R. Vasaitis’s research while affiliated with Swedish University of Agricultural Sciences and other places

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Publications (132)


Map of Europe showing the distribution of Pinus sylvestris and Pinus halepensis and the 4 sampling sites. Coding for the sampling sites is given in Table 1.
Rarefaction curves showing the relationship between the cumulative number of fungal OTUs and ITS2 rDNA sequences from each country (LT—Lithuania; MN—Montenegro; UA—Ukraine) as well as all data combined.
The Venn diagram, based on operational taxonomic units (OTUs), represents the common and unique OTUs in each group, showing the distributional differences in fungi among the three countries: Lithuania (LT), Montenegro (MN) and Ukraine (UA). Violet, salmon and green colors indicate unique OTUs in Lithuania (LT), Montenegro (MN) and Ukraine (UA) respectively. Rose color indicates common OTUs in Lithuania (LT) and Montenegro (MN), blue color indicates common OTUs in Lithuania (LT) and Ukraine (UA), beige color indicates common OTUs in Montenegro (MN) and Ukraine (UA). Dusty purple color Montenegro (MN), blue color indicates common OTUs among the three countries: Lithuania (LT), Montenegro (MN) and Ukraine (UA).
Ordination diagram based on detrended correspondence analysis of fungal communities from Tomicus species collected in Lithuania (polygon—purple; points—squares), Montenegro (polygon—rose; points—circles), and Ukraine (polygon—green; points—diamonds). The size of individual points (squares, diamonds or circles) represents the relative richness of fungal OTUs.
Non-metric multidimensional scaling (NMDS) plot of the fungal community. NMDS of dissimilarity based on a Bray–Curtis distance matrix of rarefied fungal OTU abundances. The dashed lines on the NMDS plot indicate the relative positioning of fungal communities in ordination space. The values (−1, 0, 1 for NMDS1 and −1, −0.5, 0, 0.5, 1 for NMDS2) correspond to the ordination scores, which reflect differences in fungal community composition based on Bray–Curtis dissimilarity. Closer points represent more similar communities, while greater distances indicate higher dissimilarity.

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Fungal Pathogens Associated with Tomicus Species in European Forests: Regional Variations and Impacts on Forest Health
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March 2025

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64 Reads

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Denys Baturkin

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Simple Summary This study explored the relationship between pine bark beetles, Tomicus spp. and fungi in forests of three European countries (Lithuania, Ukraine, Montenegro) with different climates. Bark beetles are tiny insects that attack trees, often carrying fungi that can harm forests. The study aimed to understand the types of fungi associated with these beetles and their potential impact on forest health. The results revealed significant differences in fungal communities across the regions, influenced by local environmental conditions and tree species. Some fungi, such as those causing blue-stain or shoot blight, are harmful to tree health and were found in high numbers in certain areas. Interestingly, some fungi that typically support beetle development were also discovered, showing how these fungi–beetle interactions can be both beneficial and harmful. The findings highlight the role of these beetles in spreading harmful fungi, especially as climate change weakens trees and creates favorable conditions for beetles. This research is valuable for understanding how beetles and fungi interact and showing the risks they pose to forests. It provides important information for forest management strategies to protect trees from these threats, especially in the face of environmental changes that could exacerbate the problem.

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Infestation by Ips amitinus (Eichhoff, 1872), Its Associated Fungi, and Butt Rots in Stands of Pinus sibirica in South-Western Siberia

December 2023

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93 Reads

In 2019, the bark beetle Ips amitinus (native to central Europe) was identified in south-western Siberia at a distance exceeding 2500 km east of its previously known easternmost location in the European part of Russia. In Siberia, its invasive populations are characterised by high abundance and harmfulness. Here, I. amitinus accomplishes primary attacks on standing vital trees of Pinus sibirica with a lethal outcome. This invasion has already resulted in massive dieback in stands of pine over a large geographic territory. By, 2021, the invaded area was estimated to cover at least 31,200 km². The objectives of this study were to investigate fungi associated with/vectored by I. amitinus in its invasive area in south-western Siberia and wood decay fungi that cause root and butt rots to P. sibirica. This led to the following conclusions: (i) DNA analysis of sixty adult beetles of Ips amitinus collected from P. sibirica in south-west Siberia revealed the presence of 143 fungal taxa; (ii) species richness was significantly higher in beetles collected from dead branches than from (more recently infested) dying branches; (iii) fungal communities were >90% dominated by yeasts, among which the most common were Nakazawaea holstii, Kuraishia molischiana, and N. ambrosiae; (iv) entomopathogenic Beauveria bassiana s.l. was the most common fungus isolated from dead/mycosed beetles of I. amitinus, followed by Lophium arboricola and four Ophiostoma spp.; and (v) Heterobasidion parviporum was the most common decay fungus detected, which was causing heart rot in stems of P. sibirica.


