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Ecology and Disease in Natural Forests

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... dombeyi. El género Armillaria (Fr.) Staude incluye especies que se encuentran entre las causas más comunes e importantes de pudrición de raíces en plantas leñosas en todo el mundo (Fox, 2000;Kile et al., 1991). Sin embargo, aunque la mayoría de las especies de Armillaria tienen el potencial de infectar árboles sanos y estresados, difieren en su patogenicidad según el huésped y también pueden actuar como necrótrofos y saprobios (Hood et al., 1991;Kile et al., 1991). ...
... El género Armillaria (Fr.) Staude incluye especies que se encuentran entre las causas más comunes e importantes de pudrición de raíces en plantas leñosas en todo el mundo (Fox, 2000;Kile et al., 1991). Sin embargo, aunque la mayoría de las especies de Armillaria tienen el potencial de infectar árboles sanos y estresados, difieren en su patogenicidad según el huésped y también pueden actuar como necrótrofos y saprobios (Hood et al., 1991;Kile et al., 1991). ...
Thesis
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Temperate forests have been threatened, in recent decades, by events of death and decay throughout the globe. The causes have been investigated without, in many cases, revealing a primary cause of the deterioration. This gave rise to a concept that is increasingly used to evaluate these phenomena, which is that of complex diseases. In this context, the endophytic mycobiome of trees plays an important role as a recent object of study and integration in interpretations of forest health. In recent decades, a phenomenon of grouped death of unknown etiology has been observed in the two Nothofagus species with the widest distribution in the forests of North Patagonia: N. dombeyi and N. pumilio. Within the framework of this thesis, the phenomenon of the grouped death of Nothofagus in Los Alerces National Park (Chubut) was described, characterizing for the first time in the field the symptoms, signs, damage and characteristics of the site and the stand. The endophytic mycobiota of stem and root wood of Nothofagus was analyzed in stands with clustered death and beta diversity was described along environmental and host variables as a contribution to its autoecology and in search of possible associations between the mycobiota and the phenomenon of grouped death. The study of these communities was developed through two methodological approaches: one by isolation prospecting and another by massive sequencing of the ITS1 molecular marker from environmental DNA using the MiSeq Illumina platform. The diversity results obtained by both approaches were analyzed, but the performance of the new technologies was also evaluated using the database of sequences obtained by crop prospecting as a comparison criterion. The application of second-generation sequencing technologies for the study of endophytic fungal communities in wood proved to be complementary to culture-dependent approaches and allowed deepening of beta diversity patterns, detecting greater heterogeneity than with culture. Although more pathogenic organisms were recovered from symptomatic than asymptomatic N. dombeyi in the culture study, the independent approach allowed us to detect that both species of Nothofagus carry a wide diversity of potential fungal pathogens in their sapwood, both in trees that present visible symptoms and those of healthy appearance. The grouped death of Nothofagus is revealed as a complex phenomenon in which many biotic and abiotic variables are interacting and in which, in this framework, the wood endophytic mycobiota would be playing a secondary role. Both species of Nothofagus carry different fungal assemblages, with very little overlap, but both very diverse. In N. dombeyi, the site turned out to be the most important factor in the differentiation of the assemblages. The variations observed were related with site-features such as climate, topography and water availability, reflecting the greater heterogeneity of sites in which this species is distributed. Nothofaugs pumilio showed a strong susceptibility to seasonality and greater compartmentalization between root and stem, which translates into an increase in rotting and degrading fungal organisms with warm and drier periods. Similarly, during cold periods, the root acts as a reservoir of diversity of fungal organisms with other trophic strategies. This is definitely something that we are interested in continuing to investigate, given the general context of climate change that temperate forests are facing around the globe.
... dombeyi. El género Armillaria (Fr.) Staude incluye especies que se encuentran entre las causas más comunes e importantes de pudrición de raíces en plantas leñosas en todo el mundo (Fox, 2000;Kile et al., 1991). Sin embargo, aunque la mayoría de las especies de Armillaria tienen el potencial de infectar árboles sanos y estresados, difieren en su patogenicidad según el huésped y también pueden actuar como necrótrofos y saprobios (Hood et al., 1991;Kile et al., 1991). ...
... El género Armillaria (Fr.) Staude incluye especies que se encuentran entre las causas más comunes e importantes de pudrición de raíces en plantas leñosas en todo el mundo (Fox, 2000;Kile et al., 1991). Sin embargo, aunque la mayoría de las especies de Armillaria tienen el potencial de infectar árboles sanos y estresados, difieren en su patogenicidad según el huésped y también pueden actuar como necrótrofos y saprobios (Hood et al., 1991;Kile et al., 1991). ...
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Temperate forests have been threatened, in recent decades, by events of death and decay throughout the globe. The causes have been investigated without, in many cases, revealing a primary cause of the deterioration. This gave rise to a concept that is increasingly used to evaluate these phenomena, which is that of complex diseases. In this context, the endophytic mycobiome of trees plays an important role as a recent object of study and integration in interpretations of forest health. In recent decades, a phenomenon of grouped death of unknown etiology has been observed in the two Nothofagus species with the widest distribution in the forests of North Patagonia: N. dombeyi and N. pumilio. Within the framework of this thesis, the phenomenon of the grouped death of Nothofagus in Los Alerces National Park (Chubut) was described, characterizing for the first time in the field the symptoms, signs, damage and characteristics of the site and the stand. The endophytic mycobiota of stem and root wood of Nothofagus was analyzed in stands with clustered death and beta diversity was described along environmental and host variables as a contribution to its autoecology and in search of possible associations between the mycobiota and the phenomenon of grouped death. The study of these communities was developed through two methodological approaches: one by isolation prospecting and another by massive sequencing of the ITS1 molecular marker from environmental DNA using the MiSeq Illumina platform. The diversity results obtained by both approaches were analyzed, but the performance of the new technologies was also evaluated using the database of sequences obtained by crop prospecting as a comparison criterion. The application of second-generation sequencing technologies for the study of endophytic fungal communities in wood proved to be complementary to culture-dependent approaches and allowed deepening of beta diversity patterns, detecting greater heterogeneity than with culture. Although more pathogenic organisms were recovered from symptomatic than asymptomatic N. dombeyi in the culture study, the independent approach allowed us to detect that both species of Nothofagus carry a wide diversity of potential fungal pathogens in their sapwood, both in trees that present visible symptoms and those of healthy appearance. The grouped death of Nothofagus is revealed as a complex phenomenon in which many biotic and abiotic variables are interacting and in which, in this framework, the wood endophytic mycobiota would be playing a secondary role. Both species of Nothofagus carry different fungal assemblages, with very little overlap, but both very diverse. In N. dombeyi, the site turned out to be the most important factor in the differentiation of the assemblages. The variations observed were related with site-features such as climate, topography and water availability, reflecting the greater heterogeneity of sites in which this species is distributed. Nothofaugs pumilio showed a strong susceptibility to seasonality and greater compartmentalization between root and stem, which translates into an increase in rotting and degrading fungal organisms with warm and drier periods. Similarly, during cold periods, the root acts as a reservoir of diversity of fungal organisms with other trophic strategies. This is definitely something that we are interested in continuing to investigate, given the general context of climate change that temperate forests are facing around the globe.
... Many trees and shrubs in southern Australian eucalypt forests are susceptible to A. luteobubalina infection. Kile et al. (1991), list 81 species from 21 families that are killed by A. luteobubalina in Australia. In the karri forest only 12 species from 6 families, including suppressed eucalypts, have been recorded as hosts (Pearce et al. 1986). ...
... Unless A. luteobubalina can access, infect and colonise stumps within 3-4 years it is excluded by other microorganisms (Kile 1981). Clear felling in disease patches could therefore be a better management practice by reducing the number of stumps infected and the disease level in the subsequent crop (Kile et al. 1991). ...
Conference Paper
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Armillaria luteobubalina is an endemic pathogen of dry sclerophyll forests and woodlands of southern Australia. The fungus infects the roots of susceptible hosts causing a root and butt rot, commonly referred to as Armillaria root disease (ARD). In high quality regrowth karri stands, ARD causes significant losses due to mortality and defect, which are exacerbated by intensive management practices such as thinning. This review gives a brief outline of what is known about Armillaria luteobubalina and Armillaria root disease in karri regrowth forests of Western Australia and discusses options available for disease control.
