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Hymenoscyphus fraxineus is an introduced ascomycete fungus which causes ash dieback and has resulted in widespread mortality of ash throughout Europe. Although H. fraxineus has been present on the continent for at least four decades, it was not identified until 2006. The first record of the pathogen in Britain came in 2012 although it was probably present a decade earlier. The most common host European ash (Fraxinus excelsior L.) is economically and ecologically important to Britain where the cost of ash dieback is estimated at billions of pounds. The impact of ash dieback has stimulated a major research response which we review with the aim of providing up-to-date information relevant to Britain and identifying knowledge gaps where research would contribute to improved disease mitigation. Hymenoscyphus fraxineus is an outcrossing fungus with high genotypic diversity; ascospores produced via sexual reproduction are critical to aerial dispersal and infection. Temperature, moisture, and ground cover influence pathogen fruit body development, the timing of ascospore release, and extent of ascospore germination; they also interact together to affect the likelihood of infection. In addition, stand characteristics, including tree density, tree height, and landscape fragmentation, affect disease dynamics with increased disease severity on moist sites with high ash density. Efforts at finding natural resistance in ash have identified genetic markers associated with disease tolerance, and gene expression analysis is providing insights into the basis of that resistance. Mainland European findings indicate that ash dieback makes trees more vulnerable to other pathogens, whilst endophytes in the ash phyllosphere can suppress infection by H. fraxineus. Possible tools for long-term control of ash dieback include (1) deployment of resistant ash, (2) quantitatively informed management practices based on microclimate models and better understanding of the pathogen life cycle, and (3) manipulation of biocontrol agents from the ash microbiome or pathogen mycoviruses. [ABSTRACT FROM AUTHOR]

The increase in introduced insect pests and pathogens due to anthropogenic environmental changes has become a major concern for tree species worldwide. Common ash (Fraxinus excelsior L.) is one of such species facing a significant threat from the invasive fungal pathogen Hymenoscyphus fraxineus. Some studies have indicated that the susceptibility of ash to the pathogen is genetically determined, providing some hope for accelerated breeding programs that are aimed at increasing the resistance of ash populations. To address this challenge, we used a genomic selection strategy to identify potential genetic markers that are associated with resistance to the pathogen causing ash dieback. Through genome‐wide association studies (GWAS) of 300 common ash individuals from 30 populations across Poland (ddRAD, dataset A), we identified six significant SNP loci with a p‐value ≤1 × 10−4 associated with health status. To further evaluate the effectiveness of GWAS markers in predicting health status, we considered two genomic prediction scenarios. Firstly, we conducted cross‐validation on dataset A. Secondly, we trained markers on dataset A and tested them on dataset B, which involved whole‐genome sequencing of 20 individuals from two populations. Genomic prediction analysis revealed that the top SNPs identified via GWAS exhibited notably higher prediction accuracies compared to randomly selected SNPs, particularly with a larger number of SNPs. Cross‐validation analyses using dataset A showcased high genomic prediction accuracy, predicting tree health status with over 90% accuracy across the top SNP sets ranging from 500 to 10,000 SNPs from the GWAS datasets. However, no significant results emerged for health status when the model trained on dataset A was tested on dataset B. Our findings illuminate potential genetic markers associated with resistance to ash dieback, offering support for future breeding programs in Poland aimed at combating ash dieback and bolstering conservation efforts for this invaluable tree species. [ABSTRACT FROM AUTHOR]

Abstract The increase in introduced insect pests and pathogens due to anthropogenic environmental changes has become a major concern for tree species worldwide. Common ash (Fraxinus excelsior L.) is one of such species facing a significant threat from the invasive fungal pathogen Hymenoscyphus fraxineus. Some studies have indicated that the susceptibility of ash to the pathogen is genetically determined, providing some hope for accelerated breeding programs that are aimed at increasing the resistance of ash populations. To address this challenge, we used a genomic selection strategy to identify potential genetic markers that are associated with resistance to the pathogen causing ash dieback. Through genome‐wide association studies (GWAS) of 300 common ash individuals from 30 populations across Poland (ddRAD, dataset A), we identified six significant SNP loci with a p‐value ≤1 × 10−4 associated with health status. To further evaluate the effectiveness of GWAS markers in predicting health status, we considered two genomic prediction scenarios. Firstly, we conducted cross‐validation on dataset A. Secondly, we trained markers on dataset A and tested them on dataset B, which involved whole‐genome sequencing of 20 individuals from two populations. Genomic prediction analysis revealed that the top SNPs identified via GWAS exhibited notably higher prediction accuracies compared to randomly selected SNPs, particularly with a larger number of SNPs. Cross‐validation analyses using dataset A showcased high genomic prediction accuracy, predicting tree health status with over 90% accuracy across the top SNP sets ranging from 500 to 10,000 SNPs from the GWAS datasets. However, no significant results emerged for health status when the model trained on dataset A was tested on dataset B. Our findings illuminate potential genetic markers associated with resistance to ash dieback, offering support for future breeding programs in Poland aimed at combating ash dieback and bolstering conservation efforts for this invaluable tree species.

