Volkmar Timmermann
Research Scientist
Authors
Mark A. Anthony Leho Tedersoo Bruno De Vos Luc Croisé Henning Meesenburg Markus Wagner Henning Andreae Frank Jacob Paweł Lech Anna Kowalska Martin Greve Genoveva Popova Beat Frey Arthur Gessler Marcus Schaub Marco Ferretti Peter Waldner Vicent Calatayud Roberto Canullo Giancarlo Papitto Aleksander Marinšek Morten Ingerslev Lars Vesterdal Pasi Rautio Helge Meissner Volkmar Timmermann Mike Dettwiler Nadine Eickenscheidt Andreas Schmitz Nina Van Tiel Thomas W. Crowther Colin AverillAbstract
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Marco Ferretti Arthur Gessler Nathalie Cools Stefan Fleck Rossella Guerrieri Tamara Jakovljević Manuel Nicolas Tiina M. Nieminen Diana Pitar Nenad Potočić Stephan Raspe Marcus Schaub Kai Schwärzel Volkmar Timmermann Monika Vejpustková Lars Vesterdal Petteri Vanninen Peter Waldner Lothar Zimmermann Tanja GM SandersAbstract
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Arne Verstraeten Nicolas Bruffaerts Fabiana Cristofolini Elena Vanguelova Johan Neirynck Gerrit Genouw Bruno De Vos Peter Waldner Anne Thimonier Anita Nussbaumer Mathias Neumann Sue Benham Pasi Rautio Liisa Ukonmaanaho Päivi Merilä Antti-Jussi Lindroos Annika Saarto Jukka Reiniharju Nicholas Clarke Volkmar Timmermann Manuel Nicolas Maria Schmitt Katrin Meusburger Anna Kowalska Idalia Kasprzyk Katarzyna Kluska Łukasz Grewling Małgorzata Malkiewicz Lars Vesterdal Morten Ingerslev Miklós Manninger Donat Magyar Hugues Titeux Gunilla Pihl Karlsson Regula Gehrig Sandy Adriaenssens Agneta Ekebom Åslög Dahl Marco Ferretti Elena GottardiniAbstract
The effects of tree pollen on precipitation chemistry are not fully understood and this can lead to misinterpretations of element deposition in European forests. We investigated the relationship between forest throughfall (TF) element fluxes and the Seasonal Pollen Integral (SPIn) using linear mixed-effects modelling (LME). TF was measured in 1990–2018 during the main pollen season (MPS, arbitrary two months) in 61 managed, mostly pure, even-aged Fagus, Quercus, Pinus, and Picea stands which are part of the ICP Forests Level II network. The SPIn for the dominant tree genus was observed at 56 aerobiological monitoring stations in nearby cities. The net contribution of pollen was estimated as the TF flux in the MPS minus the fluxes in the preceding and succeeding months. In stands of Fagus and Picea, two genera that do not form large amounts of flowers every year, TF fluxes of potassium (K+), ammonium-nitrogen (NH4+-N), dissolved organic carbon (DOC), and dissolved organic nitrogen (DON) showed a positive relationship with SPIn. However- for Fagus- a negative relationship was found between TF nitrate-nitrogen (NO3−-N) fluxes and SPIn. For Quercus and Pinus, two genera producing many flowers each year, SPIn displayed limited variability and no clear association with TF element fluxes. Overall, pollen contributed on average 4.1–10.6% of the annual TF fluxes of K+ > DOC > DON > NH4+-N with the highest contribution in Quercus > Fagus > Pinus > Picea stands. Tree pollen appears to affect TF inorganic nitrogen fluxes both qualitatively and quantitatively, acting as a source of NH4+-N and a sink of NO3−-N. Pollen appears to play a more complex role in nutrient cycling than previously thought.
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Chatchai Kosawang Isabella Børja Maria-Luz Herrero Nina Elisabeth Nagy Lene R. Nielsen Halvor Solheim Volkmar Timmermann Ari HietalaAbstract
Introduction: The ascomycete Hymenoscyphus fraxineus, originating from Asia, is currently threatening common ash (Fraxinus excelsior) in Europe, massive ascospore production from the saprotrophic phase being a key determinant of its invasiveness. Methods: To consider whether fungal diversity and succession in decomposing leaf litter are affected by this invader, we used ITS-1 metabarcoding to profile changes in fungal community composition during overwintering. The subjected ash leaf petioles, collected from a diseased forest and a healthy ash stand hosting the harmless ash endophyte Hymenoscyphus albidus, were incubated in the forest floor of the diseased stand between October 2017 and June 2018 and harvested at 2–3-month intervals. Results: Total fungal DNA level showed a 3-fold increase during overwintering as estimated by FungiQuant qPCR. Petioles from the healthy site showed pronounced changes during overwintering; ascomycetes of the class Dothideomycetes were predominant after leaf shed, but the basidiomycete genus Mycena (class Agaricomycetes) became predominant by April, whereas H. albidus showed low prevalence. Petioles from the diseased site showed little change during overwintering; H. fraxineus was predominant, while Mycena spp. showed increased read proportion by June. Discussion: The low species richness and evenness in petioles from the diseased site in comparison to petioles from the healthy site were obviously related to tremendous infection pressure of H. fraxineus in diseased forests. Changes in leaf litter quality, owing to accumulation of host defense phenolics in the pathogen challenged leaves, and strong saprophytic competence of H. fraxineus are other factors that probably influence fungal succession. For additional comparison, we examined fungal community structure in petioles collected in the healthy stand in August 2013 and showing H. albidus ascomata. This species was similarly predominant in these petioles as H. fraxineus was in petioles from the diseased site, suggesting that both fungi have similar suppressive effects on fungal richness in petiole/rachis segments they have secured for completion of their life cycle. However, the ability of H. fraxineus to secure the entire leaf nerve system in diseased forests, in opposite to H. albidus, impacts the general diversity and successional trajectory of fungi in decomposing ash petioles.
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Volkmar TimmermannAbstract
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Mladen Ognjenović Ivan Seletković Nenad Potočić Mia Marušić Melita Perčec Tadić Mathieu Jonard Pasi Rautio Volkmar Timmermann Lucija Lovreškov Damir UgarkovićAbstract
European beech (Fagus sylvatica L.) forests provide multiple essential ecosystem goods and services. The projected climatic conditions for the current century will significantly affect the vitality of European beech. The expected impact of climate change on forest ecosystems will be potentially stronger in southeast Europe than on the rest of the continent. Therefore, our aim was to use the long-term monitoring data of crown vitality indicators in Croatia to identify long-term trends, and to investigate the influence of current and previous year climate conditions and available site factors using defoliation (DEF) and defoliation change (DDEF) as response variables. The results reveal an increasing trend of DEF during the study period from 1996 to 2017. In contrast, no significant trend in annual DDEF was observed. The applied linear mixed effects models indicate a very strong influence of previous year drought on DDEF, while climate conditions have a weak or insignificant effect on DEF. The results suggest that site factors explain 25 to 30% DEF variance, while similar values of conditional and marginal R2 show a uniform influence of drought on DDEF. These results suggest that DEF represents the accumulated impact of location-specific stressful environmental conditions on tree vitality, while DDEF reflects intense stress and represents the current or recent status of tree vitality that could be more appropriate for analysing the effect of climate conditions on forest trees.
