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Pyrophilous fungi detected after wildfires in the Great Smoky Mountains National Park expand known species ranges and biodiversity estimates
Mycologia
Abstract
Following a late fall wildfire in 2016 in the Great Smoky Mountains National Park, pyrophilous fungi in burn zones were documented over a 2-y period with respect to burn severity and phenology. Nuc rDNA internal transcribed spacer (ITS1-5.8S-ITS2 = ITS) barcodes were obtained to confirm morphological evaluations. Forty-one taxa of Ascomycota and Basidiomycota were identified from burn sites and categorized as fruiting only in response to fire or fruiting enhanced by fire. Twenty-two species of Pezizales (Ascomycota) were among the earliest to form ascomata in severe burn zones, only one of which had previously been documented in the Great Smoky Mountains National Park. Nineteen species of Basidiomycota, primarily Agaricales, were also documented. Among these, only five species (Coprinellus angulatus, Gymnopilus decipiens, Lyophyllum anthracophilum, Pholiota carbonicola, and Psathyrella pennata) were considered to be obligate pyrophilous taxa, but fruiting of two additional taxa (Hygrocybe conica and Mycena galericulata) was clearly enhanced by fire. Laccaria trichodermophora was an early colonizer of severe burn sites and persisted through the winter of 2017 and into spring and summer of 2018, often appearing in close association with Pinus pungens seedlings. Fruiting of pyrophilous fungi peaked 4–6 mo post fire then diminished, but some continued to fruit up to 2.5 y after the fire. In all, a total of 27 previously unrecorded taxa were added to the All Taxa Biodiversity Inventory (ATBI) database (~0.9%). Most pyrophilous fungi identified in this study are either cosmopolitan or have a Northern Hemisphere distribution, but cryptic endemic lineages were detected in Anthracobia and Sphaerosporella. One new combination, Hygrocybe spadicea var. spadicea f. odora, is proposed.
... Another hypothesis is that the reduction in fruiting after the first year is intrinsic to the fungi themselves. A few fungal taxa are known to be pyrophilous and require fire to reproduce (Seaver 1909;Wicklow 1973Wicklow , 1975Petersen 1974;Hughes et al., 2020) and some parasitic ascomycetes increase sporocarp production in response to post-fire damage of their hosts (Robinson et al., 2008;Sapphire JM et al., 2011). Perhaps after a large sporocarp production event following a burn, the following year or two will have smaller yields, similar to lowered production in fruit and nut trees following a big season (masting events). ...
... Several Mycena are now known to be parasitic (Thoen et al., 2020), and based on its strong association with grass roots, we suspect M. citrinomarginata is parasitic. Sporocarp production in at least three other Mycena species is known to be stimulated by fire (McMullan-Fisher et al., 2002;Hughes et al., 2020;Kouki and Salo 2020), and at least one of these species is also strongly host associated: Mycena galericulata. When found growing with Rhododendron and Kalmia, M. galericulata "fruited in exceptionally large numbers in severe burn areas" (Hughes et al., 2020). ...
... Sporocarp production in at least three other Mycena species is known to be stimulated by fire (McMullan-Fisher et al., 2002;Hughes et al., 2020;Kouki and Salo 2020), and at least one of these species is also strongly host associated: Mycena galericulata. When found growing with Rhododendron and Kalmia, M. galericulata "fruited in exceptionally large numbers in severe burn areas" (Hughes et al., 2020). This massive fruiting is similar to what we observed and is in opposition to the usual response of saprobic Mycena, which typically decrease immediately after fire in response to loss of woody material and litter (McMullan-Fisher et al., 2002;Boberg et al., 2011;Crabtree 2014;Semenova-Nelsen et al., 2019). ...
... These differences may facilitate or inhibit the movement and resilience of wildlife for conservation (e.g., elk) or hunting (e.g., white-tailed deer) when faced with disturbances like the recent wildfires. Similar changes in ranges in response to wildfire have been quantified in the park for fungi (Hughes et al. 2020). Jurisdictional contributions to biodiversity and other important ecological properties can provide a heuristic that further research is needed into the role of the social system across this landscape and other protected area-centered ecosystems. ...
