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Fires in coniferous forests throughout the northern United States alter ecosystem processes and ecological communities, including the diversity and composition of microbial communities living in the soil. In addition to its influence on ecosystem processes and functions, the soil microbiome can interact with soilborne pathogens to facilitate or suppress plant disease development. Altering the microbiome composition to promote taxa that inhibit pathogenic activity has been suggested as a management strategy for forest diseases, including Armillaria root disease caused by Armillaria solidipes , which causes growth loss and mortality of conifers. These forest ecosystems are experiencing increased wildfire burn severity that could influence A. solidipes activity and interactions of the soil microbiome with Armillaria root disease. In this research, we examine changes to the soil microbiome following three levels of burn severity in a coniferous forest in northern Idaho, United States, where Armillaria root disease is prevalent. We further determine how these changes correspond to the soil microbiomes associated with the pathogen A. solidipes, and a putatively beneficial species, A. altimontana . At 15-months post-fire, we found significant differences in richness and diversity between bacterial communities associated with unburned and burned areas, yet no significant changes to these metrics were found in fungal communities following fire. However, both bacterial and fungal communities showed compositional changes associated with burn severity, including microbial taxa with altered relative abundance. Further, significant differences in the relative abundance of certain microbial taxa in communities associated with the three burn severity levels overlapped with taxa associated with various Armillaria spp. Following severe burn, we observed a decreased relative abundance of beneficial ectomycorrhizal fungi associated with the microbial communities of A. altimontana, which may contribute to the antagonistic activity of this soil microbial community. Additionally, A. solidipes and associated microbial taxa were found to dominate following high-severity burns, suggesting that severe fires provide suitable environmental conditions for these species. Overall, our results suggest that shifts in the soil microbiome and an associated increase in the activity of A. solidipes following high-severity burns in similar conifer forests may result in priority areas for monitoring and proactive management of Armillaria root disease., Importance: With its influence on ecosystem processes and functions, the soil microbiome can interact with soilborne pathogens to facilitate or suppress plant disease development. These forest ecosystems are experiencing increased wildfire frequency and burn severity that could influence the fungal root pathogen, Armillaria solidipes , and interactions with the soil microbiome. We examined changes to the soil microbiome following three levels of burn severity, and examined how these changes correspond with A. solidipes , and a putatively beneficial species, A. altimontana . Following severe burn, there was a decreased relative abundance of ectomycorrhizal fungi associated A. altimontana . A. solidipes and associated microbial taxa dominated following high-severity burns, suggesting that severe fires provide suitable environmental conditions for these species. Our results suggest that shifts in the soil microbiome and an associated increase in the activity of A. solidipes following high-severity burns in conifer forests may result in priority areas for monitoring and proactive management of Armillaria root disease., Competing Interests: The authors declare no conflict of interest.