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European ash (Fraxinus excelsior) and narrow-leafed ash (F. angustifolia) are keystone forest tree species with a broad ecological amplitude and significant economic importance. Besides global warming both species are currently under significant threat by an invasive fungal pathogen that has been spreading progressively throughout the continent for almost three decades. Ash dieback caused by the ascomycete Hymenoscyphus fraxineus is capable of damaging ash trees of all age classes and often ultimately leads to the death of a tree after years of progressively developing crown defoliation. While studies at national and regional level already suggested rapid decline of ash populations as a result of ash dieback, a comprehensive survey at European level with harmonized crown assessment data across countries could shed more light into the population decline from a pan-European perspective and could also pave the way for a new conservation strategy beyond national boarders. Here we present data from the ICP Forests Level I crown condition monitoring from 27 countries resulting in > 36,000 observations. We found a substantial increase in defoliation and mortality over time indicating that crown defoliation has almost doubled during the last three decades. Hotspots of mortality are currently situated in southern Scandinavia and north-eastern Europe. Overall survival probability after nearly 30 years of infection has already reached a critical value of 0.51, but with large differences among regions (0.20–0.86). Both a Cox proportional hazard model as well as an Aalen additive regression model strongly suggest that survival of ash is significantly lower in locations with excessive water regime and which experienced more extreme precipitation events during the last two decades. Our results underpin the necessity for fast governmental action and joint rescue efforts beyond national borders since overall mean defoliation will likely reach 50% as early as 2030 as suggested by time series forecasting.
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Volkmar TimmermannAbstract
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Volkmar TimmermannAbstract
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Lena Wohlgemuth Pasi Rautio Bernd Ahrends Alexander Russ Lars Vesterdal Peter Waldner Volkmar Timmermann Nadine Eickenscheidt Alfred Fürst Martin Greve Peter Roskams Anne Thimonier Manuel Nicolas Anna Kowalska Morten Ingerslev Päivi Merilä Sue Benham Carmen Iacoban Günter Hoch Christine Alewell Martin JiskraAbstract
Despite the importance of vegetation uptake of atmospheric gaseous elemental mercury (Hg(0)) within the global Hg cycle, little knowledge exists on the physiological, climatic, and geographic factors controlling stomatal uptake of atmospheric Hg(0) by tree foliage. We investigate controls on foliar stomatal Hg(0) uptake by combining Hg measurements of 3569 foliage samples across Europe with data on tree species' traits and environmental conditions. To account for foliar Hg accumulation over time, we normalized foliar Hg concentration over the foliar life period from the simulated start of the growing season to sample harvest. The most relevant parameter impacting daily foliar stomatal Hg uptake was tree functional group (deciduous versus coniferous trees). On average, we measured 3.2 times higher daily foliar stomatal Hg uptake rates in deciduous leaves than in coniferous needles of the same age. Across tree species, for foliage of beech and fir, and at two out of three forest plots with more than 20 samples, we found a significant (p<0.001) increase in foliar Hg values with respective leaf nitrogen concentrations. We therefore suggest that foliar stomatal Hg uptake is controlled by tree functional traits with uptake rates increasing from low to high nutrient content representing low to high physiological activity. For pine and spruce needles, we detected a significant linear decrease in daily foliar stomatal Hg uptake with the proportion of time during which water vapor pressure deficit (VPD) exceeded the species-specific threshold values of 1.2 and 3 kPa, respectively. The proportion of time within the growing season during which surface soil water content (ERA5-Land) in the region of forest plots was low correlated negatively with foliar Hg uptake rates of beech and pine. These findings suggest that stomatal uptake of atmospheric Hg(0) is inhibited under high VPD conditions and/or low soil water content due to the regulation of stomatal conductance to reduce water loss under dry conditions. Other parameters associated with forest sampling sites (latitude and altitude), sampled trees (average age and diameter at breast height), or regional satellite-observation-based transpiration product (Global Land Evaporation Amsterdam Model: GLEAM) did not significantly correlate with daily foliar Hg uptake rates. We conclude that tree physiological activity and stomatal response to VPD and soil water content should be implemented in a stomatal Hg model to assess future Hg cycling under different anthropogenic emission scenarios and global warming.
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Mladen Ognjenovic Ivan Seletkovic Mia Marušic Mathieu Jonard Pasi Rautio Volkmar Timmermann Melita Percec Tadic Miran Lanšcak Damir Ugarkovic Nenad PotocicAbstract
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Authors
Ari Hietala Ahto Agan Nina Elisabeth Nagy Isabella Børja Volkmar Timmermann Rein Drenkhan Halvor SolheimAbstract
The populations of European ash and its harmless fungal associate Hymenoscyphus albidus are in decline owing to ash dieback caused by the invasive Hymenoscyphus fraxineus, a fungus that in its native range in Asia is a harmless leaf endophyte of local ash species. To clarify the behavior of H. albidus and its spatial and temporal niche overlap with the invasive relative, we used light microscopy, fungal species-specific qPCR assays, and PacBio long-read amplicon sequencing of the ITS1-5.8S-ITS2 region to examine fungal growth and species composition in attached leaves of European ash. The plant material was collected from a healthy stand in central Norway, where ash saplings in late autumn showed leaflet vein necrosis like that commonly related to H. fraxineus. For reference, leaflet samples were analyzed from stands with epidemic level of ash dieback in southeastern Norway and Estonia. While H. albidus was predominant in the necrotic veins in the healthy stand, H. fraxineus was predominant in the diseased stands. Otherwise, endophytes with pathogenic potential in the genera Venturia (anamorph Fusicladium), Mycosphaerella (anamorph Ramularia), and Phoma, and basidiomycetous yeasts formed the core leaflet mycobiome both in the healthy and diseased stands. In necrotic leaf areas with high levels of either H. albidus or H. fraxineus DNA, one common feature was the high colonization of sclerenchyma and phloem, a region from which the ascomata of both species arise. Our data suggest that H. albidus can induce necrosis in ash leaves, but that owing to low infection pressure, this first takes place in tissues weakened by autumn senescence, 1–2 months later in the season than what is characteristic of H. fraxineus at an epidemic phase of ash dieback. The most striking difference between these fungi would appear to be the high fecundity of H. fraxineus. The adaptation to a host that is phylogenetically closely related to European ash, a tree species with high occurrence frequency in Europe, and the presence of environmental conditions favorable to H. fraxineus life cycle completion in most years may enable the build-up of high infection pressure and challenge of leaf defense prior to autumn senescence.