... Among the seven Armillaria spp. found in Europe, six are widely distributed wood-decay fungi with great ecological and economical importance (KILE et al. 1991; GUILLAUMIN et al. 1993). The distribution and ecological characteristics of Armillaria spp. ...
... rarely in fruit orchards, causing occasional mortality on cherry, apple and pear in the Korçe district in southern Albania, and the Diber district in the north. Armillaria ostoyae, the most important Armillaria pathogen in gymnosperm forests of Europe and North America (KILE et al. 1991), occurs in most European gymnosperm forests with continental or oceanic-type climates, excluding the northernmost boreal forests. In central Europe the fungus is common at low altitudes, whereas in the Mediterranean area it is found only at high altitudes (GUILLAUMIN et al. 1993). ...
Article
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Summary Species of Armillaria were identified from 645 isolates obtained in a nation-wide survey in Albania. The material was collected from ca. 250 permanent plots, established for monitoring forest health, and from forests and orchards attacked by Armillaria. Armillaria mellea s.s. occurred on several coniferous and broadleaved trees in most areas examined, although it was absent above 1100-1200 m in northern Albania. This species damaged Abies and Quercus spp. and, to a lesser extent, other forest trees. A. mellea was also commonly recorded causing damage in orchards and vineyards. Armillaria gallica was a common saprophyte or weak pathogen in coniferous and deciduous forests at altitudes from 600 to 1600 m, and less commonly on oaks at lower altitudes. Armillaria ostoyae was rare in central and southern Albania, but common in northern Albania, causing significant damage to pine and other conifers, mostly at altitudes from 600 to 1800 m. Armillaria cepistipes was recorded at altitudes from 800 to 1800 m as a saprophyte or weak pathogen on conifers and deciduous trees, mostly in beech and silver fir forests. Armillaria tabescens was found in oak forests at altitudes from sea level to 900 m. In orchards, A. tabescens occasionally attacked almond and pear trees. Armillaria borealis was found in a few locations in northern Albania, at altitudes from 800 to 1800 m.
... Armillaria root disease, long recognized as a serious problem in both angiosperm-and gymnosperm-dominated forest ecosystems (Shaw and Kile 1991;Guillamin et al. 2005), is caused by a group of fungi that occurs worldwide in boreal, temperate and tropical forests. Globally, approximately 36 species of Armillaria are recognized ( Kile et al. 1991;Volk and Burdsall 1995;Pegler 2000;Guillamin et al. 2005). ...
... Among the seven Armillaria spp. found in Europe, six are widely distributed wood- decay fungi with great ecological and economical importance ( Kile et al. 1991;Guillaumin et al. 1993). The distribution and ecological characteristics of Armillaria spp. ...
Article
Full-text available
Species of Armillaria were identified from 645 isolates obtained in a nation-wide survey in Albania. The material was collected from ca. 250 permanent plots, established for monitoring forest health, and from forests and orchards attacked by Armillaria. Armillaria mellea s.s. occurred on several coniferous and broadleaved trees in most areas examined, although it was absent above 1100–1200 m in northern Albania. This species damaged Abies and Quercus spp. and, to a lesser extent, other forest trees. Armillaria mellea was also commonly recorded causing damage in orchards and vineyards. Armillaria gallica was a common saprophyte or weak pathogen in coniferous and deciduous forests at altitudes from 600 to 1600 m, and less commonly on oaks at lower altitudes. Armillaria ostoyae was rare in central and southern Albania, but common in northern Albania, causing significant damage to pine and other conifers, mostly at altitudes from 600 to 1800 m. Armillaria cepistipes was recorded at altitudes, from 800 to 1800 m as a saprophyte or weak pathogen on conifers and deciduous trees, mostly in beech and silver fir forests. Armillaria tabescens was found in oak forests at altitudes from sea level to 900 m. In orchards, A. tabescens occasionally attacked almond and pear trees. Armillaria borealis was found in a few locations in northern Albania, at altitudes from 800 to 1800 m.
... Armillaria root disease, long recognized as a serious problem in both angiosperm-and gymnosperm-dominated forest ecosystems (Shaw and Kile 1991; Guillamin et al. 2005), is caused by a group of fungi that occurs worldwide in boreal, temperate and tropical forests. Globally, approximately 36 species of Armillaria are recognized (Kile et al. 1991; Volk and Burdsall 1995; Pegler 2000; Guillamin et al. 2005). Among the seven Armillaria spp. ...
... Among the seven Armillaria spp. found in Europe, six are widely distributed wooddecay fungi with great ecological and economical importance (Kile et al. 1991; Guillaumin et al. 1993). The distribution and ecological characteristics of Armillaria spp. ...
... Armillaria root disease, long recognized as a serious problem in both angiosperm-and gymnosperm-dominated forest ecosystems (Shaw and Kile 1991; Guillamin et al. 2005), is caused by a group of fungi that occurs worldwide in boreal, temperate and tropical forests. Globally, approximately 36 species of Armillaria are recognized (Kile et al. 1991; Volk and Burdsall 1995; Pegler 2000; Guillamin et al. 2005). Among the seven Armillaria spp. ...
... Among the seven Armillaria spp. found in Europe, six are widely distributed wooddecay fungi with great ecological and economical importance (Kile et al. 1991; Guillaumin et al. 1993). The distribution and ecological characteristics of Armillaria spp. ...
Article
Full-text available
Species of Armillaria were identified from 645 isolates obtained in a nation-wide survey in Albania. The material was collected from ca. 250 permanent plots, established for monitoring forest health, and from forests and orchards attacked by Armillaria. Armillaria mellea s.s. occurred on several coniferous and broadleaved trees in most areas examined, although it was absent above 1100-1200 m in northern Albania. This species damaged Abies and Quercus spp. and, to a lesser extent, other forest trees. A. mellea was also commonly recorded causing damage in orchards and vineyards. Armillaria gallica was a common saprophyte or weak pathogen in coniferous and deciduous forests at altitudes from 600 to 1600 m, and less commonly on oaks at lower altitudes. Armillaria ostoyae was rare in central and southern Albania, but common in northern Albania, causing significant damage to pine and other conifers, mostly at altitudes from 600 to 1800 m. Armillaria cepistipes was recorded at altitudes from 800 to 1800 m as a saprophyte or weak pathogen on conifers and deciduous trees, mostly in beech and silver fir forests.Armillaria tabescens was found in oak forests at altitudes from sea level to 900 m. In orchards, A. tabescens occasionally attacked almond and pear trees. Armillaria borealis was found in a few locations in northern Albania, at altitudes from 800 to 1800 m.
... Armillaria root disease has been commonly associated with root decay and mortality of deciduous and coniferous trees and shrubs and has been reported to affect more than 600 species of woody plants ( Garraway et al. 1991, Morrison et al. 1991. This damaging root disease has been found in most forests of the world, but it has been reported to be more common and abundant in temperate and boreal forests than in tropical regions ( Kile et al. 1991). It has been reported to occur in nearly every state of the continental United States and also reported widely in Canada and Mexico (Williams et al. 1986). ...
... Some species of Armillaria reportedly survive as saprophytes in woody substrates within soil or standing dead material, while others have been demonstrated to be aggressive pathogens of living trees. The pathogenic species can kill living roots, eventually killing the infected tree, and then continue to live in dead material as saprophytes for several years ( Kile et al. 1991). Species of Armillaria can be spread by sexually produced basidiospores, or spread vegetatively by rhizomorphs and by transfer from infected roots to healthy roots via contacts and/or grafts ( Kile et al. 1991, Redfern andFilip 1991). ...
Article
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Fungi in the genus Armillaria are associated with an important disease of deciduous and coniferous trees and shrubs in western North America. This study examined the distribution of Armillaria by forest habitat types on the Kaibab National Forest and northern Coconino National Forest, Arizona. Over 400 trees were examined for Armillaria in 76 Interior West Forest Inventory and Analysis permanent plots representing 17 different habitat types. Samples of the fungus associated with Armillaria root disease were collected from 23 trees and identified using DNA sequencing. All samples were determined to consist of a single species, Armillaria solidipes Peck [= A. ostoyae (Romagnesi) Herink]. Only 10 of the 76 plots and 5 of the 17 habitat types sampled had Armillaria solidipes present on one or more trees. A. solidipes was more commonly found in mixed-conifer and subalpine forests and was rare in ponderosa pine (Pinus ponderosa Doug. Ex. Larson & C. Larson) forests, which was consistent with previous root disease studies conducted in the Southwest. However, our estimates of Armillaria presence may be conservative since we did not examine entire root systems.
