturbance ecology to analyze modelling approaches for drought, wind, forest fires, insect pests and ungulate browsing. Modelling approaches were reviewed by disturbance agent and mechanism, and a set of general disturbance modelling concepts was deduced. We found that although the number of disturbance modelling approaches emerging over the last 15 years has increased strongly, statistical concepts for descriptive modelling are still largely prevalent over mechanistic concepts for explanatory and predictive applications. Yet, considering the increasing importance of disturbances for forest dynamics and ecosystem stewardship under anthropogenic climate change, the latter concepts are crucial tool for understanding and coping with change in forest ecosystems. Current challenges for disturbance modelling in forest ecosystems are thus (i) to overcome remaining limits in process understanding, (ii) to further a mechanistic foundation in disturbance modelling, (iii) to integrate multiple disturbance processes in dynamic ecosystem models for decision support in forest management, and (iv) to bring together scaling capabilities across several levels of organization with a representation of system complexity that captures the emergent behaviour of disturbance regimes. (C) 2010 Elsevier B.V. All rights reserved

European Spruce Bark Beetle, Ips typographus (L.) Males Are Attracted to Bark Cores of Drought-Stressed Norway Spruce Trees with Impaired Defenses in Petri Dish Choice Experiments

The European spruce bark beetle, Ips typographus (L.), is a major pest of Norway spruce. During outbreaks, the beetles can colonize moderately stressed trees via mass attacks mediated by aggregation pheromones, while at endemic population levels, beetles infest trees with impaired defenses. I. typographus introduces ophiostomatoid fungi into the phloem, which can support host colonization. Low-density fungal infections are locally contained by hypersensitive wound reactions; larger necrotic lesions indicate lower tree resistance. Here, we made links between drought stress, susceptibility to fungal infections, and the attractiveness of spruce for host-searching I. typographus males. We sampled bark cores from roofed, non-roofed and untreated control trees of a rainfall exclusion field site. Drought stress was assessed using pre-dawn twig water potentials, and tree defenses were assessed using inoculations with Grosmannia penicillata. Subsequently, we performed Petri dish arena choice tests in the lab, where male beetles could choose between the bark samples of differentially stressed trees. We found that the attractiveness of bark cores increased with drought stress and the extent of hypersensitive wound reactions to fungal infection. Furthermore, beetles stayed longer in those Petri dish sections with the sample of their final choice. The bioassays provide evidence for the primary attraction of male I. typographus to tissues of Norway spruce and preference of beetles for stressed trees

European Spruce Bark Beetle, <i>Ips typographus</i> (L.) Males Are Attracted to Bark Cores of Drought-Stressed Norway Spruce Trees with Impaired Defenses in Petri Dish Choice Experiments

The European spruce bark beetle, Ips typographus (L.), is a major pest of Norway spruce. During outbreaks, the beetles can colonize moderately stressed trees via mass attacks mediated by aggregation pheromones, while at endemic population levels, beetles infest trees with impaired defenses. I. typographus introduces ophiostomatoid fungi into the phloem, which can support host colonization. Low-density fungal infections are locally contained by hypersensitive wound reactions; larger necrotic lesions indicate lower tree resistance. Here, we made links between drought stress, susceptibility to fungal infections, and the attractiveness of spruce for host-searching I. typographus males. We sampled bark cores from roofed, non-roofed and untreated control trees of a rainfall exclusion field site. Drought stress was assessed using pre-dawn twig water potentials, and tree defenses were assessed using inoculations with Grosmannia penicillata. Subsequently, we performed Petri dish arena choice tests in the lab, where male beetles could choose between the bark samples of differentially stressed trees. We found that the attractiveness of bark cores increased with drought stress and the extent of hypersensitive wound reactions to fungal infection. Furthermore, beetles stayed longer in those Petri dish sections with the sample of their final choice. The bioassays provide evidence for the primary attraction of male I. typographus to tissues of Norway spruce and preference of beetles for stressed trees

Expansion of geographic range in the pine processionary moth caused by increased winter temperatures