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In 2021, mean defoliation remained at approximately the same level as in 2020 with no change for broadleaves and only a very slight increase for conifers. Deciduous temperate oaks had the highest increase in mean defoliation (+1.4%), while common beech had the largest decrease (-1.7%). Based on the data of the past 20 years, trends show a considerable increase in defoliation of Austrian pine and evergreen oaks (7.1% and 6.7%, respectively). On the other hand, the increase in defoliation for deciduous temperate oaks (2.6%) and common beech (3.4%) has been relatively low and the trend for Scots pine and Norway spruce shows a moderate increase in defoliation of 4.3 and 3.8%, respectively. No trend was detected for deciduous (sub-) Mediterranean oaks. There was again a decrease in the number of observed damage symptoms compared to last year. As in previous years, the number of recorded damage symptoms per assessed tree was substantially higher for broadleaves than for conifers. Insects, abiotic causes, and fungi were the most common damage agent groups for all species, comprising altogether more than half of all damage records. Tree mortality increased again slightly in 2021, mainly due to abiotic factors.
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Mladen Ognjenovic Ivan Seletkovic Nenad Potocic M. Perčec-Tadić R. Sokol Jurković M. Marušić K. Sever D. Ugarković Volkmar Timmermann Pasi Rautio Mathieu JonardAbstract
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Volkmar TimmermannAbstract
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Nenad Potocic Volkmar Timmermann Mladen Ognjenovic Anne-Katrin Prescher Kai Schwärzel Marco FerrettiAbstract
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Volkmar TimmermannAbstract
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Arne Verstraeten Elena Gottardini Nicolas Bruffaerts Fabiana Cristofolini Elena Vanguelova Johan Neirynck Gerrit Genouw Bruno de Vos Peter Waldner Anne Thimonier Anita Nussbaumer Mathias Neumann Sue Benham Pasi Rautio Liisa Ukonmaanaho Päivi Merilä Annika Saarto Jukka Reiniharju Peter Roskams Geert Sioen Nathalie Cools Nicholas Clarke Volkmar Timmermann Hans-Peter Dietrich Manuel Nicolas Maria Schmitt Katrin Meusburger Silvio Schüler Anna Kowalska Idalia Kasprzyk Katarzyna Kluska Łukasz Grewling Joanna Święta-Musznicka Małgorzata Latałowa Marcelina Zimny Małgorzata Malkiewicz Lars Vesterdal Miklós Manninger Donat Magyar Hugues Titeux Gunilla Pihl-Karlsson Marco FerrettiAbstract
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Volkmar TimmermannAbstract
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The ascomycete Hymenoscyphus fraxineus has spread across most of the host range of European ash with a high level of mortality, causing important economic, cultural and environmental effects. We present a novel method combining a Monte-Carlo approach with a generalised additive model that confirms the importance of meteorology to the magnitude and timing of H. fraxineus spore emissions. The variability in model selection and the relative degree to which our models are over- or under-fitting the data has been quantified. We find that both the daily magnitude and timing of spore emissions are affected by meteorology during and prior to the spore emission diurnal peak. We found the daily emission magnitude has the strongest associations to weekly average net radiation and leaf moisture before the emission, soil temperature during the day before emission and net radiation during the spore emission. The timing of the daily peak in spore emissions has the strongest associations to net radiation both during spore emission and in the day preceding the emission. The seasonal peak in spore emissions has a near-exponential increase/decrease, and the mean daily emission peak is approximately Gaussian.
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Volkmar TimmermannAbstract
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Volkmar TimmermannAbstract
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The scope of this study was to provide an update on fluoride (F) emission effects on vegetation around three aluminium smelters. We visited Årdal and Sunndal smelters in 2019-2020 and Mosjøen in 2020, assessed and documented the visual symptoms of F-damage on vegetation and related these to detected values of F in plant tissue. Three plant species showed qualities as useful indicators: Rowan, pine and St. John’s wort. Because male-fern accumulated extreme F-values and showed clear grazing damage, the monitoring of this species may be warranted because of the potential health hazard for the grazing animals. In Årdal and Sunndal, during 2019 and 2020, we detected the highest F-values in male-fern, ranging from 94 to 925 mg F/kg. In rowan, the highest F-concentration was detected in trees growing within the Årdal smelter (1161 mg F/kg) but on all other locations the F-concentrations in rowan ranged from 4 to 327 mg F/kg. In pine, the F-concentrations ranged from 6-351 mg F/kg for all needle ages, but older needles always accumulated more F than younger ones. In St. John’s wort the accumulated F-values ranged from 10-84 mg F/kg. At all smelters there was a gradient of decreasing F-concentration in vegetation with increasing distance from the smelter. F-emissions in Årdal (12 and 11 kg F/hour in 2019 and 2020, respectively) and in Sunndal during 2019 (12 kg F/hour) were only slightly higher than the recommended limits (10 kg F/hour) for damage on vegetation, while in Mosjøen the F-emissions were 7 kg F/hour in 2020. The presence of F-damage on vegetation was consistent with the reported emission-levels. On basis of this evaluation, reductions in emissions are still advisable in Årdal and Sunndal, while the situation is acceptable in Mosjøen.
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Olalla Díaz-Yáñez Blas Mola-Yudego Volkmar Timmermann Mari Mette Tollefsrud Ari Hietala Jonas OlivaAbstract
Determining the impacts of invasive pathogens on tree mortality and growth is a difficult task, in particular in the case of species occurring naturally at low frequencies in mixed stands. In this study, we quantify such effects by comparing national forest inventory data collected before and after pathogen invasion. In Norway, Fraxinus excelsior is a minor species representing less than 1% of the trees in the forests and being attacked by the invasive pathogen Hymenoscyphus fraxineus since 2006. By studying deviations between inventories, we estimated a 74% higher-than-expected average ash mortality and a 13% slower-than-expected growth of the surviving ash trees, indicating a lack of compensation by the remaining ash. We could confidently assign mortality and growth losses to ash dieback as no mortality or growth shifts were observed for co-occurring tree species in the same plots. The mortality comparisons also show regional patterns with higher mortality in areas with the longest disease history in Norway. Considering that ash is currently mostly growing in mixed forests and that no signs of compensation were observed by the surviving ash trees, a significant habitat loss and niche replacement could be anticipated in the mid-term.