... When comparing the findings of our study with prior studies, it is necessary to keep in mind that we did not identify the causative agents of decay, nor did we determine the trees infected by the same fungal individual. However, experienced harvester operators recognize decay columns associated with Armillaria species owing to their dark colour and certain other characteristic features and tend to buck the stem base in such cases as the associated stem decay columns are normally below 1 m in height (Kile et al., 1991). ...
Article
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Root and butt rot caused by pathogenic fungi in the genera Heterobasidion and Armillaria is a pressing issue in managed Norway spruce forests. The disease results in financial losses for the forest owners and reduces the volume of wood that can be used in long‐lived products. Pathogenic wood decay fungi spread either with the aid of airborne spores or via mycelial growth among neighbouring trees, the latter leading to clustering (tendency of decayed trees to be in close proximity relative to their neighbouring trees) of decay‐affected trees in forests. Understanding the spatial patterns of the decay‐affected trees at the forest stand level is vital for designing management strategies to address this problem. We examined decay clustering in 273 clear‐cut Norway spruce stands in Norway using harvester‐recorded data on spatial occurrence of decayed and healthy Norway spruce trees. We tested clustering using three global‐cluster tests that account for population density and distribution, evaluating clustering without identifying specific cluster locations. The proportions of clustered and non‐clustered stands differed depending on the statistical test used for clustering assessment, resulting in overall agreement of 32.8% for clustered and 36.9% for non‐clustered. Clustered stands exhibited a median cluster distance (maximum distance between the decay‐affected trees within a cluster) of 12 m (Inter‐Quantile Range, IQR, 6–20 m) and a median of 6 (IQR 3–16) nearest neighbour trees (number of decayed trees forming a cluster), estimates comparable with prior studies focused on assessment of trees infected by mycelial spread of the same fungal individual. The decay incidence in the clustered stands was 16.24%, while the non‐clustered stands had a butt‐rot incidence of 20.97%. In clustered stands the average number of trees per hectare was higher (693) than in non‐clustered stands (553). Synthesis and applications: Our study demonstrates that Norway spruce stands display a diverse range of spatial patterns of butt rotted trees. We found that higher densities of Norway spruce trees probably facilitate the vegetative spread of pathogenic wood decay fungi, leading to clustering of decay‐affected trees. To disrupt the spread of decay fungi between tree generations, precision planting of trees other than Norway spruce around infested stumps of prior generation trees has been recommended by earlier studies. We discussed the potential of using harvester‐derived geoposition data for butt‐rotted trees upon planning and execution of forest regeneration.
... Armillaria root disease is one of the major damaging factors in forests worldwide (Coetzee et al., 2018;Guillaumin et al., 1985;Kile and Watling, 1983;Lockman and Kearns, 2016;Pavlov, 2015;Shaw and Kile, 1991). As a result of root systems colonization by pathogens of the genus Armillaria, a decline in stand productivity has been reported as tree resources are used for defence responses rather than growth (Franceschi et al., 2005) as well as increased tree mortality (Bendz Hellgren and Stenlid, 1995;Bloomberg and Morrison, 1989;Cruickshank, 2011;Cruickshank et al., 2011;Filip et al., 2010;Kile et al., 1991;Shaw and Toes, 1977). Economic losses occur, reaching a value of up to 1000 Euros/ha/y in the most heavily infested stands (Kaliszewski et al., 2007;Smith, 1984). ...
Article
Full-text available
Armillaria root disease affects forests around the world. It occurs in many habitats and causes losses in the infested stands. Weather conditions are important factors for growth and development of Armillaria species. Yet, the relation between occurrence of damage caused by Armillaria disease and weather variables are still poorly understood. Thus, we used generalized linear mixed models to determine the relationship between weather conditions of current and previous year (temperature, precipitation and their deviation from long-term averages , air humidity and soil temperature) and the incidence of Armillaria-induced damage in young (up to 20 years old) and older (over 20 years old) coniferous stands in selected forest districts across Poland. We used unique data, gathered over the course of 23 years (1987-2009) on tree damage incidence from Armillaria root disease and meteorological parameters from the 24-year period (1986-2009) to reflect the dynamics of damage occurrence and weather conditions. Weather parameters were better predictors of damage caused by Armillaria disease in younger stands than in older ones. The strongest predictor was soil temperature, especially that of the previous year growing season and the current year spring. We found that temperature and precipitation of different seasons in previous year had more pronounced effect on the young stand area affected by Armillaria. Each stand's age class was characterized by a different set of meteorological parameters that explained the area of disease occurrence. Moreover, forest district was included in all models and thus, was an important variable in explaining the stand area affected by Armillaria.
... The genus Armillaria includes species that are among the most common and important causes of root rot in woody plants, worldwide Fox, 2000). However, although most Armillaria species have the potential to infect healthy and stressed trees, they differ in their pathogenicity according to the host and may also act as necrotrophs and saprobes (Hood et al., 1991;Kile et al., 1991). To date, native species in Patagonia were known to act as saprophytes in native forest and to be able to become aggressive pathogens of exotic tree species (Pildain et al., 2009). ...
Article
Temperate forests ecosystems are threatened by declines and diseases worldwide. Causes have been investigated during the last decades allowing, in some cases, to relate such deterioration to climate change or pathogens but, in many cases, causes have not yet been elucidated. In recent decades, a phenomenon of grouped mortality, whose etiology remains unknown, has been observed in the two most distributed Nothofagus species of North Patagonian forests. This study aimed to assess sapwood-inhabiting fungal diversity of N. dombeyi and N. pumilio trees in healthy and affected stands in order to determine whether health condition shape these fungal communities, to characterize such patterns and, to seek for the likely pathogens associated with tree damage. Seven sites in Los Alerces National Park (Chubut, Argentina), were sampled seasonally for two years. Eighty-eight fungal taxa were recovered and identified. Spatial heterogeneity across plant compartments was found, involving community composition and structure, with stem harbouring greater diversity than root. It was found that the sapwood in both Nothofagus species was inhabited by different fungal species, in high richness and showing different patterns according to health condition and plant compartment. Health condition was a stronger driver of N. dombeyi sapwood-inhabiting fungal community than it was for N. pumilio; the latter evidenced plant compartment as a stronger driver of such community. Nothofagus pumilio presented higher frequency of decomposition agents in both affected and healthy trees than it was in N. dombeyi, indicating a greater wood deterioration of its stands. More decomposition agents and potential pathogens have been found in symptomatic stands of N. dombeyi than in healthy stands, although their frequency patterns do not allow the inference of conclusions about the primary fungal agent causing tree mortality. Our findings suggest a secondary role of living-wood-inhabiting mycobiota in tree damage processes as an expression of tree stress caused by climatic factors.
... Armillaria species occur all across Europe [85,121,[123][124][125][126][127][128][129]. The most common Armillaria species on pines in north-east Europe are A. borealis and A. ostoyae [130,131], while in Maritime and south-east Europe, A. cepistipes, A. gallica, A. mellea, and A. ostoyae are the most common [121,122,129,132,133]. Armillaria species can be aggressive primary pathogens as well as secondary pathogens of stressed trees and saprophytes [134]. ...
Article
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Pines are major components of native forests and plantations in Europe, where they have both economic significance and an important ecological role. Diseases of pines are mainly caused by fungal and oomycete pathogens, and can significantly reduce the survival, vigor, and yield of both individual trees and entire stands or plantations. Pine pitch canker (PPC), caused by Fusarium circinatum (Nirenberg and O’Donnell), is among the most devastating pine diseases in the world, and is an example of an emergent invasive disease in Europe. The effects of microbial interactions on plant health, as well as the possible roles plant microbiomes may have in disease expression, have been the focus of several recent studies. Here, we describe the possible effects of co-infection with pathogenic fungi and oomycetes with F. circinatum on the health of pine seedlings and mature plants, in an attempt to expand our understanding of the role that biotic interactions may play in the future of PPC disease in European nurseries and forests. The available information on pine pathogens that are able to co-occur with F. circinatum in Europe is here reviewed and interpreted to theoretically predict the effects of such co-occurrences on pine survival, growth, and yield. Beside the awareness that F. circinatum may co-occurr on pines with other pathogens, an additional outcome from this review is an updating of the literature, including the so-called grey literature, to document the geographical distribution of the relevant pathogens and to facilitate differential diagnoses, particularly in nurseries, where some of them may cause symptoms similar to those induced by F. circinatum. An early and accurate diagnosis of F. circinatum, a pathogen that has been recently introduced and that is currently regulated in Europe, is essential to prevent its introduction and spread in plantings and forests.