... When managers pursue conservation or resource extraction objectives focused on individual species, such changes can threaten their success (Schwartz et al. 2019). The Great Smoky Mountains PACE has similar challenges with exotic invasions (Johnson et al. 2005;Resnik 2018) and, more recently, wildfires (Hughes et al. 2020) to many PACEs in the western United States. Biological invasions and wildfires are two management challenges that cross boundaries, so the behaviors and decisions of neighboring land managers can have direct impact on a focal management unit's levels of invasion or wildfire risk (Fischer and Charnley 2012;Epanchin-Niell and Wilen 2015;Ager et al. 2018;Fischer et al. 2019;Charnley et al. 2020;Dayer et al. 2020;Latombe et al. 2020). ...
... Some landowners may be unaware of the risks and management options at hand (Marzano et al. 2015). Management collaboration and education among neighbors may help reduce the risk, for example, from devastating wildfires like those that impacted the GRSM PACE, destroying homes and businesses, in 2016 (Hughes et al. 2020). ...
Land management decisions have profound impacts on biological communities. Various administrative units are likely to implement different management decisions, with the result that consistent biological patterns that track administrative boundaries may emerge. To assess these relationships, we collected data on land cover, disturbance evidence, and tree communities in randomized sampling sites located in National Park, US Forest Service Wilderness, and US Forest Service Non-wilderness distributed across the Great Smoky Mountains National Park Protected Area-Centered Ecosystem (GRSM PACE). We examined how present-day values for these ecological responses were linked to both biophysical site characteristics (as the foundation for ecological communities) and jurisdiction type (as categories that integrate the combination of dynamic social decisions and management practices since initial designation). The variable representing administrative unit, jurisdiction type, acted as an umbrella indicator of the management trajectory and social influences that have affected a site and emerged as an important predictor of groundcover types (grass and litter), tree diversity, and wildlife sign at sites across the GRSM PACE. This was an exploratory study with limited sample size, so more data are needed to explore the mechanisms driving these relationships. These initial findings, however, suggest that differences in management activities alter the ecological trajectory of sites across the PACE. Given dynamic and diverse management practices, our data suggest jurisdiction overlaid on biophysical site information may be useful to rapidly assess how socially defined landscape activities impact biological responses, particularly when both are stressed by global change.
... Pyrophilous fungi have historically only been found fruiting on burned soil, and they are known fire-responsive fungal colonizers of post-fire soil . Many of these fungi are Ascomycota of the Pezizales order (El-Abyad & Webster, 1968;Hughes et al., 2020), although some Basidiomycota are also represented in the pyrophilous guild (Steindorff et al., 2021). These fungi are known to fruit in a predictable order after fire, and observations of this phenomenon have been made at various sites world-wide (Petersen, 1970;Warcup, 1990;Bruns et al., 2020;Fischer et al., 2021). ...
... The soil colonization after fire events happens in successionnot all of the species grow simultaneously Hughes et al., 2020). We considered pyrotolerant fungi opportunists that occupy, at a later stage than pyrophilous fungi, substrates and habitats made available by fires (Fox et al., 2022). ...
... In addition, we performed a comprehensive comparison using 124 Dikarya genomes, including pyrophilous Agaricales genomes (Steindorff et al., 2021) (Table S6). Even though most pyrophilous fungi analyzed in this study belong to the Pyronemataceae family, they colonize the post-fire soil in different periods Hughes et al., 2020). Hence, to make an intra-Pezizales enrichment comparison, we separated the data into early and late colonizers. ...
Wildfires drastically impact the soil environment, altering the soil organic matter, forming pyrolyzed compounds, and markedly reducing the diversity of microorganisms. Pyrophilous fungi, especially the species from the orders Pezizales and Agaricales, are fire‐responsive fungal colonizers of post‐fire soil that have historically been found fruiting on burned soil and thus may encode mechanisms of processing these compounds in their genomes.
Pyrophilous fungi are diverse. In this work, we explored this diversity and sequenced six new genomes of pyrophilous Pezizales fungi isolated after the 2013 Rim Fire near Yosemite Park in California, USA: Pyronema domesticum, Pyronema omphalodes, Tricharina praecox, Geopyxis carbonaria, Morchella snyderi, and Peziza echinospora.
A comparative genomics analysis revealed the enrichment of gene families involved in responses to stress and the degradation of pyrolyzed organic matter. In addition, we found that both protein sequence lengths and G + C content in the third base of codons (GC3) in pyrophilous fungi fall between those in mesophilic/nonpyrophilous and thermophilic fungi.