Armillaria species show considerable variation in ecological roles and virulence, from mycorrhizae and saprophytes to important root pathogens of trees and horticultural crops. We studied two Armillaria species that can be found in coniferous forests of northwestern USA and southwestern Canada. Armillaria altimontana not only is considered as a weak, opportunistic pathogen of coniferous trees, but it also appears to exhibit in situ biological control against A. solidipes, formerly North American A. ostoyae, which is considered a virulent pathogen of coniferous trees. Here, we describe their genome assemblies and present a functional annotation of the predicted genes and proteins for the two Armillaria species that exhibit contrasting ecological roles. In addition, the soil microbial communities were examined in association with the two Armillaria species within a 45-year-old plantation of western white pine (Pinus monticola) in northern Idaho, USA, where A. altimontana was associated with improved tree growth and survival, while A. solidipes was associated with reduced growth and survival. The results from this study reveal a high similarity between the genomes of the beneficial/non-pathogenic A. altimontana and pathogenic A. solidipes; however, many relatively small differences in gene content were identified that could contribute to differences in ecological lifestyles and interactions with woody hosts and soil microbial communities., (© 2022. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Armillaria possesses several intriguing characteristics that have inspired wide interest in understanding phylogenetic relationships within and among species of this genus. Nuclear ribosomal DNA sequence-based analyses of Armillaria provide only limited information for phylogenetic studies among widely divergent taxa. More recent studies have shown that translation elongation factor 1-α (tef1) sequences are highly informative for phylogenetic analysis of Armillaria species within diverse global regions. This study used Neighbor-net and coalescence-based Bayesian analyses to examine phylogenetic relationships of newly determined and existing tef1 sequences derived from diverse Armillaria species from across the Northern Hemisphere, with Southern Hemisphere Armillaria species included for reference. Based on the Bayesian analysis of tef1 sequences, Armillaria species from the Northern Hemisphere are generally contained within the following four superclades, which are named according to the specific epithet of the most frequently cited species within the superclade: (i) Socialis/Tabescens (exannulate) superclade including Eurasian A. ectypa, North American A. socialis (A. tabescens), and Eurasian A. socialis (A. tabescens) clades; (ii) Mellea superclade including undescribed annulate North American Armillaria sp. (Mexico) and four separate clades of A. mellea (Europe and Iran, eastern Asia, and two groups from North America); (iii) Gallica superclade including Armillaria Nag E (Japan), multiple clades of A. gallica (Asia and Europe), A. calvescens (eastern North America), A. cepistipes (North America), A. altimontana (western USA), A. nabsnona (North America and Japan), and at least two A. gallica clades (North America); and (iv) Solidipes/Ostoyae superclade including two A. solidipes/ostoyae clades (North America), A. gemina (eastern USA), A. solidipes/ostoyae (Eurasia), A. cepistipes (Europe and Japan), A. sinapina (North America and Japan), and A. borealis (Eurasia) clade 2. Of note is that A. borealis (Eurasia) clade 1 appears basal to the Solidipes/Ostoyae and Gallica superclades. The Neighbor-net analysis showed similar phylogenetic relationships. This study further demonstrates the utility of tef1 for global phylogenetic studies of Armillaria species and provides critical insights into multiple taxonomic issues that warrant further study.

Desarmillaria caespitosa, a North American vicariant species of European D. tabescens, is redescribed in detail based on recent collections from the USA and Mexico. This species is characterized by morphological features and multilocus phylogenetic analyses using portions of nuc rDNA 28S (28S), translation elongation factor 1-alpha (tef1), the second largest subunit of RNA polymerase II (rpb2), actin (act), and glyceraldehyde-3-phosphate dehydrogenase (gpd). A neotype of D. caespitosa is designated here. Morphological and genetic differences between D. caespitosa and D. tabescens were identified. Morphologically, D. caespitosa differs from D. tabescens by having wider basidiospores, narrower cheilocystidia, which are often irregular or mixed (regular, irregular, or coralloid), and narrower caulocystidia. Phylogenetic analyses of five independent gene regions show that D. caespitosa and D. tabescens are separated by nodes with strong support. The new combination, D. caespitosa, is proposed. [ABSTRACT FROM AUTHOR]