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Ahto Agan Rein Drenkhan Kalev Adamson Leho Tedersoo Halvor Solheim Isabella Børja Iryna Matsiakh Volkmar Timmermann Nina Elisabeth Nagy Ari HietalaAbstract
European ash (Fraxinus excelsior) is threatened by the invasive ascomycete Hymenoscyphus fraxineus originating from Asia. Ash leaf tissues serve as a route for shoot infection but also as a sporulation substrate for this pathogen. Knowledge of the leaf niche partitioning by indigenous fungi and H. fraxineus is needed to understand the fungal community receptiveness to the invasion. We subjected DNA extracted from unwashed and washed leaflets of healthy and diseased European ash to PacBio sequencing of the fungal ITS1-5.8S-ITS2 rDNA region. Leaflets from co-inhabiting rowan trees (Sorbus aucuparia) served as a reference. The overlap in leaflet mycobiomes between ash and rowan was remarkably high, but unlike in rowan, in ash leaflets the sequence read proportion, and the qPCR-based DNA amount estimates of H. fraxineus increased vigorously towards autumn, concomitant with a significant decline in overall fungal richness. The niche of ash and rowan leaves was dominated by epiphytic propagules (Vishniacozyma yeasts, the dimorphic fungus Aureobasidion pullulans and the dematiaceous hyphomycete Cladosporium ramotenellum and H. fraxineus), and endophytic thalli of biotrophs (Phyllactinia and Taphrina species), the indigenous necrotroph Venturia fraxini and H. fraxineus. Mycobiome comparison between healthy and symptomatic European ash leaflets revealed no significant differences in relative abundance of H. fraxineus, but A. pullulans was more prevalent in symptomatic trees. The impacts of host specificity, spatiotemporal niche partitioning, species carbon utilization profiles and life cycle traits are discussed to understand the ecological success of H. fraxineus in Europe. Further, the inherent limitations of different experimental approaches in the profiling of foliicolous fungi are addressed.
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Arne Verstraeten Elena Gottardini Nicolas Bruffaerts Bruno De Vos Elena Vanguelova Fabiana Cristofolini Sue Benham Pasi Rautio Liisa Ukonmaanaho Päivi Merilä Annika Saarto Peter Waldner Marijke Hendrickx Gerrit Genouw Peter Roskams Nathalie Cools Johan Neirynck Anita Nussbaumer Mathias Neumann Nicholas Clarke Volkmar Timmermann Karin Hansen Hans-Peter Dietrich Manuel Nicolas Maria Schmitt Anne Thimonier Katrin Meusburger Silvio Schüler Anna KowalskaAbstract
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Arne Verstraeten Elena Gottardini Nicolas Bruffaerts Bruno De Vos Elena Vanguelova Fabiana Cristofolini Gerrit Genouw Anita Nussbaumer Mathias Neumann Sue Benham Pasi Rautio Liisa Ukonmaanaho Päivi Merilä Annika Saarto Jukka Reiniharju Peter Waldner Marijke Hendrickx Peter Roskams Nathalie Cools Johan Neirynck Arthur De Haeck Yvan De Bodt Geert Sioen Nicholas Clarke Volkmar Timmermann Karin Hansen Hans-Peter Dietrich Manuel Nicolas Maria Schmitt Anne Thimonier Katrin Meusburger Silvio Schüler Anna Kowalska Idalia Kasprzyk Katarzyna Borycka Łukasz Grewling Joanna Święta-Musznicka Małgorzata Latałowa Marcelina Zimny Małgorzata Malkiewicz Lars Vesterdal Iben Margrete Thomsen Miklós Manninger Donat Magyar Gergely Mányoki Hugues TiteuxAbstract
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Ari Hietala Isabella Børja Hugh B. Cross Nina Elisabeth Nagy Jørn Henrik Sønstebø Volkmar Timmermann Adam Vivian-Smith Halvor SolheimAbstract
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Ari Hietala Isabella Børja Hugh Cross Nina Elisabeth Nagy Halvor Solheim Volkmar Timmermann Adam Vivian-SmithAbstract
European ash (Fraxinus excelsior), a keystone species with wide distribution and habitat range in Europe, is threatened at a continental scale by an invasive alien ascomycete, Hymenoscyphus fraxineus. In its native range of Asia, this fungus is a leaf endophyte with weak parasitic capacity and robust saprobic competence in local ash species that are closely related to European ash. In European ash, H. fraxineus has a similar functional role as in Asia, but the fungus also aggressively kills shoots, resulting in crown dieback and tree death. H. fraxineus is a typical invasive species, as its spread relies on high propagule pressure. While crown dieback of European ash is the most obvious symptom of ash dieback, the annual colonization of ash leaves is a crucial key dependency for the invasiveness of H. fraxineus, since its fruiting bodies are formed on overwintered leaf vein tissues in soil debris. Leaves of European ash host a wide range of indigenous epiphytes, endophytes, facultative parasites and biotrophic fungi, including Hymenoscyphus albidus, a relative of H. fraxineus that competes for the same sporulation niche as the invader. At face value, leaves of European ash are colonized by a large and diverse indigenous mycobiome. In order to understand why this invader became successful in Europe, we discuss and summarize the current knowledge of diversity, seasonal dynamics and traits of H. fraxineus and indigenous fungi associated with leaves of European ash.
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Anita Nussbaumer Peter Waldner Vladislav Apuhtin Fatih Aytar Sue Benham Filippo Bussotti Johannes Eichhorn Nadine Eickenscheidt Petr Fabianek Lutz Falkenried Stefan Leca Martti Lindgren María José Manzano Serrano Stefan Neagu Seppo Nevalainen Jozef Pajtik Nenad Potočić Pasi Rautio Geert Sioen Vidas Stakėnas Celal Tasdemir Iben Margrete Thomsen Volkmar Timmermann Liisa Ukonmaanaho Arne Verstraeten Sören Wulff Arthur GesslerAbstract
Mast seeding, the synchronised occurrence of large amounts of fruits and seeds at irregular intervals, is a reproductive strategy in many wind-pollinated species. Although a series of studies have investigated mast year (MY) patterns in European forest tree species at the regional scale, there are few recent evaluations at a European scale on the impact of weather variables (weather cues) and resource dynamics on mast behaviour. Thus the main objective of this study is to investigate the impact of specific weather conditions, as environmental drivers for MYs, on resources in Fagus sylvatica L., Quercus petraea (Matt.)Liebl., Quercus robur L., Picea abies (L.) Karst. and Pinus sylvestris L. at a European level and to explore the robustness of the relationships in smaller regions within Europe. Data on seed production originating from the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) were analysed. Three beta regression models were applied to investigate the impact of seasonal weather variables on MY occurrence, as well as the influence of fruiting intensity levels in the years prior to MYs. Resource dynamics are analysed at three different spatial scales (continent, countries and ecoregions). At a European scale, important weather cues for beech MYs were a cold and wet summer two years before a MY, a dry and warm summer one year before a MY and a warm spring in the MY. For spruce, a cold and dry summer two years prior to a MY and a warm and dry summer in the year before the MY showed the strongest associations with the MY. For oak, high spring temperature in the MY was the most important weather cue. For beech and spruce, and to some extent also for oak species, the best fitting models at European scale were well reflected by those found at smaller scales. For pine, best fitting models were highly diverse concerning weather cues. Fruiting levels were high in all species two years before the MY and also high one year before the MY in the oak species and in pine. In beech, fruiting levels one year before the MY were not important and in spruce, they were inconsistent depending on the region. As a consequence, evidence of resource depletion could only be seen in some regions for spruce.