... At least 1.3 million hectares in the Northern Rockies have moderate to severe root disease, with up to 60% caused by Armillaria ostoyae (USFS 2007). Armillaria kills conifers of all species when they are young, but it is especially damaging to Douglas-fir, subalpine fir, and grand fir (Abies grandis) because these species remain susceptible throughout their lives (Kile et al. 1991). Armillaria and other root diseases influence forest species composition, structure, and successional trajectories by accelerating a transition to species that are more tolerant of root disease or by maintaining stands of more susceptible species in early seral stages (Byler and Hagle 2000). ...
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Disturbances alter ecosystem, community, or population structures and change elements of the biological and/or physical environment. Climate changes can alter the timing, magnitude, frequency, and duration of disturbance events, as well as the interactions of disturbances on a landscape, and climate change may already be affecting disturbance events and regimes. Interactions among disturbance regimes, such as the co-occurrence in space and time of bark beetle outbreaks and wildfires, can result in highly visible, rapidly occurring, and persistent changes in landscape composition and structure. Understanding how altered disturbance patterns and multiple disturbance interactions might result in novel and emergent landscape behaviors is critical for addressing climate change impacts and for designing land management strategies that are appropriate for future climates. This chapter describes the ecology of important disturbance regimes in the Northern Rockies region, and potential shifts in these regimes as a consequence of observed and projected climate change. We summarize five disturbance types present in the Northern Rockies that are sensitive to a changing climate—wildfires, bark beetles, white pine blister rust (Cronartium ribicola), other forest diseases, and nonnative plant invasions—and provide information that can help managers anticipate how, when, where, and why climate changes may alter the characteristics of disturbance regimes.
... Gr1886/13, 24-IV-13, Gr2164/13, 25-IV-13. Obs.: Especie de interés para el sector forestal, ya que existen reportes de distintas especies de Armillaria que producen pudrición de raíces en árboles en pie (Kile et al. 1991). no comestible, ingerida cruda o mal cocida puede causar trastornos estomacales y hemólisis (Gamundí & Horak, 1994 : Singer, 1952;Singer, 1954;Horak, 1964;Lazo, 1971;Horak, 1980;Godeas et al., 1993;Valenzuela, 1993;Valenzuela et al., 1998 Ref. : cotton, 1915: cotton, , Horak, 1980Pegler et al., 1980;Horak, 1982;Godeas et al., 1993;Valenzuela, 1993;Valenzuela et al., 1998 Ref. : Singer, 1953;Singer, 1954;Singer, 1969;Horak, 1980; M.E.: ArGEntInA, cHu, dpto. ...
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Se han realizado abundantes expediciones micológicas en Patagonia, pero los sitios relevados han cubierto solo una porción minima de esta extensa región. Así, las expediciones micológicas se centraron en los extremos norte (Neuquén y Rio Negro en Argentina, Santiago y Maule en Chile) y sur (Tierra del Fuego argentina y chilena) de los bosques subantárticos. El territorio comprendido entre estas provincias permanence inexplorado para los hongos agaricoides. El objetivo central de este trabajo fue evaluar la riqueza de especies de hongos agaricoides en bosques puros de Nothofagus pumilio (lenga) en Chubut (Argentina). Se realizaron 4 viajes de muestreo entre 2012 y 2014 en 3 bosques en la provincia del Chubut, Argentina. Se encontraron 130 especies de hongos agaricoides. Entre ellas, 125 pertenecen al orden Agaricales, 2 al orden Boletales y 3 al orden Russulales. La familia Cortinariaceae (Agaricales) fue la más rica, con 57 especies. Se presenta una clave para todas las especies halladas. Fueron encontrados 4 nuevos registros a nivel nacional y 124 a nivel provincial.
... Besides being pathogens, Armillaria species also play a critical role as decomposers. Extensive infection observed in stumps and roots and long possession of the substrate suggest that Armillaria species contribute significantly to decomposition and mineral cycling within many forests [6]. ...
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Background Armillaria is a globally distributed mushroom-forming genus composed primarily of plant pathogens. Species in this genus are prolific producers of rhizomorphs, or vegetative structures, which, when found, are often associated with infection. Because of their importance as plant pathogens, understanding the evolutionary origins of this genus and how it gained a worldwide distribution is of interest. The first gasteroid fungus with close affinities to Armillaria—Guyanagaster necrorhizus—was described from the Neotropical rainforests of Guyana. In this study, we conducted phylogenetic analyses to fully resolve the relationship of G. necrorhizus with Armillaria. Data sets containing Guyanagaster from two collecting localities, along with a global sampling of 21 Armillaria species—including newly collected specimens from Guyana and Africa—at six loci (28S, EF1α, RPB2, TUB, actin-1 and gpd) were used. Three loci—28S, EF1α and RPB2—were analyzed in a partitioned nucleotide data set to infer divergence dates and ancestral range estimations for well-supported, monophyletic lineages. Results The six-locus phylogenetic analysis resolves Guyanagaster as the earliest diverging lineage in the armillarioid clade. The next lineage to diverge is that composed of species in Armillaria subgenus Desarmillaria. This subgenus is elevated to genus level to accommodate the exannulate mushroom-forming armillarioid species. The final lineage to diverge is that composed of annulate mushroom-forming armillarioid species, in what is now Armillaria sensu stricto. The molecular clock analysis and ancestral range estimation suggest the most recent common ancestor to the armillarioid lineage arose 51 million years ago in Eurasia. A new species, Guyanagaster lucianii sp. nov. from Guyana, is described. Conclusions The armillarioid lineage evolved in Eurasia during the height of tropical rainforest expansion about 51 million years ago, a time marked by a warm and wet global climate. Species of Guyanagaster and Desarmillaria represent extant taxa of these early diverging lineages. Desarmillaria represents an armillarioid lineage that was likely much more widespread in the past. Guyanagaster likely evolved from a gilled mushroom ancestor and could represent a highly specialized endemic in the Guiana Shield. Armillaria species represent those that evolved after the shift in climate from warm and tropical to cool and arid during the late Eocene. No species in either Desarmillaria or Guyanagaster are known to produce melanized rhizomorphs in nature, whereas almost all Armillaria species are known to produce them. The production of rhizomorphs is an adaptation to harsh environments, and could be a driver of diversification in Armillaria by conferring a competitive advantage to the species that produce them. Electronic supplementary material The online version of this article (doi:10.1186/s12862-017-0877-3) contains supplementary material, which is available to authorized users.
... An example is the association between the mountain pine beetle (Dendroctonus ponderosae) and the blue-stain fungus Grosmannia clavigera causing the decline of lodgepole pines in North America, where the fungus attacks the living cell network, gradually spreading throughout the sapwood and causing the death of the tree through starvation (Kirisits, 2007). However, disease can be caused directly by pathogens on trees that are stressed, which may turn latent fungi pathogenic (Griffith and Boddy, 1990;Desprez-Loustau et al., 2006;Parfitt et al., 2010), or on otherwise healthy trees where conditions favor the pathogen, such as is the case with the opportunistic fungus Armillaria (Kile et al., 1991). ...
Article
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This review examines the roles that ray and axial parenchyma (RAP) plays against fungal pathogens in the secondary xylem of wood within the context of the CODIT model (Compartmentalization of Decay in Trees), a defense concept first conceived in the early 1970s by Alex Shigo. This model, simplistic in its design, shows how a large woody perennial is highly compartmented. Anatomical divisions in place at the time of infection or damage, (physical defense) alongside the ‘active’ response by the RAP during and after wounding work together in forming boundaries that function to restrict air or decay spread. The living parenchyma cells play a critical role in all of the four walls (differing anatomical constructs) that the model comprises. To understand how living cells in each of the walls of CODIT cooperate, we must have a clear vision of their complex interconnectivity from a three-dimensional perspective, along with knowledge of the huge variation in ray parenchyma (RP) and axial parenchyma (AP) abundance and patterns. Crucial patterns for defense encompass the symplastic continuum between both RP and AP and the dead tissues, with the latter including the vessel elements, libriform fibers, and imperforate tracheary elements (i.e., vasicentric and vascular tracheids). Also, the heartwood, a chemically altered antimicrobial non-living substance that forms the core of many trees, provides an integral part of the defense system. In the heartwood, dead RAP can play an important role in defense, depending on the genetic constitution of the species. Considering the array of functions that RAP are associated with, from capacitance, through to storage, and long-distance water transport, deciding how their role in defense fits into this suite of functions is a challenge for plant scientists, and likely depends on a range of factors. Here, we explore the important role of RAP in defense against fungal pathogens and the trade-offs involved from a viewpoint for structure-function relations, while also examining how fungi can breach the defense system using an array of enzymes in conjunction with the physically intrusive hyphae.