In light of the increase of environmental pollution, we tested the effect of cadmium (Cd) and lead (Pb) soil contamination on ash dieback. The experiment included the inoculation of Hymenoscyphus fraxineus on Fraxinus excelsior and Fraxinus angustifolia seedlings growing on unpolluted soil, soil contaminated with cadmium (Cd), and soil contaminated with lead (Pb). At the end of the experiment, 173 days after soil contamination and 50 days since inoculation, all F. excelsior and F. angustifolia seedlings inoculated with H. fraxineus showed ash dieback symptoms in comparison to their control groups. However, both F. excelsior and F. angustifolia seedlings grown on contaminated soil had significantly increased necrotic lesions in comparison to the seedlings grown on uncontaminated soil. Our results showed for the first time that cadmium (Cd) and lead (Pb) soil contamination can significantly contribute to ash dieback and increase damage to F. excelsior and F. angustifolia seedlings. [ABSTRACT FROM AUTHOR]

European ash dieback caused by the alien, invasive ascomycete species Hymenoscyphus fraxineus currently represents, along with its side effects, the greatest threat to common ash (Fraxinus excelsior) trees in Europe. The disease is widely distributed in Germany and present in all regions where common ash is growing. In order to study European ash dieback in Germany and to develop management strategies to conserve common ash as a forest tree species, the concerted, interdisciplinary research project FraxForFuture was initiated. FraxForFuture consist of an association of five sub-networks: FraxConnect, FraxMon, FraxGen, FraxPath, and FraxSilva. In total, 27 individual projects conduct research on the epidemiology of the disease, the pathogen and the preservation of ash, including various control, breeding, and silvicultural strategies. The main goals of the FraxForFuture network and the sub-networks as well as essential information about the network of common research plots are provided. [ABSTRACT FROM AUTHOR]

Ash dieback caused by the invasive fungus Hymenoscyphus fraxineus leads to massive mortality among common ash (Fraxinus excelsior) in Europe. To select tolerant genotypes, inoculation experiments are frequently conducted with isolates of the fungus. The aim of this study was to improve the inoculation methodology for evaluating susceptibility of ash genotypes to ash dieback through (i) testing the virulence of different isolates of Hymenoscyphus fraxineus and (ii) investigating temporal lesion development using three widely applied cultivars: Altena, Atlas and Westhof's Glorie. First, an experiment was conducted with a repeated measure design in which virulence of five different isolates was tested. Second, a subset of two isolates that induced the longest downward lesion length in experiment one was used in combination with a slightly adjusted inoculum protocol. Significant differences were found between isolates for downward lesion length, but a significant interaction effect of isolates and cultivars was absent. Also, the inoculation position within the stem affected lesion length; the largest lesions were found on the highest inoculation position within the stem. Furthermore, we found that cryopreserved isolates can remain virulent over years. The timing of inoculation at the end of the growing season was effective as large lesions already occurred during winter dormancy. For future inoculation studies, we propose to use: (i) isolates that induce large lesions, as less virulent isolates induced not only shorter but also fewer lesions; (ii) a similar inoculation position to better compare inoculations within and between experiments; (iii) cryopreserved isolates for testing over years; (iv) mycelial suspensions for inoculum preparation to cover wood chips more evenly; (v) reference clone—like the studied cultivars—to standardize research outcomes between years and research groups. [ABSTRACT FROM AUTHOR]

European ash (Fraxinus excelsior) is highly affected by the pathogenic fungus Hymenoscyphus fraxineus in all of Europe. Increases in plant’s secondary metabolite (SM) production is often linked tol enhanced resistance to stress, both biotic and abiotic. Moreover, plant-associated bacteria have been shown to enhance SM production in inoculated plants. Thus, our hypothesis is that bacteria may boost ash SM production, hence priming the tree’s metabolism and facilitating higher levels of resilience to H. fraxineus. We tested three different ash genotypes and used Paenibacillus sp. and Pseudomonas sp. for inoculation in vitro. Total phenol (TPC), total flavonoid (TFC) and carotenoid contents were measured, as well as the chlorophyll a/b ratio and morphometric growth parameters, in a two-stage trial, whereby seedlings were inoculated with the bacteria during the first stage and with H. fraxineus during the second stage. While the tested bacteria did not positively affect the morphometric growth parameters of ash seedlings, they had a statistically significant effect on TPC, TFC, the chlorophyll a/b ratio and carotenoid content in both stages, thus confirming our hypothesis. Specifically, in ash genotype 64, both bacteria elicited an increase in carotenoid content, TPC and TFC during both stages. Additionally, Pseudomonas sp. inoculated seedlings demonstrated an increase in phenolics after infection with the fungus in both genotypes 64 and 87. Our results indicate that next to genetic selection of the most resilient planting material for ash reforestation, plant-associated bacteria could also be used to boost ash SM production. [ABSTRACT FROM AUTHOR]