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Nenad Potocic Ivan Seletkovic Mladen Ognjenovic Tamara Jakovljevic Melita Percec Tadic Volkmar TimmermannAbstract
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Dieback of European ash, caused by the ascomycete Hymenoscyphus fraxineus originating from Asia, has rapidly spread across Europe, and is threatening this keystone tree at a continental scale. High propagule pressure is characteristic to invasive species. Consistently, the enormous production of windborne ascospores by H. fraxineus in an ash forest with epidemic level of disease obviously facilitates its invasiveness and long distance spread. To understand the rate of build-up of propagule pressure by this pathogen following its local introduction, during 2011–2017 we monitored its sporulation at a newly infested ash stand in south-western Norway characterized with mild winters and cool summers. We also monitored the propagule pressure by Hymenoscyphus albidus, a non-pathogenic native species that competes for the same sporulation niche with H. fraxineus. During the monitoring period, crown condition of ash trees had impaired, and 20% of the dominant trees were severely damaged in 2017. H. fraxineus showed an exponential increase in spore production between 2012 and 2015, followed by drastic decline in 2016 and 2017. During 2011–2013, the two Hymenoscyphus species showed similar sporulation level, but thereafter spores of H. albidus were no longer detected. The data suggest that following local introduction, the population of H. fraxineus reaches rapidly an exponential growth stage if the local weather conditions are favorable for ascomata maturation across years. In the North Atlantic climate, summer temperatures critically influence the pathogen infection pressure, warm summers allowing the population to grow according to its biotic potential, whereas cold summers can cause a drastic decline in propagule pressure.
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Arne Verstraeten Elena Gottardini Nicolas Bruffaerts Bruno de Vos Elena Vanguelova Fabiana Cristofolini Sue Benham Pasi Rautio Liisa Ukonmaanaho Päivi Merilä Peter Waldner Marijke Hendrickx Gerrit Genouw Peter Roskams Nathalie Cools J Neirynck Anita Nussbaumer Mathias Neumann Nicholas Clarke Volkmar Timmermann Karin Hansen Hans-Peter Diettrich Manuel Nicolas Maria Schmitt Anne Thimonier Katrin Meusburger Silvio Schueler Anna KowalskaAbstract
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Isabella Børja Volkmar Timmermann Ari Hietala Mari Mette Tollefsrud Nina Elisabeth Nagy Adam Vivian-Smith Hugh Cross Jørn Henrik Sønstebø Tor Myking Halvor SolheimAbstract
In Norway the common ash (Fraxinus excelsior L.) has its northernmost distribution in Europe. It grows along the coastal range as small fragmented populations. The first occurrence of ash dieback caused by Hymenoscyphus fraxineus in Norway was reported in 2008. At that time, the disease had already spread through large areas of southern and south-eastern parts of Norway. Since then the disease continued spreading with a speed of about 50- 60 km per year along the western coastal range. To monitor the disease development over time, we established eight permanent monitoring plots in south-eastern and western Norway in 2009 and 2012, respectively. In all plots tree mortality was high, especially among the youngest trees in south-eastern Norway. The extent of crown damage has continually increased in all diameter classes for both regions. In 2009, 76.8 % of all trees on the five monitoring plots in south-eastern Norway were considered to be healthy or slightly damaged, and only 8.9 % to be severely damaged. In 2015, 51.7 % were dead, 13.5 % severely damaged and only 25.7 % remained healthy or slightly damaged. To assess the infection pressure and spore dispersal patterns of the pathogen, we used a Burkard volumetric spore sampler placed in an infested ash stand in southern Norway. We examined the airborne ascospores of H. fraxineus and H. albidus captured on the sampling tape microscopically and with real-time PCR assays specific to these fungi. We detected very few ascospores of H. albidus, whereas ascospores of H. fraxineus dominated throughout entire sampling periods of 2009, 2010 and 2011. Spore discharge occurred mainly between the hours of 5 and 8 a.m., though the distinctive sporulation had yearly variation between 5-7 a.m. We observed the same diurnal pattern throughout the entire sampling period, with a seasonal peak in spore liberation between mid-July and midAugust, after which the number of ascospores decreased substantially. Similar diurnal patterns were observed throughout the sampling period except that after mid-August the number of trapped ascospores substantially decreased. To compare the genetic pattern of common ash in the northern and central ranges of Europe we analyzed the Norwegian samples together with available samples from central Europe by using chloroplast and nuclear microsatellite markers. We found that the northern range of common ash was colonized via a single migration route that originated in eastern or south-eastern Europe with little influence originating from other southern or western European refugia. In the northern range margins, genetic diversity decreased and population differentiation increased, coherent with a post-glacial colonization history characterized by founder events and population fluctuations. Based on our findings we discuss the future management and conservational implications.
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Ash dieback, caused by the ascomycete Hymenoscyphus fraxineus, has been spreading throughout Europe since the early 1990s, threatening European ash at a continental scale. Little is known about the development of the disease in individual forest trees and in different age classes. In this study we monitored ash dieback on trees of different diameter classes in five permanent plots in ash stands in south-eastern Norway from 2009 to 2016, and from 2012 to 2016 in three plots in western Norway with a shorter disease history. Our results showed that more than 80% of the youngest and more than 40% of the intermediate future crop trees in the plots in south-eastern Norway were dead by 2016, while the disease development in large, dominant trees was slower. Although less damage has been observed in the plots in western Norway, the trend for the juvenile trees is the same as in south-eastern Norway with rapidly increasing damage and mortality. Most dead trees in south-eastern Norway were found at sites with high soil moisture and showed symptoms of root-rot caused by Armillaria species. Infected trees, both young and old ones, are weakened by the disease and appear to be more susceptible to other, secondary pathogens, especially under unfavourable site conditions.