... and Heterobasidion spp., are predicted to become more severe and move into new geographic regions during drought because these pathogens most successfully colonize stressed trees (Sturrock et al., 2011;Olatinwo et al., 2013). Armillaria root rot is caused by primary and secondary pathogens that infect conifers and hardwoods (Kile et al., 1991). Infections cause wood decay, growth reduction, and tree mortality, and increase tree susceptibility to colonization by insects (Sturrock et al., 2011). ...
... Armillaria root rot belongs to the most dangerous diseases of tree root systems in the moderate and subtropical climatic zones [Kile et al. 1991, Mańka 2005, Rykowski 1990]. So far, in the subject-related world literature, over 40 species of Armillaria have been described [Żółciak 2005]. ...
Article
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The objective of the presented st udies was the determination of Armillaria spe- cies causing root rot in stands growing in habi tats of a higher fertility in the Siemianice Experimental Forest District. With the application of the mycelium intersterility test, the occurrence of A. gallica and A. ostoyae was found on the area of the mentioned Forest District, while in the nearby situated Mansion Park, Armillaria cepistipes was identified
... As soon as the mycelial fans reach the root collar and have colonized the entire circumference, the host will die within a short time ( ). In cases where the host is able to restrict the Armillaria infection to single roots or only parts of the root system, Armillaria commonly advances to the heartwood and causes butt rot, which might reduce the productivity and stability of the host tree ( Kile et al. 1991). To date, the control of Armillaria root disease is challenging due to its hidden growth in the soil or infected plant tissue and its persistence in woody debris in soils for many years ( Baumgartner et al. 2011). ...
Article
Mycelial fan formation was studied in five Armillaria cepistipes, ten A. borealis and ten diploid and six haploid A. ostoyae strains on excised stem segments of Picea abies. Stem segments were either non-autoclaved or autoclaved, representing dying and dead wood, respectively. To confirm the identity of mycelial fans on non-autoclaved stem segments, re-isolations were made and isolates characterized with microsatellite markers. Mycelial fan formation on autoclaved stem segments was fast and reliable for most of the tested Armillaria strains. On non-autoclaved stem segments, mycelial fan formation was slower, more erratic and less predictable. Mycelial fan formation was fastest in A. cepistipes closely followed by A. borealis and was slowest in A. ostoyae. For two A. cepistipes and four A. ostoyae strains (all diploid), growth rates of mycelial fans were estimated in a time course experiment. They ranged between 5.1 and 8.7 mm/day for autoclaved and between 1.4 and 4.7 mm/day for non-autoclaved stem segments. The haploid A. ostoyae strains also formed mycelial fans on autoclaved stem segments, but typically slower and less reliably than the diploid strains. Whether haploid strains are able to produce mycelial fans on non-autoclaved stem segments remains unknown because of accidental diploidization of the original haploid strains which was likely caused by basidiospores introduced into the study system on the non-autoclaved stems. Overall, the method developed in this study may be useful for further investigations into the genetic, physiological and biochemical nature of mycelial fan formation in the genus Armillaria.
... The establishment and maintenance of a diversity of tree species in forests and urban plantings can help Incidence of root disease and butt rot caused by A. ostoyae is likely to increase as temperatures increase and precipitation decreases, resulting in increased direct and indirect tree mortality and larger impacts on tree growth. Kile et al., 1991;Cruickshank et al., 1997;Goheen and Otrosina, 1998;Morrison et al., 2000;La Porta et al., 2008;Dukes et al., 2009;Kliejunas, 2011 Phytophthora root rot ~ Phytophthora cinnamomi ...
Article
The health of the earth's forests and urban green spaces is increasingly challenged by the outcomes of human activities, including global climate change. As climate changes, the role and impact of diseases on trees in both forest ecosystems and in urban settings will also change. Knowledge of relationships between climate variables and diseases affecting forest and urban trees is reviewed, with specific emphasis on those affecting foliage, shoots, and stems. Evidence that forest diseases are already responding to the earth's changing climate is examined (e.g., Dothistroma needle blight in northern British Columbia) as are predicted scenarios for future changes in impact on forests by other tree diseases. Outbreaks of tree diseases caused by native and alien pathogens are predicted to become more frequent and intense - this and other general predictions about the effects of climate change on forest and tree diseases are discussed. Despite the uncertainty that accompanies such predictions it is imperative that researchers, forest and urban tree managers, and policy makers work together to develop and implement management strategies that enhance the resilience of the worlds' forests and urbanized trees. Strategies discussed include monitoring, forecasting, planning, and mitigation.
... Some Armillaria species are primary pathogens, causing the disease generally referred to as Armillaria root rot, which is considered amongst the most serious diseases of trees in boreal and temperate forests and various species damage high-value crops. Other species are important components of woody ecosystems by virtue of their saprophytic life strategy, where they contribute significantly to wood degradation (Gregory et al. 1991;Kile et al. 1991). Armillaria species also have an important role in the traditions of various Asian cultures as a source of nutrients or linked to traditional medicine (Hobbs 1986). ...
Article
Fourteen Chinese Biological Species (CBS) of Armillaria were previously identified in a collection of Chinese isolates. CBS C, F, G, H, J, L, N and O remained unnamed, while the remaining isolates included A. borealis, A. cepistipes, A. gallica, A. mellea, A. sinapina and A. tabescens. CBS F was suggested to represent A. singula based on basidiocarp morphology. In this study, phylogenetic relationships between Chinese Armillaria isolates and those from other parts of the world were determined based on DNA sequence data. Results of this study suggest that CBS F might not represent A. singula, and that A. monadelpha (a name applied to the North American form of A. tabescens by some authors) and A. tabescens should be treated as a single species. Four main phylogenetic lineages, referred to as the A. ostoyae, A. gallica, A. tabescens and A. mellea clusters, were identified on the phylogenetic trees. The unnamed biological species grouped within the “A. gallica cluster” and were phylogenetically closely related. The results of this study contribute to our current understanding of the systematics of Armillaria from South East Asia where these fungi are relatively poorly known.
... It would be very interesting to find out if A. mellea is really so rare in Polish forests . The natural range of the species covers the whole of Poland (Kile et al. 1991) and A. mellea could be expected to occur in many deciduous tree stands. On the basis of the author's permanent observations this Armillaria species produces basidiomes rather seldom. ...
Article
Armillaria root rot is the most important disease of woody plants in forest, hor-ticulture plantations, and in house gardens all over the world. Armillaria spp. can infect more than 600 species of trees, shrubs and herbaceous plants (Hood et al. 1991, Kile et al. 1991, Mańka 2005). The Armillaria genus includes more than 40 species. In Europe seven species were noted and in Poland six (Domański et al. 1967, Pegler 2000, Żółciak 2005). In Polish forests Armillaria ostoyae is the most common species occurring in al-most all forest sites. Armillaria borealis and A. cepistipes have been noted on conifer-ous and deciduous trees or shrubs. Armillaria gallica is most common species in oak stands and plays an important role in the complex of factors causing oak decline in many oak stands. Armillaria mellea was found only in one forest ecosystem (oak stand, Zielonka Forest District; Łakomy 2001) and in three other places – vineyard and two small groups of poplars out of forest (Żółciak 1999 a, b). The sixth species – A. tabescens was described in 1967 (Domański et al. 1967) from beech stumps in two forest districts in south-eastern Poland (Bieszczady Mountains, Stuposiany and Wetlina Forest Districts). Four Armillaria species (excluding A. tabescens and A. mellea) were noted by many authors (Rykowski 1990, Żółciak 1991, Mańka and Łakomy 1995, Kowalski et al. 1997, Łakomy 1998, Żółciak 1999 a, b, Łakomy and Siwecki 2000, Szewczyk and Mańka 2002, Łakomy 2004). In 2006 two groups of A. mellea basidiomes were found in two localities in Strzelce Krajeńskie Forest District (compartment 177c and on the border between compartments 177c and 177f). One group was growing in the middle of a forest path (2 m from an oak stump), which was the border between a mixed stand (Larix decidua, Pinus sylvestris 55-year-old, Quercus petraea, Fagus sylvatica, Carpinus betulus 62-year-old, 177f) and a 64-year-old Scots pine stand with single birch (Betula pendula) and oak trees (177c, Phot. 1). In the second locality basidiomes appeared close to a birch stump in a Scots pine stand (177c, Phot. 2). These two localities were less than 100 m distant from each other. In the pine stand basidiomes were found close to a neighbouring oak stand (about 50 m). The two groups of basidiomes differed from each other. Pilei of basidiomes from the path were bigger than those found on stump, and open. The margin of basidiomes growing close to the stump was rolled down.