Sixty-three-year-old resin tapping wounds on 135-year-old stem of Pinus sylvestris (Site 1—Kalsnava); a wound on a standing tree; b close photo of wound surface; note width of scars of approx. 0.5 cm, and “step” between scars of approx. 1 cm; c cross section of two resin tapping wounds showing approx. 3–4 cm lateral closure from each margin in about 4 decades; d longitudinal section of a wound showing up to 1 cm-deep reddish sapwood discoloration
Mean tree ring width (µm) for resin tapped and control pines. Orange bold line—tree ring width of resin tapped trees, gray line—tree ring width of control trees
Long-term pathological consequences of resin tapping wounds on stems of Scots pine (Pinus sylvestris L.)

May 2022

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193 Reads

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12 Citations

Trees

Key message After 5–6 decades since inflicting resin tapping wounds, overmature (> 120 years old) Pinus sylvestris stems remain undecayed and vigorous. Abstract Overmature trees of Pinus sylvestris bearing large wounds made by resin tapping decades ago are still present in woodlands of south-eastern Baltic Sea region. The aim of the present study was to investigate health condition of those trees focusing on fungal infections and to estimate impact of the injury on radial stem growth. The study was conducted in Latvia in three overmature stands of P. sylvestris , resin-tapped in 1950–1970 s. On the studied ninety 120–167-year-old trees, exposed sapwood constituted from 1140 to 7755 cm ² per individual stem. Of the 127 wounds sampled, 52 (41%) showed wood discoloration. The discoloration in its extent was limited, expanding beyond wound margins approx. 1 (max 3) cm in radial, and 6–7 cm in longitudinal directions. Of the 127 wood samples/wounds subjected to fungal isolations, 96% resulted in fungal growth, yielding 236 fungal isolates that represented 47 fungal taxa. The most common among macro-fungi was basidiomycete Porodaedalea pini , which was isolated from 9% of stems. The fungus is currently classed not as a tree pathogen, but instead as an indicator species for woodland sites to be considered for nature conservation. Data from tree ring widths have revealed that tree reacted to the resin tapping injury by increasing radial increment of the un-affected part of the circumference of the stem. Current study demonstrated that even on the long term, resin tapping has little influence on health condition and vitality of P. sylvestris , even at the very old age. This should be considered as a supporting message in case resin taping practices in the region are to be revived.



Similarities in fungal community structures between the study sites (Sørensen qualitative similarity index).
Pathogenicity tests with ophiostomatoid fungi (isolated from wood beyond Ips sexdentatus galleries and beetles) inoculated to 3-4-year-old saplings of Pinus sylvestris.
Fungal Communities Vectored by Ips sexdentatus in Declining Pinus sylvestris in Ukraine: Focus on Occurrence and Pathogenicity of Ophiostomatoid Species

December 2021

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135 Reads

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12 Citations

Drought-induced stress and attacks by bark beetle Ips sexdentatus currently result in a massive dieback of Pinus sylvestris in eastern Ukraine. Limited and fragmented knowledge is available on fungi vectored by the beetle and their roles in tree dieback. The aim was to investigate the fungal community vectored by I. sexdentatus and to test the pathogenicity of potentially aggressive species to P. sylvestris. Analysis of the fungal community was accomplished by combining different methods using insect, plant, and fungal material. The material consisted of 576 beetles and 96 infested wood samples collected from six sample plots within a 300 km radius in eastern Ukraine and subjected to fungal isolations and (beetles only) direct sequencing of ITS rDNA. Pathogenicity tests were undertaken by artificially inoculating three-to-four-year-old pine saplings with fungi. For the vector test, pine logs were exposed to pre-inoculated beetles. In all, 56 fungal taxa were detected, 8 exclusively by isolation, and 13 exclusively by direct sequencing. Those included nine ophiostomatoids, five of which are newly reported as I. sexdentatus associates. Two ophiostomatoid fungi, which exhibited the highest pathogenicity, causing 100% dieback and mortality, represented genera Graphium and Leptographium. Exposure of logs to beetles resulted in ophiostomatoid infections. In conclusion, the study revealed numerous I. sexdentatus-vectored fungi, several of which include aggressive tree pathogens.