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Ari Hietala Hugh Cross Halvor Solheim Volkmar Timmermann Nina Elisabeth Nagy Isabella Børja Adam Vivian-Smith Jørn Henrik SønstebøAbstract
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Hugh Cross Jørn Henrik Sønstebø Nina Elisabeth Nagy Volkmar Timmermann Halvor Solheim Isabella Børja Håvard Kauserud Tor Carlsen Barbara Rzepka Katarzyna Wasak Adam Vivian-Smith Ari HietalaAbstract
High biodiversity is regarded as a barrier against biological invasions. We hypothesized that the invasion success of the pathogenic ascomycete Hymenoscyphus fraxineus threatening common ash in Europe relates to differences in dispersal and colonization success between the invader and the diverse native competitors. Ash leaf mycobiome was monitored by high-throughput sequencing of the fungal internal transcribed spacer region (ITS) and quantitative PCR profiling of H. fraxineus DNA. Initiation of ascospore production by H. fraxineus after overwintering was followed by pathogen accumulation in asymptomatic leaves. The induction of necrotic leaf lesions coincided with escalation of H. fraxineus DNA levels and changes in proportion of biotrophs, followed by an increase of ubiquitous endophytes with pathogenic potential. H. fraxineus uses high propagule pressure to establish in leaves as quiescent thalli that switch to pathogenic mode once these thalli reach a certain threshold – the massive feedback from the saprophytic phase enables this fungus to challenge host defenses and the resident competitors in mid-season when their density in host tissues is still low. Despite the general correspondence between the ITS-1 and ITS-2 datasets, marker biases were observed, which suggests that multiple barcodes provide better overall representation of mycobiomes.
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Anita Nussbaumer Peter Waldner Sophia Etzold Arthur Gessler Sue Benham Iben Margrete Thomsen Bruno Bilde Jørgensen Volkmar Timmermann Arne Verstraeten Geert Sioen Pasi Rautio Liisa Ukonmaanaho Mitja Skudnik Vladislav Apuhtin Sabine Braun Alexandra WauerAbstract
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Johannes Eichhorn Peter Roskams Nenad Potocic Volkmar Timmermann Marco Ferretti Volker Mues Andras Szepesi Dave Durrant Ivan Seletkovic Hans‐Werner Schroeck Seppo Nevalainen Filippo Bussotti Paloma Garcia Sören WulffAbstract
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Ari Hietala Isabella Børja Hugh Cross Volkmar Timmermann Nina Elisabeth Nagy Halvor Solheim Jørn Henrik Sønstebø adam vivian smithAbstract
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Ari Hietala Isabella Børja Hugh Cross Nina Elisabeth Nagy Halvor Solheim Jørn Henrik Sønstebø Volkmar Timmermann Adam Vivian-SmithAbstract
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Marco Ferretti Karin Hansen Vicent Calatayud Marta Camino-Serrano Nathalie Cools Bruno de Vos Tiina M. Nieminen Nenad Potocic Pasi Rautio Marcus Schaub Volkmar Timmermann Liisa Ukonmaanaho Peter WaldnerAbstract
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Ari Hietala Hugh Cross Isabella Børja Nina Elisabeth Nagy Volkmar Timmermann Halvor Solheim Jørn Henrik Sønstebø Adam Vivian-SmithAbstract
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Marco Ferretti Tanja GM Sanders Alexa Michel Vicent Calatayud Nathalie Cools Elena Gottardini Matthias Haeni Karin Hansen Nenad Potocic Marcus Schaub Volkmar Timmermann Serina Trotzer Elena VanguelovaAbstract
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Walter Seidling Serina Trotzer Tanja GM Sanders Volkmar Timmermann Nenad Potocic Alexa MichelAbstract
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Ari M. Hietala, Volkmar Timmermann, Isabella Børja & Halvor Solheim Norwegian Forest and Landscape Institute. PO Box 115, 1431 Ås, Norway: ari.hietala@skogoglandskap.no Owing to the Gulf Stream, the northernmost European populations of several tree species are found in Norway. Common ash (Fraxinus excelsior), the only native ash species in Norway, is present in the lowlands in the southeastern part with continental climate and in southern and southwestern coastal regions with North Atlantic climate up to Central Norway. The current standing volume of ash in Norway is ca 3 mill m3 (broadleaved trees in total 220 mill m3). The first documentation of Ash Dieback (ADB) is from 2008 from a nursery in the southeastern part of the country. A survey later that year showed that dieback symptoms were present over a distance of nearly 400 km in the southeastern region. In addition to nurseries and forests, ADB symptoms were observed on roadside, alley, garden and park trees. Based on the presence of old ADB-like stem lesions detected in 2008, the pathogen must have arrived to Norway no later than 2006. In 2008, the Norwegian Food Safety Authority laid down regulations with the aim of preventing further spread of ADB. These regulations divide the country into quarantine, observation and infection-free zones, and prohibit the export of ash seedlings, seed and wood from the quarantine zone. Despite of these regulations, the disease spread rapidly along the western coast in the period between 2009 and 2013, and currently only the ash stands in Central Norway are free of the disease. The rapid spread of the disease in Norway is obviously due to airborne dispersal of pathogen ascospores. In our experimental stand in SE Norway the number of pathogen fruit bodies can be as high as 10,000 per m2 in the peak season, mid-July to mid-August. During the early morning hours the amount of pathogen ascospores at a diseased stand can exceed 100,000 ascospores per m3 air. The first symptoms of the disease, necrotic lesions on leaf blade and petiole, appear typically during the first two weeks of August in SE Norway. To observe long-term impacts of ADB, eight monitoring plots have been established in continental and North Atlantic climate zones. In SE Norway with the oldest disease history, above 60 % of the trees with a breast height diameter (BHD) below 12.5 cm have so far died or suffer from severe defoliation, 1/3 of the larger trees being affected to a similar degree. The proportions of healthy (no signs of defoliation) small and larger trees are 20% and 37%, respectively. In SW Norway with more recent disease history a similar trend is observed but the proportion of dead trees is still small. As a consequence of ADB, the Norwegian nurseries no longer grow ash seedlings. There are currently no practical control options for the disease in forestland. Several European countries have reported that even at heavily diseased ash stands there are often some ash trees that show little symptoms. This may be due to genetic variation between trees in disease resistance, a hypothesis that is currently being investigated in several European projects. Thus implementation of forest management practices that eliminate ash could have a negative effect as survival of the tree ultimately depends on selection of trees with increased disease resistance. Bibliography for Ari M. Hietala Ari M. Hietala is a Senior Forest Pathologist at the Norwegian Forest and Landscape Institute, which is a primarily government funded organisation providing scientific research and services to government, non-governmental and commercial organisations. He has worked with a range of fungal root and shoot diseases occurring on broadleaved trees and conifers indigenous to the Nordic countries. Ari and the rest of the group participate currently in several European consortia engaged in ash dieback research.