• Stand density and the presence of D. confusus are associated with mortality. • Density (basal area), elevation, and Armillaria are directly associated with the presence of D. confusus. • Increased summer temperatures and slope are associated with the presence of Armillaria. • Increased stand density and summer temperatures cause stress in trees. • Stress promotes Armillaria spp. and D. confusus , leading to subalpine fir mortality. Subalpine fir mortality complex has caused significant damage to high elevation forests within Colorado. For the past two decades the climate of spruce-fir forests has trended towards being warmer and drier, which indirectly affects subalpine fir mortality. We examined potential links among abiotic (i.e. deviations in temperature and precipitation), biotic [Armillaria root disease (Armillaria spp.), Western balsam bark beetle (Dryocoetes confusus), a black stain fungus (Ophiostoma dryocoetidis)] and forest structure factors with mortality in subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii) dominated forests of Colorado. Sampling was performed in two separate field surveys: 1. Statewide characterization plots and 2. Stand health monitoring plots. The statewide characterization plots were compared to aerial surveys and to establish suitable locations for stand health plots. Stand health monitoring plots collected direct measurements of abiotic and biotic factors affecting forests, with an emphasis on identifying the presence of D. confusus and Armillaria spp. The objectives of this study were to determine: (1) Is spruce-fir morality observed at ground level similar to that estimated through aerial surveys? (2) Is the presence dead subalpine fir and Englemann spruce on a plot associated with site and stand characteristics?, (3) What factors are associated with the incidence of Armillaria spp. , Dr. confusus, and Dendroctonus rufipennis ?, and (4) Do trends in warming temperatures and reductions in precipitation in the past five years influence subalpine fir mortality and/or the presence of biotic agents? Our results suggested that the presence of biotic agents (Dr. confusus , De. rufipennis, Armillaria spp., and O. dryocoetidis) and stand density are associated with spruce-fir mortality, whereas climatic factors had a direct association to the presence of Armillaria and were indirectly associated with mortality via Dr. confusus. Increasing maximum summer temperatures were found to be associated with the presence of Armillaria spp. The climatic variables investigated in this study did not significantly link to incidence of Dr. confusus , yet stand density was associated with increasing prevalence of Dr. confusus. The presence of Armillaria spp. and Dr. confusus were significantly related to tree decline, but several other factors are also associated with subalpine fir mortality, suggesting complex interactions of abiotic and biotic factors are likely involved. We hypothesized that increasing stand density, drought and the presence of Dr. confusus and Armillaria spp. are threatening Colorado spruce-fir forests. Understanding the factors that are involved in subalpine fir mortality are important for management. [ABSTRACT FROM AUTHOR]

Research Highlights: Two genets of Armillaria altimontana Brazee, B. Ortiz, Banik, and D.L. Lindner and five genets of Armillaria solidipes Peck (as A. ostoyae [Romagnesi] Herink) were identified and spatially mapped within a 16-year-old western white pine (Pinus monticola Doug.) plantation, which demonstrated distinct spatial distribution and interspecific associations. Background and Objectives: A. solidipes and A. altimontana frequently co-occur within inland western regions of the contiguous USA. While A. solidipes is well-known as a virulent primary pathogen that causes root disease on diverse conifers, little has been documented on the impact of A. altimontana or its interaction with A. solidipes on growth, survival, and the Armillaria root disease of conifers. Materials and Methods: In 1971, a provenance planting of P. monticola spanning 0.8 ha was established at the Priest River Experimental Forest in northern Idaho, USA. In 1987, 2076 living or recently dead trees were measured and surveyed for Armillaria spp. to describe the demography and to assess the potential influences of Armillaria spp. on growth, survival, and the Armillaria root disease among the study trees. Results: Among the study trees, 54.9% were associated with Armillaria spp. The genets of A. altimontana and A. solidipes comprised 82.7% and 17.3% of the sampled isolates (n = 1221) from the study plot, respectively. The mapped distributions showed a wide, often noncontiguous, spatial span of individual Armillaria genets. Furthermore, A. solidipes was found to be uncommon in areas dominated by A. altimontana. The trees colonized by A. solidipes were associated with a lower tree growth/survival and a substantially higher incidence of root disease than trees colonized only by A. altimontana or trees with no colonization by Armillaria spp. Conclusions: The results demonstrate that A. altimontana was not harmful to P. monticola within the northern Idaho planting. In addition, the on-site, species-distribution patterns suggest that A. altimontana acts as a long-term, in situ biological control of A. solidipes. The interactions between these two Armillaria species appear critical to understanding the Armillaria root disease in this region. [ABSTRACT FROM AUTHOR]