Development of effective biocontrol procedures using ash endophytes to combat an ash pathogen Hymenoscyphus fraxineus would be an appropriate contribution to the ongoing effort to protect European ash stands against ash decline. In this study we investigated the biocontrol potential of two ash endophytes, Thielavia basicola and Minimidochium sp., against H. fraxineus using in vitro plant-fungus and fungus-fungus dual cultures approach in three biocontrol models. The tests aimed to determine whether the endophytes show antagonism toward Fraxinus excelsior and F. pennsylvanica, to assess the level of antagonism of the endophytes toward H. fraxineus and to identify potential secondary metabolites induced by the presence of H. fraxineus. The results that dual culture experiments modeled according to our design may be a very useful tool to precisely study biocontrol potential of fungi, i.e., without the impact of environmental factors. Such experiments also enable the selection of most resistant ash genotypes and rapid propagation, producing large numbers of pathogen-free seedlings. It should be noted, however, that both of the endophytes tested in the dual cultures strongly inhibited the growth of H. fraxineus. Their growth under the influence of callus/seedlings was also inhibited. Comparison of HPLC profiles showed that the presence of H. fraxineus in the post-culture medium induced the production of an unknown secondary metabolite in this species. Such results suggest that some of the plant–fungus combinations examined in this study may have potential to be developed as biocontrol methods, thus increasing the survivability of ash stands under natural conditions.

European ash (Fraxinus excelsior) is highly affected by the pathogenic fungus Hymenoscyphus fraxineus in all of Europe. Increases in plant’s secondary metabolite (SM) production is often linked tol enhanced resistance to stress, both biotic and abiotic. Moreover, plant-associated bacteria have been shown to enhance SM production in inoculated plants. Thus, our hypothesis is that bacteria may boost ash SM production, hence priming the tree’s metabolism and facilitating higher levels of resilience to H. fraxineus. We tested three different ash genotypes and used Paenibacillus sp. and Pseudomonas sp. for inoculation in vitro. Total phenol (TPC), total flavonoid (TFC) and carotenoid contents were measured, as well as the chlorophyll a/b ratio and morphometric growth parameters, in a two-stage trial, whereby seedlings were inoculated with the bacteria during the first stage and with H. fraxineus during the second stage. While the tested bacteria did not positively affect the morphometric growth parameters of ash seedlings, they had a statistically significant effect on TPC, TFC, the chlorophyll a/b ratio and carotenoid content in both stages, thus confirming our hypothesis. Specifically, in ash genotype 64, both bacteria elicited an increase in carotenoid content, TPC and TFC during both stages. Additionally, Pseudomonas sp. inoculated seedlings demonstrated an increase in phenolics after infection with the fungus in both genotypes 64 and 87. Our results indicate that next to genetic selection of the most resilient planting material for ash reforestation, plant-associated bacteria could also be used to boost ash SM production.

Tree pathogens are a major threat to forest ecosystems. Conservation management strategies can exploit natural mechanisms of resistance, such as tree genotype and host‐associated microbial communities. However, fungal and bacterial communities are rarely looked at in the same framework, particularly in conjunction with host genotype. Here, we explore these relationships and their influence on ash dieback disease, caused by the pathogen Hymenoscyphus fraxineus, in European common ash trees.We collected leaves from UK ash trees and used microsatellite markers to genotype trees, quantitative PCR to quantify H. fraxineus infection load and ITS and 16S rRNA amplicon sequencing to identify fungal and bacterial communities, respectively.There was a significant association between H. fraxineus infection intensity and ash leaf fungal and bacterial community composition. Higher infection levels were positively correlated with fungal community alpha‐diversity, and a number of fungal and bacterial genera were significantly associated with infection presence and intensity. Under higher infection loads, leaf microbial networks were characterized by stronger associations between fewer members than those associated with lower infection levels. Together these results suggest that H. fraxineus disrupts stable endophyte communities after a particular infection threshold is reached, and may enable proliferation of opportunistic microbes. We identified three microbial genera associated with an absence of infection, potentially indicating an antagonistic relationship with H. fraxineus that could be utilized in the development of anti‐pathogen treatments.Host genotype did not directly affect infection, but did significantly affect leaf fungal community composition. Thus, host genotype could have the potential to indirectly affect disease susceptibility through genotype × microbiome interactions, and should be considered when selectively breeding trees.Synthesis. We show that the diversity, composition and network structure of ash leaf microbial communities are associated with the severity of infection from ash dieback disease, with evidence of disease‐induced dysbiosis. We also show that host genotype influences leaf fungal community composition, but does not directly influence tree infection. These findings help to elucidate relationships between host genetics, the microbiome and a tree pathogen, highlighting potential resistance mechanisms and possible co‐infection concerns that could inform ash tree management. [ABSTRACT FROM AUTHOR]