... Known also as the shoe-string fungus, honey mushroom, or oak root rot fungus, Armillaria spp. attack a wide range of woody plant species including perennials, commercial trees, forest trees, and shrubs (Hood et al., 1991;Kile et al., 1991). Some specific plants that Armillaria spp. ...
Article
The genus Armillaria contains white-rot basidiomycetes pathogenic to woody plant hosts worldwide. Of particular interest are the species found in southeastern peach (Prunus persica) orchards causing Armillaria root rot. Isolates collected during the past four years from orchards in Georgia, South Carolina, and Alabama were identified to species and grouped within species based on molecular analysis of the internal transcribed spacer (ITS) regions and the intergenic spacer (IGS) regions of the ribosomal DNA (rDNA). Thirty isolates of A. tabescens and nine isolates of A. mellea were identified from these orchards. Restriction fragment length polymorphism (RFLP) analysis of the IGS region with Alu I revealed two groups of A. tabescens and one group of A. mellea. Similarly, the ITS region was amplified with primer sets At-ITS1/Am-ITS1/ITS-2 and ITS-1/ITS-4 and subsequently restricted with Mbo II and Hha I. These restrictions indicated two groups of A. tabescens and four groups of A. mellea.
... Because of their ubiquity and wide host range, pathogenic Armillaria spp. are of considerable importance to the ecology and management of natural and planted forests in boreal, temperate and tropical zones (Kile et al. 1991). ...
Article
The overall aim of this study was to develop a new, reliable and rapid diagnostic assay for differentiating six European Armillaria species based on variation in their elongation factor‐1 alpha (EF‐1 α) gene sequences and to verify a set of species‐specific primers on 61 Armillaria isolates from Europe. Partial sequences of the EF‐1 α gene obtained in Armillaria borealis, Armillaria cepistipes, Armillaria gallica, Armillaria mellea, Armillaria ostoyae and Armillaria tabescens revealed sufficient interspecific variation to distinguish among species using nested primers. These primers gave unambiguous bands when tested on representative isolates of five of these species. However, the EF‐1 α sequences of European A. borealis isolates clustered into two distinct clades, termed here AbX and AbY. Specific primers were subsequently designed and tested successfully on both AbX‐type and AbY‐type A. borealis isolates. The taxonomy of A. borealis needs to be elucidated to determine whether a new, as yet unnamed Armillaria taxon exists in Europe. Three A. borealis isolates were also found to have heterozygous sites in their EF‐1 α sequences, which suggests that the gene could exist in more than one copy or that these isolates contain hybrid sequences. A pyrosequencing method was also developed, targeting a small region of EF‐1 α intron 4, which was able to differentiate European Armillaria isolates to the species level and additionally could distinguish AbX‐type and AbY‐type A. borealis isolates.
... In natural stands infested with ARD, dominant healthy trees limit infection (Kile et al., 1991) but mortality is occasionally encountered in overmature veterans. In regrowth Eucalypt stands, A. luteobubalina is capable of infecting and killing dominant trees (Edgar et al., 1976; Shearer and Tippett, 1988). ...
Article
The incidence of Armillaria root disease was recorded during routine measurement of a silvicultural experiment designed to test the effect of thinning and nitrogen fertilizer application on the growth of karri regenerated after clearfelling. The experiment was established in 1984, when the stand was 12-year-old. Ten years after thinning, the level of disease increased significantly with increased thinning intensity, and disease accounted for 51% of the mortality in the plots thinned to 200 stems/ha. Fifteen years after thinning, the level of infection had increased in all treatments but was still significantly lower in the unthinned treatment. In the thinned treatments, 54–63% of the trees ranked in the largest 200 stems/ha were infected, and 50–100% of the mortality within these trees was attributed to Armillaria luteobubalina. In the unthinned treatment, no mortality within the dominant trees was associated with disease. Ten years after thinning and fertilizer treatments, it could not be determined whether fertilizer application had had any effect on the level of disease. Whole tree thinning, which results in stump and root removal, is discussed as a viable management option in high-quality karri regrowth stands infested with A. luteobubalina.
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The occurrence of Armillaria mellea (Vahl) P. Kumm. and the ecological characteristics of this fungus were studied in Kocaeli, Yuvacik dam basin mixed-broad leaved forests. During the surveys, we analyzed the sporocarps (fruiting bodies) of A. mellea growing up on woody plants in plots selected by cluster sampling in the Yuvacik dam watershed dominated by broad-leaved forests. The Runs test results showed that randomness rules complied in the selection of the plots, and there was no tendency (p= 0.109 > 0.05, z= -1.603). The presence/absence of A. mellea and environmental variables were tested with Chi-square analysis, and the temperature differed among these environmental variables. To the dendrogram, A. mellea was mainly seen in the south of the study area and preferred western aspects. It is understood that this macrofungus prefers the south of the study area because of the altitude. Our data showed that sporocarps of A. mellea generally occurred in the western aspect, at temperatures of 15–20°C, >80% humidity and 800–1000 m altitude. Our logistic regression analysis model (z=-9.508+0.307×temperature+0.081×humidity) showed that if the temperature and humidity change by 1 unit in the region, sporocarp formation is affected by 36% and 8.4%, respectively.
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This datasheet on Armillaria novae-zelandiae covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Impacts, Uses, Prevention/Control, Further Information.
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This datasheet on Armillaria limonea covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Impacts, Uses, Prevention/Control, Further Information.
Thesis
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Armillaria es un género de hongos de la pudrición blanca y constituye uno de los patógenos más ampliamente estudiados por las pérdidas que ocasiona en la actividad forestal. En los últimos años se ha comenzado a utilizar la batería enzimática ligninolítica de este organismo en actividades de biorremediación que involucran la producción de enzimas extracelulares y la transformación de colorantes. En Argentina este patógeno está asociado a bosque nativo e implantado de Patagonia y recientemente han sido dilucidadas su taxonomía y filogenia estableciéndose cuatro linajes correspondientes a cuatro especies. El objetivo de este trabajo fue caracterizar la enzimología ligninolítica de especies de Armillaria en Patagonia y su capacidad de biodegradar compuestos contaminantes. Se cuantificaron las actividades lacasa y manganeso peroxidasa basales de 17 aislamientos y se evaluó su inducción por aditivos xenobióticos (guayacol y cobre) en el cultivo en 6 aislamientos representativos de las cuatro especies. Se determinaron los perfiles electroforéticos de isoenzimas de lacasa de 12 aislamientos y de tres aislamientos cultivados con los distintos xenobióticos. Se evaluó la capacidad de decoloración del colorante trifenilmetano verde de malaquita. La actividad lacasa, su perfil de isoenzimas, la susceptibilidad de ser inducida y la capacidad degradadora del colorante sugirieron ser caracteres quimiotaxonómicos útiles para Armillaria en Patagonia, no así la actividad manganeso peroxidasa que no resultó de interés a estos fines y registró actividades muy bajas o indetectables en concordancia con otras especies de la pudrición blanca citadas en bibliografía. Todas las especies demostraron ser efectivas en degradar el colorante verde de malaquita pero A. montagnei y A. novae-zelandiae exhibieron las mayores capacidades degradadoras, asociadas a las mayores actividades lacasa. En A. umbrinobrunnea y A. sparrei la degradación no estuvo asociada a actividades lacasa significativas, por lo que otras ligninasas u otros procesos oxidativos podrían ser los responsables de la transformación del contaminante. A. novae-zelandiae presentó la mayor tendencia a inducción de la enzima lacasa, resultando condicionante la presencia de aditivos xenobióticos para el registro de actividades enzimáticas elevadas en condiciones de cultivo; la inducción evidenció ser diferencial para algunas isoformas de lacasa en esta especie. El perfil electroforético reveló tres isoenzimas de lacasa, dos de ellas nuevas para el género, y la heterogeneidad interespecífica suficiente para resultar útil en caracterizar las especies patagónicas; ambos elementos resultan novedosos en el marco actual de conocimiento del género.