Fig. 1. Agar plate pairings with compatible mated monokaryotic mycelia from Hapalopilus croceus. Clamp connections were recorded both from plates without (A) and with a barrage zone (B). Compatible pairings were indistinguishable without microscopy. (C) Mated hyphae of H. croceus with clamp connections (black arrows). Note the presence of chlamydospores (asterisks). (D) Unmated hyphae of H. croceus, lacking clamp connections.
The mating type system of the rare polypore Hapalopilus croceus

June 2020

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492 Reads

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1 Citation

Fungal Ecology

Hapalopilus croceus is a rare, wood-decay polypore associated with old coarse broadleaf trees, primarily Quercus. It is mainly distributed throughout the temperate zone of the Northern Hemisphere with its main distribution in Europe. Mating systems in fungi influence the probability of in- or outbreeding and may affect the species’ fitness. We found the mating system of H. croceus to be bipolar by pairing 18 single spore isolates from one sporocarp. We discuss this finding from the perspective of H. croceus as a threatened species with a very small and fragmented current population associated with a declining habitat and globally red-listed as Vulnerable (VU).


Natural infection and colonization of pre-commercially cut stumps of Picea abies and Pinus sylvestris by Heterobasidion rot and its biocontrol fungus Phlebiopsis gigantea

April 2020

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80 Reads

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10 Citations

Biological Control

Fungi from the genus Heterobasidion are among the most important pathogens of forest trees in Northern Hemisphere causing root rot and wood decay, while Phlebiopsis gigantea is a very common saprotrophic wood decay fungus. Both fungi are primary colonizers of freshly cut conifer stumps (through which Heterobasidion spp. accomplishes primary infections of tree root systems), thus both fungi are competing for the substrate. To date, P. gigantea is widely used as Heterobasidion spp. biocontrol agent. Hypothesis has been proposed that natural colonization of stumps by P. gigantea might also to some extent restrict infections by the pathogen. The main aim of the study was to assess the potential of natural infections of P. gigantea to restrict infection and spread of Heterobasidion spp. in Picea abies and Pinus sylvestris stumps. In total, 793 P. abies stumps and 1158 P. sylvestris stumps were examined in 24 sample plots located in the eastern part of Latvia. Of these, 325 (41.0%) P. abies stumps were infected by Heterobasidion spp., and 59 (7.4%) by P. gigantea, and 168 (14.5%) P. sylvestris stumps were infected by Heterobasidion spp., and 846 (73.1%) by P. gigantea. In P. abies, the observed Heterobasidion spp. infection frequencies were significantly (p < 0.05) higher than those of P. gigantea, while the respective situation in P. sylvestris was reverse and P. gigantea infections were more frequent (p < 0.05). The mean surface area colonized by Heterobasidion spp. in P. abies and P. sylvestris stumps was 5.7 and 5.3 cm² and did not differ significantly (mean coverage of stump surface area respectively 18% and 13%; p = 0.41). In contrast, the mean surface area colonized by P. gigantea was significantly different in the two tree species, respectively, 3.9 and 21.3 cm² (16% and 59%; p < 0.05). The mean surface area colonized by Heterobasidion spp. in P. abies stumps was significantly larger (p < 0.05) than the area colonized by P. gigantea, while conversely, the mean area colonized by P. gigantea in P. sylvestris stumps was significantly larger (p < 0.001) than that colonized by Heterobasidion spp. Both fungi were co-occurring in P. abies stumps in 33 cases (4.2% of all investigated stumps), and in P. sylvestris stumps in 138 cases (11.9%). There were no correlations between the sizes of colonized areas of Heterobasidion spp. and P. gigantea in P. abies stumps (r = 0.06; p = 0.76), or P. sylvestris stumps (r = 0.009; p = 0.27). In conclusion, the results of this study strongly suggest that even in stumps of P. sylvestris, that otherwise are much preferred for natural colonization by airborne spores of the biocontrol agent P. gigantea, natural colonization by P. gigantea is not able to restrict infections by Heterobasidion spp. This clearly indicates that for effective biocontrol of Heterobasidion spp. infections, the necessity for thorough treatment coverage of cut P. abies and P. sylvestris stumps at early stages of plantation management, during pre-commercial thinning.