Novel pests and diseases are becoming increasingly common, and often cause additional mortality to host species in the newly contacted communities. This can alter the structure of the community up to, and including, the extinction of host species. In the last 20 years, ash dieback (ADB) disease has spread into Europe from East Asia. It has caused substantial mortality in ash tree (Fraxinus excelsior L.) populations. However, a proportion of the individuals in most populations appear to be less susceptible to ADB and resistance seems to have high heritability. These observations have led to suggestions that ash populations may be sustainable after the disease. In order to test this hypothesis, I modified an existing model of UK woodland (parametrized for Wytham Woods, Oxfordshire) to take into account the impact of ADB and allowed offspring to inherit resistance traits from their parent. The results suggest that ash populations would still exist in 100 years, but at lower levels than they are currently. For example, when the initial proportion of resistant individuals is about 10% and heritability of resistance is 0.5, then the population of ash falls to about one-third of present levels. The proportion of individuals initially resistant to ADB had a larger effect on population size after 100 years than the heritability of resistance. The fact that the initial size of the resistant population is important to achieve a high population size in the presence of ADB suggests that a selective breeding programme with the intention of augmenting the natural ash populations would be beneficial.

Temperate European forests are currently largely under attack by the infection with Hymenoscyphus fraxineus, a fungal pathogen introduced from Asia since at least the early 1990s and causing a major dieback of common ash (Fraxinus excelsior) throughout Europe. At present, ash dieback evokes major problems for forestry, in particular in sensitive forest remnants in Northern Germany, where the disease causes serious concerns for ecosystem conservation. This makes ash dieback a focal area of ecological research. In the present study, we quantified the extent of ash dieback in adult and in young ash trees in Northern Schleswig-Holstein, Germany, in relation to community composition and associated biotic and abiotic factors. Data collection was carried out in 37 plots in communities of ash-rich forests and included floristic inventory, rating of adult and young ash individuals and recording of light and soil conditions. Data were analyzed using non-metric multidimensional scaling and general linear mixed effects models. Forest type was the strongest significant predictor for variation in crown defoliation of adult ash trees. Damage was highest in communities of wet alder-ash forests and lowest in ash-rich beech forests. A further significant predictor of adult crown defoliation was individual height of the ash tree with larger trees being less affected than smaller ones. For juveniles, total species richness displayed a significant positive relationship with the proportional abundance of fungal infection, while the mean damage proportion per individual significantly increased with increasing relative light intensity in the understorey. The study clearly shows a strong relationship between forest type and ecosystem vulnerability to ash dieback. In particular, communities belonging to the species-rich wet alder-ash forests were most severely affected by ash disease, thereby deserving special attention among the vulnerable fragmented forest remnants in Schleswig-Holstein. Co-varying factors, however, seem to differ between juvenile and adult trees, hinting at the relative importance of tree performance for the adult trees and abiotic conditions for the juveniles. Accounting for such differences along a larger ecological gradient of ash forest communities will be necessary to more comprehensively understand effects of ash dieback on the ecosystem and needs to be addressed in future research.

Ash dieback, caused by the ascomycete fungus Hymenoscyphus fraxineus, has been rapidly expanding across Europe during the last two decades, posing a considerable threat to native ash populations. In this study, we applied regression-based models trained by field data, in conjunction with geographic information systems, to produce spatial predictions of ash dieback severity and environmental suitability for the disease in Czech forests. A model of actual ash dieback severity relates disease extent to silvicultural and environmental characteristics of forest stands and their neighbourhood, while a model of environmental suitability for the disease quantifies the relative susceptibility of sites to the disease, independent of the current silvicultural characteristics. The final predictive maps suggested that fertile lowlands and humid areas bordering Poland and Slovakia were the most endangered regions. Areas at the lowest risk of damage were concentrated in dry areas and in highland and mountain areas in the western part of the country, usually with poor soils on acid bedrock. Predictions of actual disease severity are an effective tool for guiding the current management of infested stands whereas predicting environmental suitability is useful for making long-term strategic decisions, e.g. identifying areas where future ash regeneration and cultivation may be unsuccessful. [ABSTRACT FROM AUTHOR]