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Armillaria is a globally distributed fungal genus most notably composed of economically important plant pathogens that are found predominantly in forest and agronomic systems. The genus sensu lato has more recently received attention for its role in woody plant decomposition and in mycorrhizal symbiosis with specific plants. Previous phylogenetic analyses suggest that around 50 species are recognized globally. Despite this previous work, no studies have analyzed the global species richness and distribution of the genus using data derived from fungal community sequencing datasets or barcoding initiatives. To assess the global diversity and species richness of Armillaria, we mined publicly available sequencing datasets derived from numerous primer regions for the ribosomal operon, as well as ITS sequences deposited on Genbank, and clustered them akin to metabarcoding studies. Our estimates reveal that species richness ranges from 50 to 60 species, depending on whether the ITS1 or ITS2 marker is used. Eastern Asia represents the biogeographic region with the highest species richness. We also assess the overlap of species across geographic regions and propose some hypotheses regarding the drivers of variability in species diversity and richness between different biogeographic regions.
Technical Report
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Forests in the Southern United States experience a wide variety of weather-related disturbances, from small-scale events which have management implications for one or a few landowners to major hurricanes impacting many ownerships across multiple States. The immediate impacts of catastrophic weather disturbance are obvious-trees are killed, stressed, or damaged due to wind, flooding, ice, hail, or some combination of events. How forests respond to disturbance depends on several factors such as forest types and attributes, ecoregion, local pressure from invasive plants, preexisting infestations of pests and pathogens, prior disturbance events, and other variables which interact in complex ways, influencing successional dynamics and management decisions. In this review, we synthesize the major weather perturbations affecting the forests of the Southern United States and current state of the knowledge surrounding interactions between these events, forest pests, and forest diseases. We present a compilation of non-quantitative observations between 1955 and 2018 from annual U.S. Department of Agriculture Forest Service "Major Forest Insect and Disease Conditions in the United States" reports describing where insects or diseases were found on trees that were stressed by weather disturbances. Two conceptual models are presented, one describing changes in forest structure and composition, and a generalized model of herbivorous pest population fluctuations following different severity levels of disturbance. Finally, we propose 11 questions that require additional research to better inform sustainable forest management decisions in preparation for and in response to catastrophic weather events.
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Armillaria root rot is a serious disease, chiefly of woody plants, caused by many species of Armillaria that occur in temperate, tropical and subtropical regions of the world. Very little is known about Armillaria in South America and Southeast Asia, although Armillaria root rot is well known in these areas. In this study, we consider previously unidentified isolates collected from trees with symptoms of Armillaria root rot in Chile, Indonesia and Malaysia. In addition, isolates from basidiocarps resembling A. novae-zelandiae and A. limonea, originating from Chile and Argentina, respectively, were included in this study because their true identity has been uncertain. All isolates in this study were compared, based on their similarity in ITS sequences with previously sequenced Armillaria species, and their phylogenetic relationship with species from the Southern Hemisphere was considered. ITS sequence data for Armillaria also were compared with those available at GenBank. Parsimony and distance analyses were conducted to determine the phylogenetic relationships between the unknown isolates and the species that showed high ITS sequence similarity. In addition, IGS-1 sequence data were obtained for some of the species to validate the trees obtained from the ITS data set. Results of this study showed that the ITS sequences of the isolates obtained from basidiocarps resembling A. novae-zelandiae are most similar to those for this species. ITS sequences for isolates from Indonesia and Malaysia had the highest similarity to A. novae-zelandiae but were phylogenetically separated from this species. Isolates from Chile, for which basidiocarps were not found, were similar in their ITS and IGS-1 sequences to the isolate from Argentina that resembled A. limonea. These isolates, however, had the highest ITS and IGS-1 sequence similarity to authentic isolates of A. luteobubalina and were phylogenetically more closely related to this species than to A. limonea.
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Although cells of mushroom-producing fungi typically contain paired haploid nuclei (n + n), most Armillaria gallica vegetative cells are uninucleate. As vegetative nuclei are produced by fusions of paired haploid nuclei, they are thought to be diploid (2n). Here we report finding haploid vegetative nuclei in A. gallica at multiple sites in southeastern Massachusetts, USA. Sequencing multiple clones of a single-copy gene isolated from single hyphal filaments revealed nuclear heterogeneity both among and within hyphae. Cytoplasmic bridges connected hyphae in field-collected and cultured samples, and we propose nuclear migration through bridges maintains this nuclear heterogeneity. Growth studies demonstrate among- and within-hypha phenotypic variation for growth in response to gallic acid, a plant-produced antifungal compound. The existence of both genetic and phenotypic variation within vegetative hyphae suggests that fungal individuals have the potential to evolve within a single generation in response to environmental variation over time and space.
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This is a continuation of a series focused on providing a stable platform for the taxonomy of phytopathogenic fungi and fungus-like organisms. This paper focuses on one family: Erysiphaceae and 24 phytopathogenic genera: Armillaria, Barriopsis, Cercospora, Cladosporium, Clinoconidium, Colletotrichum, Cylindrocladiella, Dothidotthia,, Fomitopsis, Ganoderma, Golovinomyces, Heterobasidium, Meliola, Mucor, Neoerysiphe, Nothophoma, Phellinus, Phytophthora, Pseudoseptoria, Pythium, Rhizopus, Stemphylium, Thyrostroma and Wojnowiciella. Each genus is provided with a taxonomic background, distribution, hosts, disease symptoms, and updated backbone trees. Species confirmed with pathogenicity studies are denoted when data are available. Six of the genera are updated from previous entries as many new species have been described.
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Armillaria mellea and A. solidipes are two of the primary causal agents of Armillaria root rot (ARR) in over 500 woody hosts. Although extensive research on this disease has been conducted over the past few decades, finding controls for ARR has been challenging. This is largely due to the lack of knowledge of the fungal infection and host response processes. It is important to understand the infection process in order to develop new strategies to control the disease. In this review, the initial steps of the infection process of A. mellea and A. solidipes are discussed, with focus on fungal attachment and penetration. A mucilaginous substance produced at the rhizomorph tip is thought to aid attachment of the pathogen to host. Toxins and cell wall degrading enzymes produced by the fungus together with mechanical pressure generated by the rhizomorph is speculated to support fungal penetration. However, a detailed description of the Armillaria spp. infection process is lacking in the literature, and this illustrates the need for further studies.
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Diploid isolates of Armillaria were collected from a 50 × 25 m plot at each of two sites in the White Mountains, New Hampshire. All living and dead trees were examined for rhizomorphs, mycelial fans and decay. Isolates were identified to species using diploid-haploid pairings, and clones were delineated using somatic incompatibility and six isozyme markers. At a predominantly spruce-fir site, only A. ostoyae was identified and evidence suggested that at least 25 trees (10%) were pathogenically colonized. Isolates from 39 trees were divided into six somatic incompatibility groups; the largest somatic incompatibility group had colonized at least 17 trees. At a mixed conifer-hardwood site, three species of Armillaria were identified. Six somatic incompatibility groups of A. ostoyae were identified among isolates from 32 of 281 trees; the largest somatic incompatibility group colonized 14 trees and was at least 25 m in diameter. At least 14 of the 32 trees from which A. ostoyae was identified were pathogenically colonized. Armillaria gemina (31 trees) and A. calvescens (29 trees) were isolated mostly from epiphytic rhizomorphs on hardwoods, were apparently pathogenic on four of the trees, and were each divided into two somatic incompatibility groups. Two juxtaposed somatic incompatibility groups of A. ostoyae had identical isozyme electromorphs, but each of the other somatic incompatibility groups had an unique combination of electromorphs. The distribution pattern of the somatic incompatibility groups and correspondence with isozyme electromorphs supported the hypothesis that the somatic incompatibility groups represent asexually spreading clones. At both sites, there was little intermingling of clones of the same species; however, at the mixed conifer-hardwood site clones of the different Armillaria species overlapped considerably.