Occurrence and pathogenicity of Corinectria spp. – an emerging canker disease of Abies sibirica in Central Siberia

March 2020

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806 Reads

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3 Citations

During recent years, a new disease of Siberian fir (A. sibirica) emerged in Central Siberia, exhibiting symptoms of stem/branch deformation, cambium necrosis, and dieback of branches and twigs, the causal agent remaining unknown. The aim was to identify agent of the disease and to investigate its pathogenicity to A. sibirica and Norway spruce (Picea abies). Symptomatic tissues of fir were subjected to pure culture isolation of anticipated pathogen(s). Obtained isolates were subjected to molecular identification, phylogenetic analyses, and pathogenicity tests with A. sibirica saplings, and seeds and seedlings of A. sibirica and P. abies. The study demonstrated that, (i) most commonly isolated fungus from canker wounds of A. sibirica exhibited Acremonium-like anamorphs; (ii) phylogeny demonstrated that investigated fungi belong to genus Corinectria, but are genetically well separated from other worldwide known Corinectria spp.; (iii) one species of isolated fungi has the capacity to cause the disease and kill A. sibirica saplings and seedlings, but also seedlings of P. abies. Guidelines for future research were defined in order to generate needed information on species description, its origin and ecology, and estimation of potential risks upon the eventual invasion of the pathogen to new geographic areas, in particular of Europe.


Identifying Fraxinus excelsior tolerant to ash dieback: Visual field monitoring versus a molecular marker

December 2019

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235 Reads

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17 Citations

Forest Pathology

Ash dieback (ADB) caused by the pathogen Hymenoscyphus fraxineus is the cause of massive mortality of Fraxinus spp. in Europe. The aim of this work was to check for the presence of the molecular marker for ADB tolerance in mapped healthy‐looking F. excelsior trees, and to compare its occurrence in trees exhibiting severe ADB symptoms. Monitoring of 135 healthy‐looking F. excelsior on the island of Gotland, Sweden, showed that after 3–4 years 99.3% of these trees had 0%–10% crown damage, thus remaining in a similar health condition as when first mapped. After 5–6 years, 94.7% of these trees had 0%–10% crown damage. Molecular analysis of leaf tissues from 40 of those showed the presence of the molecular marker in 34 (85.0%) trees, while it was absent in 6 (15.0%) trees. Analysis of leaf tissues from 40 severely ADB‐diseased trees showed the presence of the molecular marker in 17 (42.5%) trees, but its absence in 23 (57.5%) trees (p < .0001). The results demonstrated that monitoring of healthy‐looking F. excelsior is a simple and straightforward approach for the selection of presumably ADB‐tolerant ash for future breeding. The cDNA‐based molecular marker revealed moderate capacity on its own to discriminate between presumably ADB‐tolerant and susceptible F. excelsior genotypes.