Temperate European forests are currently largely under attack by the infection with Hymenoscyphus fraxineus , a fungal pathogen introduced from Asia since at least the early 1990s and causing a major dieback of common ash (Fraxinus excelsior) throughout Europe. At present, ash dieback evokes major problems for forestry, in particular in sensitive forest remnants in Northern Germany, where the disease causes serious concerns for ecosystem conservation. This makes ash dieback a focal area of ecological research. In the present study, we quantified the extent of ash dieback in adult and in young ash trees in Northern Schleswig-Holstein, Germany, in relation to community composition and associated biotic and abiotic factors. Data collection was carried out in 37 plots in communities of ash-rich forests and included floristic inventory, rating of adult and young ash individuals and recording of light and soil conditions. Data were analyzed using non-metric multidimensional scaling and general linear mixed effects models. Forest type was the strongest significant predictor for variation in crown defoliation of adult ash trees. Damage was highest in communities of wet alder-ash forests and lowest in ash-rich beech forests. A further significant predictor of adult crown defoliation was individual height of the ash tree with larger trees being less affected than smaller ones. For juveniles, total species richness displayed a significant positive relationship with the proportional abundance of fungal infection, while the mean damage proportion per individual significantly increased with increasing relative light intensity in the understorey. The study clearly shows a strong relationship between forest type and ecosystem vulnerability to ash dieback. In particular, communities belonging to the species-rich wet alder-ash forests were most severely affected by ash disease, thereby deserving special attention among the vulnerable fragmented forest remnants in Schleswig-Holstein. Co-varying factors, however, seem to differ between juvenile and adult trees, hinting at the relative importance of tree performance for the adult trees and abiotic conditions for the juveniles. Accounting for such differences along a larger ecological gradient of ash forest communities will be necessary to more comprehensively understand effects of ash dieback on the ecosystem and needs to be addressed in future research. [ABSTRACT FROM AUTHOR]

There are relatively few studies surveying how meteorological and qualitative characteristics of tree sites in urban areas influence Hymenoscyphus fraxineus (T. Kowalski) Baral et al. (≡Chalara fraxinea T. Kowalski, =Hymenoscyphus pseudoalbidus Queloz et al.) or infection by that. Based on the significance of ash trees in urban areas and the frequent use of ash species and their varieties, it becomes increasingly important to identify factors that drive the pathogen. The aim of this study was to identify the effects of both meteorological parameters and tree characteristics on the severity of the disease in the city of Leipzig, Germany. We assessed 401 ash trees along streets and in green spaces distributed across Leipzig. The assessment included scoring the level of damage by H. fraxineus , visually estimating the degree of soil sealing under a tree's crown, measuring tree height, the height of crown base and the diameter at breast height, determining sex, year of planting and species variety, as well as recording qualitative characteristics (e.g. secondary pathogens, feeding marks). Mean relative air humidity, mean annual temperature and a combined index of meteorological conditions were used as meteorological parameters. In addition, we sampled tissue of 45 ash trees for the purpose of genetically verifying the occurrence of H. fraxineus. We analysed the effects of all variables on ash dieback by using linear mixed effect models. Results show an extensive damage by H. fraxineus in the whole city area with 70% of trees being infected. Mean relative air humidity and the index of meteorological conditions show a significant positive influence on the severity of ash dieback, whereas year of planting has a negative effect. Trees with feeding marks at the trunk tend to be more damaged by the fungus. Moreover, we found a significant difference in susceptibility between F. excelsior and its variety 'Atlas'. Both meteorological conditions and the susceptibility of species and their varieties should be taken into account when planting ash trees in urban areas. [ABSTRACT FROM AUTHOR]

Ash dieback caused by an alien, invasive fungus Hymenoscyphus fraxineus is a serious disease of European ash species in many parts in Europe. In Poland, the disease was recorded in the beginning of the 1990s. This study was performed in 2016–2017 with the aim to identify fungi isolated from ash shoots showing dieback symptoms in the Wolica Nature Reserve in Poland, as well as from shoots of two years-old ash seedlings inoculated with H. fraxineus in the greenhouse. The most frequently isolated fungi from shoots of common ash (associated with the pathogenic fungus H. fraxineus) were identified on the basis of sequencing of the internal transcribed spacer region (ITS1) of fungal rDNA. In total, 19 fungal taxa were identified for ash shoots as follows: H. fraxineus, Fusarium avenaceum, Alternaria spp., Phomopsis oblonga, Diplodia mutila and other Phomopsis spp. The pathogen H. fraxineus was not found for all the shoots samples; one year after inoculation the aforementioned fungi and other species as: Alternaria alternata, Bionectria ochroleuca, Epicoccum nigrum, F. acuminatum, F. avenaceum, and Paraphaesphaeria neglecta were identified in inoculation point, as well as H. fraxineus. The same quantitative and qualitative changes of organisms were observed in the case of artificially colonised ash seedlings in the greenhouse, as well as in the shoots of adult ash trees in the forest. [ABSTRACT FROM AUTHOR]