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Matthieu Jonard Alfred Fürst Arne Verstraeten Anne Thimonier Volkmar Timmermann Nenad Potocic Peter Waldner Sue Benham Karin Hansen Päivi Merilä Quentin Ponette Ana C. de la Cruz Peter Roskams Manuel Nikolas Luc Croisé Morten Ingerslev Giorgio Matteucci Bruno Decinti Marco Bascietto Pasi RautioAbstract
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Shoot dieback disease of European ash caused by the ascomycete Hymenoscyphus pseudoalbidus threatens ash on a continental scale. A spore sampler placed in a diseased ash forest in Southern Norway, coupled with microscopy and DNA-based fungal species-specific real-time PCR assays, was employed to profile diurnal and within-season variation in infection pressure by ascospores of H. pseudoalbidus and the potentially co-existing non-pathogenic Hymenoscyphusalbidus. Hymenoscyphus pseudoalbidus was found to be predominant in the stand. Massive simultaneous liberation, by active discharge of pathogen ascospores in the morning, peaked in mid-Jul. to mid-Aug. Accumulation of pathogen DNA on leaflets of current-year leaves reached a high level plateau phase before appearance of autumn coloration, suggesting that pathogen establishment in leaves is terminated before the onset of leaf senescence.
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Winter damage caused by frost is frequently observed on common ash (Fraxinus excelsior) in Norway. In spring 2007, extensive winter damage most likely camouflaged ash dieback caused by Chalara fraxinea. In 2008, ash dieback caused by C. fraxinea had spread to large areas in the southern part of Norway. The disease was widespread in forests and nurseries, but also on roadside trees, and in gardens and parks. In 2009, the disease had spread to new areas; about 30 km into Rogaland county in southwestern Norway and also further into some valleys in southeastern Norway.
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Dieback of European ash (Fraxinus excelsior), caused by the ascomycete Hymenoscyphus pseudoalbidus (anamorph Chalara fraxinea), started around 1992 in Poland and has since then spread over large geographical areas. By November 2010, the disease had been recorded in 22 European countries. The gradual expansion and high intensity of the ash dieback epidemic in Europe may suggest that H. pseudoalbidus is an invasive alien organism. In Norway, ash dieback was first reported in spring 2008, and a survey in early summer of the same year revealed that the disease had spread over large parts of the southern and eastern regions of the country. The distance from the southernmost to the northernmost infected stands was, at that time, about 400 km. Some old necrotic lesions were also observed, indicating that the ash dieback pathogen is likely to have been present in Norway since at least 2006. In 2009, a spore sampler was installed in a diseased ash stand at Ås, South-Eastern Norway. Sampling started in late July and continued until late September. Large numbers of ascospores resembling those of H. pseudoalbidus were observed, with the maximum number of spores occurring from the end of July to mid-August. The deposition of ascospores occurred mainly between 6 and 8 a.m. Ascospores are most likely to be the primary source initiating host infections and responsible for the rapid recent spread of H. pseudoalbidus in Europe.
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Holger Lange Nicholas Clarke O. Janne Kjønaas Wenche Aas Kjell Andreassen Isabella Børja Harald Bratli Susanne Eich-Greatorex Toril Drabløs Eldhuset Kjersti Holt Hanssen Tonje Ingeborg Økland Ingvald Røsberg Trine Aulstad Sogn Tomasgaard Volkmar TimmermannAbstract
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Holger Lange Nicholas Clarke O. Janne Kjønaas Wenche Aas Kjell Andreassen Isabella Børja Harald Bratli Susanne Eich-Greatorex Toril Drabløs Eldhuset Kjersti Holt Hansen Tonje Ingeborg Økland Ingvald Røsberg Trine Aulstad Sogn Volkmar TimmermannAbstract
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Nicholas Clarke O. Janne Kjønaas Wenche Aas Kjell Andreassen Isabella Børja Harald Bratli Susanne Eich-Greatorex Toril Drabløs Eldhuset Kjersti Holt Hanssen Holger Lange Tonje Økland Ingvald Røsberg Trine Aulstad Sogn Volkmar TimmermannAbstract
In Norway, it is planned to double the stationary use of bioenergy from all sources by up to 14 TWh before 2020, with much of this increase coming from forest resources, including residues like branches and tops (which are not much used today) being removed after tree harvest. This removal will reduce the supply of nutrients and organic matter to the forest soil, and may in the longer term increase the risk for future nutrient imbalance, reduced forest production, and changes in biodiversity and ground vegetation species composition. However, field experiments have found contrasting results (e.g. Johnson and Curtis 2001; Olsson et al. 1996). Soil effects of increased biomass removal will be closely related to soil organic matter (SOM) dynamics, litter quality, and turnover rates. The SOM pool is derived from a balance between above- and below-ground input of plant material and decomposition of both plants and SOM. Harvest intensity may affect the decomposition of existing SOM as well as the build-up of new SOM from litter and forest residues, by changing factors like soil temperature and moisture as well as amount and type of litter input. Changes in input of litter with different nutrient concentrations and decomposition patterns along with changes in SOM decomposition will affect the total storage of carbon, nitrogen and other vital nutrients in the soil. To quantify how different harvesting regimes lead to different C addition to soil, and to determine which factors have the greatest effect on decomposition of SOM under different environmental conditions, two Norway spruce forest systems will be investigated in the context of a research project starting in 2008/2009, one in eastern and one in western Norway, representing different climatic and landscape types. At each location, two treatment regimes will be tested: Conventional harvesting, with residues left on-site (CH) Aboveground whole-tree harvest, with branches, needles, and tops removed (WTH). Input of different forest residues will be quantified post harvest. Soil water at 30 cm soil depth will be analysed for nutrients and element fluxes will be estimated to provide information about nutrient leaching. Soil respiration will be measured, along with lab decomposition studies under different temperature and moisture regimes. Long term in situ decomposition studies will be carried out in the WTH plots using three different tree compartments (needles, coarse twigs, fine roots) decomposing in litter bags, in order to determine their limit value. The structure of the fungal community will be determined by soil core sampling and use of molecular techniques allowing qualitative and quantitative estimation. Understorey vegetation will be sampled to determine the biomass, and the frequency of all vascular plants, bryophytes and lichens will be estimated. After harvesting, replanting will be carried out. Seedling survival, causes of mortality and potential damage, growth, and needle nutrients will be monitored. Results from these studies will be used to identify key processes explaining trends observed in two series of ongoing long-term whole-tree thinning trials. We shall combine knowledge obtained using field experiments with results of modelling and data from the Norwegian Monitoring Programme for Forest Damage and National Forest Inventory. This will help us to predict and map the ecologically most suitable areas for increased harvesting of branches and tops on a regional scale based on current knowledge, and to identify uncertainties and additional knowledge needed to improve current predictions.