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Culturable fungi from 28 fungal communities were isolated from soil, rhizosphere and thick (1 cm diam.) roots of living beech (Fagus sylvatica) trees and their stumps 1–3 years after felling. All fungi were morphotyped and identified morphologically. The frequency of fungi was 2–5× greater in stumps than in living trees. The diversity of fungi was similar in living trees and stumps. The majority of fungal species that occurred at greater frequency on/in roots of stumps reduced the growth of Armillaria ostoyae and to a smaller extent of A. gallica rhizomorphs in a soil substrate in vitro. It is suggested that the mycobiota of roots may constrain the colonization of F. sylvatica by A. ostoyae rather than by A. gallica. The significance of these findings in the epidemiology of Armillaria in beech forests is discussed.
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Honey fungus (Armillaria spp.) root rot is the disease most frequently inquired about by U.K. gardeners to the Royal Horticultural Society. Armillaria epidemiology has been studied within forestry and agriculture, but data are lacking within gardens, which have greater host plant diversity than orchards and vineyards and greater disturbance than woodlands. Which Armillaria species are responsible for garden disease, and how the broad range of susceptible ornamentals are differentially affected is not known. To address this, isolates of Armillaria were obtained from dead and dying plants from across the U.K. over a 4-year period (2004 to 2007). Species were identified by PCR-RFLP for IGS, and further verified by species-specific PCR for EF-1 α. Of the seven species known in the U.K., three were identified: A. mellea (83.1%), A. gallica (15.8%), and A. ostoyae (1.1%). Armillaria was isolated from trees, shrubs, and nonwoody plants including bulbs and vegetables, with newly recorded hosts listed herein. A. mellea was associated with infections of multiple hosts, and with all infections of the most common host, Ligustrum. In sites where more than one Armillaria species was found, the combination was of A. mellea and A. gallica, raising questions regarding the interactions of these species in U.K. gardens. © 2017, American Phytopathological Society. All rights reserved.
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In this report, we assess the unmitigated pest risk potential of importing Eucalyptus logs and chips from South America into the United States. To do this, we estimated the likelihood and consequences of introducing representative insects and pathogens of concern. Nineteen individual pest risk assessments were prepared, eleven dealing with insects and eight with pathogens. The selected organisms were representative examples of insects and pathogens found on the foliage, on the bark, in the bark, and in the wood of Eucalyptus spp. Among the insects and pathogens assessed, eight were rated a high risk potential: purple moth (Sarsina violescens), scolytid bark and ambrosia beetles (Scolytopsis brasiliensis, Xyleborus retusus, Xyleborus biconicus, Xyleborus spp.), carpenterworm (Chilecomadia valdiviana) on Eucalyptus nitens, round-headed wood borers (Chydarteres striatus, Retrachyderes thoracicus, Trachyderes spp., Steirastoma breve, Stenodontes spinibarbis), eucalyptus longhorned borer (Phoracantha semipunctata), Botryosphaeria cankers (Botryosphaeria dothidea, Botryosphaeria obtusa, Botryosphaeria ribi), Ceratocystis canker (Ceratocystis fimbriata), and pink disease (Erythricium salmonicolor). A moderate pest risk potential was assigned to eleven other organisms or groups of organisms: eucalypt weevils (Gonipterus spp.), carpenterworm (Chilecomadia valdiviana) on two Eucalyptus species other than E. nitens, platypodid ambrosia beetle (Megaplatypus parasulcatus), yellow phorancantha borer (Phoracantha recurva), subterranean termites (Coptotermes spp., Heterotermes spp.), foliar diseases (Aulographina eucalypti, Cryptosporiopsis eucalypti, Cylindrocladium spp., Phaeophleospora spp., Mycosphaerella spp.), eucalyptus rust (Puccinia psidii), Cryphonectria canker (Cryphonectria cubensis), Cytospora cankers (Cytospora eucalypticola, Cytospora eucalyptina), Coniothyrium canker (Coniothyrium zuluense), and root and stem rots (Armillaria spp., Phellinus spp., Ganoderma sp., Gymnopilus spectabilis). For those organisms of concern that are associated with logs and chips of South American Eucalyptus spp., specific phytosanitary measures may be required to ensure the quarantine safety of proposed importations.
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Numerous gap-model studies have attempted to predict responses of tree populations to climate change. Although these models incorporate various disturbance factors, the multifaceted influences of structural-root disease activity on gap dynamics have not been included [e.g., (2000)]. The fact that apparently similar models yield very different projections for the same forest [e.g., (1997)] suggests that most of these models are incomplete and/or may be inappropriately formulated (Bachelet and Neilson 2000). The existence of belowground root gaps corresponding to aboveground canopy gaps and their influence on plant competition and establishment have also received little attention [see (1997)]. Yet, Armillaria root disease and other structural-root diseases have been associated with gap dynamics [e.g., (1988); (1993); (1993), 1995); (1997); (2000)].
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In the Złotów Forest District 5 pine stands were selected and in each of which 4 observation plots were established each, consisting of 50 trees growing in clusters. The study consisted in one count of dead trees per year conducted in observation plots and in the whole experimental area. Results of the 3-year study showed the applied test tree method is not suitable in the assessment of threat to age class I pine stands posed by Ar-millaria fungi.
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The species identification of 645 Armillaria isolates was made in a nation-wide survey in Albania. The material was collected from ca. 250 permanent plots, established for monitoring forest health, and from forests and gardens attacked by Armillaria. A. mellea s.s. was found on several coniferous and broadleaved trees in most areas examined, except above 1100-1200 m in northern Albania. It causes some damage to fir and oak species and in a lesser degree to other forest trees. It is also common and damaging in fruit orchards and vineyards. A. gallica was a common saprophyte or weak pathogen in coniferous and deciduous forests at altitudes from 600 to 1600 m, and less common at lower altitudes on oaks. A. ostoyae was rare in central and southern Albania, but common in northern Albania, causing significant damage to pine and other conifers, mostly at altitudes from 600 to 1800 m. A. cepistipes was recorded at altitudes from 800 to 1800 m as a saprophyte or weak pathogen on conifers and deciduous trees, mostly in beech and silver fir forests. A. tabescens was found preferably in oak forests at altitudes from sea level to 900 m. In fruit orchards it occasionally attacked almond and pear trees. A. borealis was found on a few localities in northernmost Albania, at altitudes from 800 to1800 m. A. Mediterranea was found on a few localities in souththernmost Albania, at altitudes from 0 to100 m. The possible management of Armillaria species is necessary for healing of forests, orchards, etc. During the years in Albania the possible management Armillaria species was carried out by means of: Inoculums’ Reduction; Resistant Species; and Increasing Host Vigor.
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In Zielonka Forest District seven Scots pine plantations monitored in 1998-2001 for Armillaria ostoyae and Heterobasidion annosum disease development proved affected mainly by the former pathogen. Observations of 200 trees per plantation (every spring and fall) showed there was no correlation between the morphological features and over ground symptoms and infestation of pines by the pathogens in question and mortality of trees. The young stands were threatened more by Armillaria than by Heterobasidion annosum s.s., yet the infestation by Armillaria decreased with age of trees while the infestation by Heterobasidion increased.
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Ponderosa pine (Pinus ponderosa) in the Black Hills National Forest, SD, USA, was surveyed for Armillaria root disease (ARD). The root pathogen Armillaria ostoyae occurred on ponderosa pine seedlings, saplings, pole‐size trees and large‐diameter trees. The mean incidence of aboveground disease symptoms by stem count was low (0.2%), but in certain areas, the incidence was higher, affecting the regeneration success and tree longevity. Symptomatic ponderosa pine were in areas characterized by having higher elevation, greater annual precipitation, more seedlings, bigger large‐diameter trees and greater odds of past harvesting activity than in areas without root disease. Stump density was positively spatially correlated with root disease incidence. No particular soil type was related to disease occurrence; though, in areas with symptomatic trees, soil available water holding capacity (AWC) was greater and soil permeability was less where root disease was present. Spatial analysis confirmed the relationships found in linear correlations, with soil AWC and stump density positively and soil permeability negatively correlated with per cent infected stems ha−1 and basal area infected.
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