Societal Impact Statement: Damage to ash trees by ash dieback caused by the emerging fungal pathogen Hymenoscyphus fraxineus is impacting people across Europe. This poses challenges to: public safety; productivity of commercial forestry; green spaces and human wellbeing; and ecosystem services and carbon sequestration. Here, we seek to quantify the impact of ash dieback on tree mortality by analyzing surveys counting the proportion of trees that have died in sites across Europe. However, more and better data are needed to inform policy makers, foresters, conservationists, and other stakeholders as they plan for a long‐term future with ash dieback. Summary: The ash dieback epidemic, caused by the fungus Hymenoscyphus fraxineus, has been present in Europe for over 20 years and caused widespread damage and mortality in ash tree (Fraxinus excelsior) populations. Ash is a major natural capital asset and plays an important role in nature's contribution to people in Europe.Here, we present a meta‐analysis of surveys of ash mortality due to ash dieback, and a time‐dependent model to estimate longer term mortality. In plantations established previous to the arrival of the epidemic, we analyze 12 surveys, finding a maximum recorded mortality of ~85%. In woodlands with exposure to ash dieback of between 4 and 20 years, we analyze 36 surveys, finding a maximum recorded mortality (which may have missed some dead trees) of ~70%. We also analyze 10 surveys of naturally regenerated saplings, finding maximum recorded mortality of ~82%. We apply logistic models to these data sets to seek longer term predictions.More data are needed before our models can be relied upon for policy decisions. If survival found so far in woodlands is due in part to heritable resistance, natural selection or a breeding program may allow future recovery of ash populations in Europe. We review surveys of ash tree death due to ash dieback in Europe and find a maximum mortality so far of 70% of trees in woodlands and 85% of trees in plantations established before the epidemic. It is still hard to predict what the long‐term outcomes will be, and more data are needed. [ABSTRACT FROM AUTHOR]

Ash dieback caused by a non-native pathogenic fungus Hymenoscyphus fraxineus has been decimating populations of European Fraxinus species for over 30 years. Nevertheless, there is still a considerable amount of valuable ashes of this species in European cities and landscapes. Although it has been shown in many studies that the effect of H. fraxineus on adult and senescent trees tends to be rather chronic, the infection by this fungus also causes a substantial decrease in the lifespan of these trees. This research is dealing with the control of ashes using the tree injection technology. It is focused on the development of bark necroses, as this symptom results in the highest damage to a tree. Trees were first inoculated with two different strains of H. fraxineus. Trees with proven positive inoculation were injected with six different fungicides. Subsequently, over the course of nine months, the area of developing bark necroses was measured using image analysis methods. After the field part of the experiment was finished, the extent of the infection by H. fraxineus in tissues was determined and confirmed by PCR. Three out of six tested variants (Tr. 1: 7.1% propiconazole, Tr. 3: 7% propamocarb and 2.7% fosetyl, and Tr. 6: 0.015% sodium selenite) displayed significant deceleration of the growth of bark necroses compared to control. Tr. 3 demonstrated the highest effectiveness with only a very small increment of the necrosis area during the first three months after the injection treatment and a follow-up complete cessation of growth. Reisolations confirmed high mortality of H. fraxineus after this treatment. Hence the presented study can contribute to substantial prolongation of the lifespan of ashes in Europe. • Tree injection is an effective protection of the ash against Hymenoscyphus fraxineus. • Three preparations showed to be effective, greatly decelerating the lesion growth. • A mixture of 7% propamocarb and 2.7% fosetyl can eliminate the pathogen in tissues. [ABSTRACT FROM AUTHOR]

Damage caused by Hymenoscyphus fraxineus , the causal agent of ash dieback, was genetically evaluated based on scorings made on five occasions during the years 2006–2016 at two seed orchards of ash ( Fraxinus excelsior L.) established in southern Sweden in 1992 and 1995, respectively. The studied population included grafts originating from 106 plus-tree clones selected from 27 stands in southern Sweden. The study verified previous results of (1) high genetic control of damage traits, (2) high genotypic variation in disease susceptibility and (3) that only a small proportion of the natural population are at least partly resistant. The clonal response for dieback damage was stable over the 10-year period, i.e. the least susceptible clones selected in 2006 still belong to the healthiest ones after several years of heavy infection pressure. There were no clear trends in the development of damage over time for trees being alive at each assessment. However, the proportion of trees that died was substantial (7–8% per year), but this estimation was influenced by other factors than the disease, such as wild game and sanitary cuttings. Altogether, the results are promising for initiating breeding programs to improve the dieback resistance. However, this is a heavy and costly task, since only a very small proportion of the ash trees are partially resistant to the dieback disease. Thus, a joint breeding program involving all the northern European countries is justified. [ABSTRACT FROM AUTHOR]