International audienceGlobal warming is predicted to cause distributional changes in organisms whose geographic ranges are controlled by temperature. We report a recent latitudinal and altitudinal expansion of the pine processionary moth, Thaumetopoea pityocampa, whose larvae build silk nests and feed on pine foliage in the winter. In north-central France (Paris Basin), its range boundary has shifted by 87 km northwards between 1972 and 2004; in northern Italy (Alps), an altitudinal shift of 110–230 m upwards occurred between 1975 and 2004. By experimentally linking winter temperature, feeding activity, and survival of T. pityocampa larvae, we attribute the expansions to increased winter survival due to a warming trend over the past three decades. In the laboratory we determined the minimum nest and night air temperatures required for larval feeding and developed a mechanistic model based on these temperature thresholds. We tested the model in a translocation experiment that employed natural temperature gradients as spatial analogues for global warming. In all transects we transferred colonies of T. pityocampa larvae to sites within zones of historical distribution, recent distribution, and outside the present range. We monitored air and nest temperature, incoming solar radiation, larval phenology, feeding activity, and survival. Early-season temperature effects on phenology were evident, with delayed development of colonies in the more extreme (colder) sites. In the coldest months, our model was consistent with the observed patterns of feeding activity: Feeding was progressively reduced with increasing latitude or elevation, as predicted by the lower number of hours when the feeding threshold was reached, which negatively affected final survival. Insolation raised nest temperature and increased feeding activity on the south but not the north aspect. Prolonged temperature drops below the feeding thresholds occurred at all sites, leading to starvation and partial mortality. Nonetheless, even the most extreme sites still allowed some feeding and, consequently, up to 20% colony survival and successful pupation. Given that the present distribution of the oligophagous T. pityocampa is not constrained by the distribution of its actual or potential hosts, and that warmer winters will cause the number of hours of feeding to increase and the probability of the lower lethal temperature to decrease, we expect the trend of improved survival in previously prohibitive environments to continue, causing further latitudinal and altitudinal expansion. This work highlights the need to develop temperature-based predictive models for future range shifts of winter-limited species, with potential applications in management

Addressing a century-old hypothesis – do pioneer beetles of Ips typographus use volatile cues to find suitable host trees?

Global warming and more frequent climate extremes have caused bark beetle outbreaks of unprecedented scale of these insects in many conifer forests world-wide. Conifers that have been weakened by drought and heat or damaged by storms are highly susceptible to bark beetle infestation. A large proportion of trees with impaired defences provides good conditions for beetle population build-up of beetles, but mechanisms driving host search of pioneer beetles are still uncertain in several species, including the Eurasian spruce bark beetle Ips typographus. Despite a two-century-long history of bark beetle research, we still lack a sufficient understanding of interactions between I. typographus and its host Norway spruce (Picea abies) to forecast future disturbance regimes and forest dynamics. Depending on the scale (habitat or patch) and beetle population state (endemic or epidemic), host selection is likely driven by a combination of pre and postlanding cues, including visual selection or olfactory detection (kairomones). Here, we discuss primary attraction mechanisms and how volatile emission profiles of Norway spruce may provide cues on tree vitality and suitability for attacks by I. typographus, in particular during the endemic phase. We identify several crucial knowledge gaps and provide a research agenda addressing the experimental challenges of such investigations

Report on specific risk analysis in regional forests of Europe under various Forest Management Alternatives

The main objective of specific risk analyses conducted in EFORWOOD project is to develop and implement response functions and quantitative models to evaluate the effects of various levels of forest management - as defined in conceptual framework of FMAs in WP2.1- on the magnitude of damage in several European forests and regional case studies. Analyses focused on five major pests and abiotic hazards that cause two types of damage: tree mortality and reduction in tree growth. The modelling described in this report was based on the hypotheses that (i) FMAs have an effect on some forest stand characteristics (e.g. composition and structure); (ii) some stand characteristics drive susceptibility to damaging agents and hazards; and (iii) some damages result in wood volume loss through tree mortality and reduced tree growth. The report describes and discusses materials, methods and modelling results obtained for 6 specific cases combining 5 regions, 4 different tree species and 5 hazards resulting in wood volume loss