Figure 1. Characteristics of study site and sampling strategy, showing (A) geographic location of site, (B) photographs of forest floor developmental gradient and (C) number of times each microhabitat type (MN = mineral soil without roots, MR = mineral soil with roots, O = organic soil) was sampled at each gradient position and within each depth category.
Figure 2. PCA of physiochemical variables [soil moisture (%), pH, P i concentration (NaHCO3 extractable PO4; μg/kg soil), cation concentration (Al, Ca, Fe, K, Mn, Mg, Na; mg/kg soil)] with relation to microhabitat type [MN = mineral soil without roots (n = 46), MR = mineral soil with roots (n = 30), O = organic soil (n = 9)]. All variables except pH were log transformed, centered and scaled prior to analysis. Percentage of variance explained by each PC is given in parentheses. Values are based on data for 85 samples from transects A, B and C. Samples with missing data were excluded in analysis. See Figure S1 (Supporting Information) for PCA displayed to show patterns relating to gradient position and depth.
Figure 3. Relationship between microhabitat type [MN = mineral soil without roots (n = 86), MR = mineral soil with roots (n = 40), O = organic soil (n = 24)] and community composition displayed using (A) nMDS based on Bray-Curtis dissimilarities between samples (stress = 0.2; n = 150 samples, 1688 OTUs) and (B) stacked bar plots of average relative proportion of ITS2 sequence reads for prominent fungal classes within each habitat. Dissimilarity matrix was calculated after transforming the OTU counts to per-sample relative sequence read abundances (A). Per-sample relative sequence read abundances were calculated before pooling samples by microhabitat type (B).
Figure 4. Association between core community OTUs and microhabitat type [MN = mineral soil without roots (n = 86), MR = mineral soil with roots (n = 40), O = organic soil (n = 24)]. Average relative ITS2 sequence read abundance (%) for each OTU within each microhabitat type is indicated within each cell. Cell color is based on row-wise centered and scaled values (mean = 0, standard deviation = 1). Warm cell colors indicate positive association. OTUs are labeled with highest assigned taxonomy and unique OTU identifier. Colors along right side indicate functional characteristics relating to trophic mode and growth morphology (ECMmat = ectomycorrhizal, medium-distance mat-forming exploration type; ECMshort = ectomycorrhizal, short-distance exploration type; root = root-associated; rootEE = root-associated, ect-endomycorrhizal; rootDSE =
Distribution patterns of fungal taxa and inferred functional traits reflect the non-uniform vertical stratification of soil microhabitats in a coastal pine forest

September 2019

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151 Reads

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11 Citations

FEMS Microbiology Ecology

In boreal systems, soil profiles typically consist of distinct stratified horizons, with organic layers at the surface overlying deeper mineral horizons providing microhabitat variation along a depth gradient, and vertical stratification of fungal communities along such soil profiles is commonly observed. We studied fungal community structure in a coastal pine forest along a gradient of decreasing influence from the coast. In this system, the vertical stratification pattern of soil microhabitats (defined here as organic, mineral with roots, and mineral without roots; O, MR, and MN, respectively) is non-uniform; organic horizons are sometimes buried under drifting sand dunes. Our results show that soil microhabitats are distinct with respect to physiochemical characteristics, community composition, and OTU richness. While community composition was partly related to depth and distance from the coastal forest edge, microhabitat appeared to have the strongest influence. A closer inspection of the OTUs with the highest relative sequence abundance within each microhabitat revealed that microhabitats support functionally distinct fungal communities with respect to trophic mode and growth morphology. These results suggest that in coastal pine forests, variation in soil microhabitats contributes to the high fungal diversity found below-ground and may play an important role in optimizing nutrient cycling.


Citations (53)


... While tapped trees occasionally matched untapped growth rates, the cumulative effects of tapping led to long-term growth reductions, as observed in Pinus sylvestris in Sweden (Zaluma et al., 2022). Growth imbalance, with narrower rings on tapped sides, mirrors findings from other studies (Zeng et al., 2021). ...

Reference:

Impact of resin tapping on the radial growth and climate sensitivity of naturally- regenerated Pinus roxburghii (Chir pine) in Western Nepal
Long-term pathological consequences of resin tapping wounds on stems of Scots pine (Pinus sylvestris L.)

Trees

... This pest is capable of inhabiting trees of numerous species within the Pinaceae family, most frequently the genus Pinus. It is widely distributed across Eurasia, including the countries of South Asia [7][8][9][10][11]. In 2023, the United States Department of Agriculture (USDA) designated this species as a priority pest with a high potential for invasion [12]. ...

Fungal Communities Vectored by Ips sexdentatus in Declining Pinus sylvestris in Ukraine: Focus on Occurrence and Pathogenicity of Ophiostomatoid Species

... In previous work, P. gigantea has been found to be abundant in native pine forests in Minnesota and it is possibly acting as a natural biological control agent (Otto, Held, Gould, & Blanchette, 2021;Otto, Held, Redford, & Blanchette, 2021). Previous research has suggested that natural colonization by P. gigantea is not able to effectively control Heterobasidion based on the extent of colonization in Norway spruce and Scots pine stumps investigated in Europe (Gaitnieks et al., 2020;Rishbeth, 1963). However, these studies in- A potential future study could compare native isolates to commercial strains to compare their efficacy. ...