Ash dieback, caused by Hymenoscyphus fraxineus, is developing rapidly across the island of Ireland. Ireland's ash (Fraxinus excelsior L.) woodlands, particularly first rotation plantations, are quite unique and are at particular risk of very rapid decline. Urgent action is required in order to minimise the economic, ecological and social impact of the disease. However, for this to happen forest owners require guidance regarding potential positive management interventions. This article outlines the wider, mainly European, experience of remedial silviculture. It presents three case studies on existing remedial silviculture trials in Ireland. In the absence of silvicultural research data specific to the evolving situation with ash dieback, this article explores the potential benefits of positive practical actions which may minimise the impact of the disease. Despite the seriousness of the situation, such silvicultural activity may even result in a positive economic outcome. It is hoped that by beginning to document potential mitigatory management options, this paper may bring some reassurance to owners and managers of ash-dominated woodlands. [ABSTRACT FROM AUTHOR]

The alien invasive pathogen Hymenoscyphus fraxineus causes large-scale decline of European ash (Fraxinus excelsior). We assessed ash dieback in Germany and identified factors that were associated with this disease. Our assessment was based on a 2015 sampling of national forest inventory plots that represent a supra-regional area. In the time from 2012 to 2015, the number of regrown ash trees corresponded to only 42% of the number of trees that had been harvested or died. Severe defoliation was recorded for almost 40% of the living trees in 2015, and more than half of the crowns mainly consisted of epicormic shoots. Necroses were present in 24% of root collars. A total of 14% of the trees were in sound condition, which sum up to only 7% of the timber volume. On average, trees of a higher social status or with a larger diameter at breast height were healthier. Collar necroses were less prevalent at sites with a higher inclination of terrain, but there was no evidence for an influence of climatic variables on collar necroses. The disease was less severe at sites with smaller proportions of the basal area of ash compared to the total basal area of all trees and in the north-eastern part of the area of investigation. The regeneration of ash decreased drastically. [ABSTRACT FROM AUTHOR]

A large part of the area in Europe in which Fraxinus excelsior is native is currently affected by ash dieback, a threatening disease caused by the ascomycetous fungus Hymenoscyphus fraxineus. Fungi other than H. fraxineus also occur in large numbers on stems of the dying ash trees. To clarify their possible role in the dieback process, six fungal species common on dying stems and twigs of ash in Poland, i.e. Cytospora pruinosa, Diaporthe eres, Diplodia mutila, Fusarium avenaceum, F. lateritium and F. solani, were tested for pathogenicity using a test based on artificial wound inoculations of 6-year-old F. excelsior plants under field conditions, with H. fraxineus included for comparison. There were significant differences in index of pathogenicity among the fungi tested. Hymenoscyphus fraxineus (mean index 5.78) was the most pathogenic. Diplodia mutila (4.23) and C. pruinosa (4.02) were significantly less pathogenic than H. fraxineus, but significantly more than the other fungi. Diaporthe eres (2.43), F. avenaceum (1.92), F. solani (1.86) and F. lateritium (1.08) were the least pathogenic ( P < 0.0001). The extent of disease symptoms caused by F. solani and F. lateritium was statistically similar to the control ( P = 0.05). All tested fungi were successfully reisolated from inoculated stems. The contribution of the results to understanding the possible role of these fungi in the ash dieback process in F. excelsior, particularly in trees weakened after primary infection by H. fraxineus, is discussed. [ABSTRACT FROM AUTHOR]

Communities of endophytic fungi in leaves of Manna ash ( Fraxinus ornus ) were examined to both the north and south of the Alps, i.e. within and beyond the native range of this tree species. Almost all leaves examined had been colonized by endophytic fungi. One hundred and two morphotypes were found, and 62 of them were identified to genus or species level using ITS sequencing and micromorphology. Venturia orni was most frequent and occurred in almost one third (32%) of the 1536 examined leaf segments. It was five times more abundant than Colletotrichum acutatum , the second most frequent endophyte. Other frequently isolated endophytes include Paraconiothyrium sp. 1, Mycosphaerella aurantia, Septoria cretae, Botryosphaeria dothidea and Boeremia exigua. The ash dieback pathogen was not isolated. The endophyte communities differed between the north and south of the Alps and the individual tree types had a distinct influence within sites. [ABSTRACT FROM AUTHOR]