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Growth of Norway spruce (Picea abies) trees and nitrogen deposition were analysed at about 500 forest plots throughout Norway in six fiveyear periods from 1977 to 2006. Growth was calculated from five repeated calliper measurements of all trees during this period and using treering series from increment cores of a subsample of trees. From the growth data a `relative growth` variable was extracted, being the deviation in % between observed and expected growth rates. The expected growth was estimated from growth models based on site productivity, age and stand density at each plot. The plots were categorized into four age classes. The nitrogen deposition was estimated for each plot for the same five year periods by geographical interpolation of deposition observations at monitoring stations made by the Norwegian Institute for Air Research. Nitrogen deposition from 1977 to 2006 ranged from 1 to 24 kg/ha/yr at the study plots, with about 15 kg/ha/yr in the southernmost region and 3 kg/ha/yr in the northern region of Norway. For the entire 30year period we found a long term relationship between growth and nitrogen deposition, corresponding to a forest growth increase of 0.7% per kg total nitrogen deposition per hectare and year (r2 = 0.13). This is in line with studies carried out on other data sets and for shorter time periods. This apparent fertilizing effect was most pronounced for the youngest forest, while the effect was weak for the oldest forest. The growth increase was observed in the southernmost part of Norway, the region with the highest nitrogen deposition. However, the relationship between nitrogen deposition and growth varied considerably between the time periods. In two of the periods the relationship was slightly negative: these periods corresponded well with summer droughts occurring in the southernmost part of Norway. Drought, as well as other climatic factors, will influence the shortterm variations in forest growth and may obscure the fertilizing effect of nitrogen deposition in some periods. In conclusion, nitrogen deposition has most likely increased growth in Norway spruce in southern Norway. However, our study also shows that inferences from such correlative studies should be drawn with care if the growth period is shorter than 10–15 years because climatic factors produce temporal variations in the relationship between nitrogen deposition and forest growth.
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The Norwegian Monitoring Programme for Forest Damage (OPS) has since its start registered damage to selected trees. The aim of the registrations has been to explain variations in crown density and crown colour. In answer to international requests, the Norwegian Forest and Landscape Institute has prepared a short guide to the determination of the most common forms of damage found in Norwegian forests...
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Volkmar TimmermannAbstract
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Volkmar TimmermannAbstract
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Volkmar TimmermannAbstract
Monitoring on the forest officers\" plots in Norway has been running since 1988, with annual assessments carried out by local forest officers. In 2005 they assessed 30277 trees on 557 plots. For 15090 of these trees on 348 plots, there exist complete records of crown condition over the past 18 years. The plots are subjectively selected, mainly in Norway spruce dominated stands, and divided into four development classes. The results from 2005 show a slight increase in mean defoliation of Norway spruce to 16.4 %. There were small, but mostly negative changes in most regions without clear trends in the long term series. In southern Norway, a strong increase in defoliation was observed. Crown colour was improving in the western and northern regions of the country, whereas discolouration was increasing in south-eastern Norway. The mean discolouration was still very low in 2005, with 90 % of the spruce trees having normal, green colour. The mortality rate was low, on the average 3.4 ‰ for all trees. Only few causal agents of crown damage were reported. Data from 18 years of monitoring on the forest officers’ plots reveal some regional patterns for defoliation of spruce in Norway, with western Norway having the lowest mean defoliation through all years, and Mid-Norway the highest. In the other regions, the trends are not so clear with greater fluctuations in defoliation and discolouration.
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Changes in the ectomycorrhizal fungus flora were studied in connection with nitrogen addition and removal experiments in a Norway spruce forest at Gårdsjön, W Sweden during a 5-year period. The above-ground ectomycorrhizal fruit body production was recorded from permanent transect plots, and the below-ground mycorrhizal fine-roots density and morphotype differentiation were studied from soil core samples from the surface root layer. The experiments were performed by adding N-enriched and N-free water, respectively, by means of sprinkling systems. Ammonium nitrate (about 35 kg N ha−1 yr−1) was added to catchment G2 NITREX, whereas at adjacent catchment G1 ROOF ambient N deposition was removed by means of a roof. The addition of N led to a rapid and substantial decrease in species diversity and fruit body production of most species in the NITREX catchment, representing one of very few biological responses to the treatments at Gårdsjön. Stress-intolerant groups such as the initially-dominant genus Cortinarius were almost absent after 5 yr of N addition. Only one dominant species (Cantharellus tubaeformis) increased fruit body production after treatment. In the nitrogen removal (G1 ROOF) experiment, the fruit body production increased strongly the first years, but then declined. No response in the below-ground mycorrhiza and fine-root density and diversity was found. All fine roots had developed ectomycorrhiza. The difference in response above ground and below ground indicates that: (1) the fruit-body producing macrofungi play a minor role below ground, and that (2) there is probably a considerable time-lag in the mycorrhizal fine-root versus fruit body production response to enhanced N levels.
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Andries W. Boxman Kai Blanck Tor Erik Brandrud B. A. Emmett Per Gundersen Rene F. Hogervorst O. Janne Kjønaas Hans Persson Volkmar TimmermannAbstract
Enhancement of the atmospheric N deposition is a serious threat for the structure and function of ecosystems. Here we evaluate the ecological effects of excess N with respect to changes in vegetation and soil biota in a series of experiments along a N gradient across Europe. The aim of this project (NITREX: N saturation EXperiments) is to assess the risk of N saturation and the reversibility of N saturation. At the experimental sites with a low-to-moderate input, N was added (n = 3), while at sites with a high input, N was removed by means of a transparent roof (n = 4). The experiments started between 1989 and 1991. Across the N gradient a positive correlation was found between the N concentration in deposition or soil solution with the N concentration in the needles and in general a negative correlation with the base cations K and Mg. In the N-addition plots there was a tendency towards a decreasing nutrient status of the needles, whereas at one site N-removal led to an improvement. Addition of N hardly affected fine-root biomass production, whereas signs of growth increase were recorded when the input was reduced. Tree growth was accelerated upon input reduction at two of three sites. Manipulation of N input did not alter the decomposition rate, although significant differences between sites were noted. Manipulation of the N input hardly affected the biomass of fungi and bacteria, but a negative relation between the N-addition and part of the soil fauna may be present among sites.