Natural infection and colonization of pre-commercially cut stumps of Picea abies and Pinus sylvestris by Heterobasidion rot and its biocontrol fungus Phlebiopsis gigantea
  • Citing Article
  • April 2020

Biological Control

... Specific single nucleotide polymorphism (SNP) and gene expression markers were found to be associated with the resistance to ash dieback in a Danish clonal trial (Harper et al. 2016;Sollars et al. 2017). However, the accuracy of these predictions that use only a few markers were low (Sollars et al. 2017;Menkis et al. 2020). Accordingly, susceptibility to ash dieback might have a polygenic basis (Kjaer et al. 2012). ...

Identifying Fraxinus excelsior tolerant to ash dieback: Visual field monitoring versus a molecular marker

Forest Pathology

... In certain studies, specific EM fungi, corresponding to characteristic exploration types, have been found to reflect different strategies of colonization along the soil profile (Anderson et al. 2014;Genney et al. 2006;Pickles and Anderson 2016). As an example, Kluting et al. (2019) reported that soil microhabitats support functionally distinct fungal communities with respect to trophic mode and growth morphology, with short-distance EM species being most closely associated with organic layers, while mat-forming EM taxa, in particular Sarcodon genera, being generally dominant in mineral horizons. Consecutively, these morphological traits of EM fungi have been proposed as functional traits that could link EM hyphal morphologies to different nutrient acquisition capacities (Agerer 2006;Defrenne et al. 2019;Zanne et al. 2020). ...

Distribution patterns of fungal taxa and inferred functional traits reflect the non-uniform vertical stratification of soil microhabitats in a coastal pine forest

FEMS Microbiology Ecology

... Present-day F. angustifolia and F. excelsior populations are susceptible to dieback caused by the fungus Hymenoscyphus fraxineus, also known as Chalara fraxinea (first reported in the 1990s), which causes high mortality, especially in young seedlings, resulting in limited natural regeneration (Enderle et al., 2019, Kirisits et al., 2010, Kowalski, 2006. In addition, ashes are also threatened by invasive pests (emerald ash borer; Agrilus planipennis), which produce massive damage and tree mortality (EFSA et al., 2020, Evans et al., 2020, Orlova-Bienkowskaja, 2014. ...

An overview of ash ( Fraxinus spp.) and the ash dieback disease in Europe.

... H. annosum s.l. primarily uses the basidiospores to infect freshly open wood surfaces and then spread to neighboring hosts by root to root contact (Zal , uma et al., 2019). For decades, the Heterobasidion root rot has been managed by use of the fungal biocontrol agent Phlebiopsis gigantea (Fr.) ...

Infection and spread of root rot caused by Heterobasidion spp. in Pinus contorta plantations in Northern Europe: three case studies

... Значительное количество микромицетов было и на гусеницах златогузки (Euproctis chrysorrhoea Linnaeus, 1758 (Erebidae)) -типичного вредителя листьев дуба (Баранов и др., 2021). Споры возбудителя язвенного рака сосны анаморфного аскомицета Fusarium circinatum Nirenberg & O'Donnell наряду с короедами распространяют сосновые слоники (Pissodes spp.), сосновый семенной клоп (Leptoglossus occidentalis Heidemann, 1910) (Hemiptera: Coreidae) и другие виды (Селиховкин и др., 2018;Selikhovkin et al., 2018;Fernández-Fernández et al., 2019). Фитопатогенный гриб Diplodia pinea (Desm.) ...

Phytopathogenic Fungus Fusarium circinatum and Potential for Its Transmission in Russia by Insects

Russian Journal of Biological Invasions

... Several studies have shown that the susceptibility of both spruce and pine stumps to Heterobasidion spore infection increases with increasing stump diameter [62][63][64][65]. At the same time, the best efficacy of urea treatment is expected in small-diameter stumps in young stands because urease activity is at its highest in young sapwood, while in dead heartwood, the activity is low [11]. ...

Infection of pre-commercially cut stumps of Picea abies and Pinus sylvestris by Heterobasidion spp. – A comparative study

Silva Fennica