Hymenoscyphus fraxineus causes a destructive invasive tree disease known as ash dieback threatening the survival of common ash not only in the forests, but also in urban and landscape settings. Pruning is a potential management practice that could help maintain tree vitality and aesthetics in parks, gardens, alleys and recreation areas, as well as maintaining veteran trees having high heritage or cultural value, or trees with high genetic importance (e.g. located in clonal seed orchards). In this study we investigated the maximum distance proximal to the lesion margin at which H. fraxineus can be detected on individual branches infected by the fungus in order to provide recommendations for pruning. Pruning of branches was carried out on 38 trees in southern Sweden. Tissue samples including bark and wood were collected from the margin of the lesion and at 5 cm intervals proximal to the lesion. Molecular analysis revealed presence of H. fraxineus in 91.3% of the investigated lesions. The proportion of lesions at which H. fraxineus could be detected declined with increasing distance from the lesion margin, with a significant reduction in the number of positive samples at 10 cm proximal to the margin. At 30 cm from the lesion edge the pathogen was never detected. Our results suggest that routine pruning may help maintain the vitality of younger trees. Pruning branches at least 35 cm from visible, active lesions in the bark should exclude the fungus and therefore reduce the probability of stem infection by H. fraxineus , however this cultural control tactic may only be economically feasible for high value amenity trees. [ABSTRACT FROM AUTHOR]

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The alien invasive pathogen Hymenoscyphus fraxineus causes large-scale decline of European ash (Fraxinus excelsior). We assessed ash dieback in Germany and identified factors that were associated with this disease. Our assessment was based on a 2015 sampling of national forest inventory plots that represent a supra-regional area. In the time from 2012 to 2015, the number of regrown ash trees corresponded to only 42% of the number of trees that had been harvested or died. Severe defoliation was recorded for almost 40% of the living trees in 2015, and more than half of the crowns mainly consisted of epicormic shoots. Necroses were present in 24% of root collars. A total of 14% of the trees were in sound condition, which sum up to only 7% of the timber volume. On average, trees of a higher social status or with a larger diameter at breast height were healthier. Collar necroses were less prevalent at sites with a higher inclination of terrain, but there was no evidence for an influence of climatic variables on collar necroses. The disease was less severe at sites with smaller proportions of the basal area of ash compared to the total basal area of all trees and in the north-eastern part of the area of investigation. The regeneration of ash decreased drastically.

Intense dieback of Fraxinus excelsior L. has been causing rapid changes in advance growth of trees and understory shrub growth of the affected stands. In this study, changes in composition and density of understory were studied in 15 permanent plots (each 235.6 m2), repeatedly sampled in 2005, 2010, and 2015. Within each plot, the number and average height of understory individuals were determined. The successional changes in understory were assessed by Detrended Correspondence Analysis. In total, 11 advance growth and 20 undergrowth species were recorded. A significant increase in the density of understory was observed only in 2015, mainly due to understorey growth of Corylus avellana L., Padus avium Mill., and Lonicera xylosteum L. Regarding advanced growth, the highest density was observed for Ulmus glabra Huds., F. excelsior and Acer platanoides L.; the density of A. platanoides and F. excelsior increased particularly in the period from 2010-2015. The observed successional changes suggested individuality of development of the affected stands according to the composition of the remaining and neighbouring canopy trees. [ABSTRACT FROM AUTHOR]

Development of effective biocontrol procedures using ash endophytes to combat an ash pathogen Hymenoscyphus fraxineus would be an appropriate contribution to the ongoing effort to protect European ash stands against ash decline. In this study we investigated the biocontrol potential of two ash endophytes, Thielavia basicola and Minimidochium sp., against H. fraxineus using in vitro plant-fungus and fungus-fungus dual cultures approach in three biocontrol models. The tests aimed to determine whether the endophytes show antagonism toward Fraxinus excelsior and F. pennsylvanica, to assess the level of antagonism of the endophytes toward H. fraxineus and to identify potential secondary metabolites induced by the presence of H. fraxineus. The results that dual culture experiments modeled according to our design may be a very useful tool to precisely study biocontrol potential of fungi, i.e., without the impact of environmental factors. Such experiments also enable the selection of most resistant ash genotypes and rapid propagation, producing large numbers of pathogen-free seedlings. It should be noted, however, that both of the endophytes tested in the dual cultures strongly inhibited the growth of H. fraxineus. Their growth under the influence of callus/seedlings was also inhibited. Comparison of HPLC profiles showed that the presence of H. fraxineus in the post-culture medium induced the production of an unknown secondary metabolite in this species. Such results suggest that some of the plant–fungus combinations examined in this study may have potential to be developed as biocontrol methods, thus increasing the survivability of ash stands under natural conditions. [ABSTRACT FROM AUTHOR]