A comparative transcriptome analysis of P. domesticum under two conditions – growing on charcoal, and during sexual development – identified modules of genes that are co‐expressed in the charcoal and light‐induced sexual development conditions. In addition, environmental sensors such as transcription factors STE12, LreA, LreB, VosA, and EsdC were upregulated in the charcoal condition.
Taken together, these results highlight genomic adaptations of pyrophilous fungi and indicate a potential connection between charcoal tolerance and fruiting body formation in P. domesticum.
... Similarly, downed wood and surviving plant tissues (e.g., leaves, roots) can provide insulation that protects substrate-or tissueinhabiting fungi from heat-induced mortality, exemplifying a "fire refugium" (see Meddens et al. 2018). In addition, some post-fire "blooms" of fungi may be due to the loss of substrate or host plants, which triggers a fruiting response (Fujimura et al. 2005;Hughes et al. 2020b;Kuo et al. 2014). Finally, repeated exposure to fires can also dictate fungal responses. ...
... Anthracobia spp. have been reported fruiting in North America following the Washington State Mount St. Helens eruption (Claridge et al. 2009) and wildfires in the western (Claridge et al. 2009;Fujimura et al. 2005) and eastern (Hughes et al. 2020b) United States. They have also been observed fruiting after wildfires in Australia (Sjm et al. 2011;Warcup 1990), the United Kingdom (Wilberforce 2005), and Denmark (Dix and Webster 1995). ...
... (reported as P. omphalodes in Warcup 1990) fruited in high abundance after wildfires in Eucalyptus forests in Australia (Warcup 1990), and P. omphalodes (= P. confluens) has been reported to fruit after fires across the United States in Iowa, New York, and North Dakota (Seaver 1909). After a 2016 wildfire in Tennessee, P. omphalodes, but not P. domesticum, was recovered both as ascomata on burned soil (Hughes et al. 2020b) and as ITS2 sequences from severely burned soils but not in great abundance and only in hemlock/pine areas (Hughes, unpublished data). Other studies report that P. domesticum and P. omphalodes accounted for approximately 60% of ITS1 sequences after a mega-fire burned down a Pinus ponderosa forest in the California Sierra Nevada ) and after a high-severity fire burned a California chaparral shrubland (FIG. ...
Fires occur in most terrestrial ecosystems where they drive changes in the traits, composition, and diversity of fungal communities. Fires range from rare, stand-replacing wildfires to frequent, prescribed fires used to mimic natural fire regimes. Fire regime factors, including burn severity, fire intensity, and timing, vary widely and likely determine how fungi respond to fires. Despite the importance of fungi to post-fire plant communities and ecosystem functioning, attempts to identify common fungal responses and their major drivers are lacking. This synthesis addresses this knowledge gap and ranges from fire adaptations of specific fungi to succession and assembly fungal communities as they respond to spatially heterogenous burning within the landscape. Fires impact fungi directly and indirectly through their effects on fungal survival, substrate and habitat modifications , changes in environmental conditions, and/or physiological responses of the hosts with which fungi interact. Some specific pyrophilous, or "fire-loving," fungi often appear after fire. Our synthesis explores whether such taxa can be considered cosmopolitan, and whether they are truly fire-adapted or simply opportunists adapted to rapidly occupy substrates and habitats made available by fires. We also discuss the possible inoculum sources of post-fire fungi and explore existing conceptual models and ecological frameworks that may be useful in generalizing fungal fire responses. We conclude with identifying research gaps and areas that may best transform the current knowledge and understanding of fungal responses to fire.
... Several common soil fungi have also been shown to degrade polyaromatic compounds (Cerniglia and Sutherland, 2010). These fungi include Neurospora crassa, which emerges from burned wood shortly after fire, and Morchella conica, which is a relative of pyrophilous Morchella species that often co-occur with Pyronema species (Lin and Kapoor, 1979;Gramss et al., 1999;Davis, 2000;Hughes et al., 2020). ...
... Fruiting bodies of the genus Pyronema are among the first macrofungi to emerge from burned soil, doing so within weeks to months after fire (Seaver, 1909;Adamczyk et al., 2012;Bruns et al., 2020;Hughes et al., 2020; Figures 1A,B). There are two currently accepted species of Pyronema: P. domesticum and P. omphalodes (= P. confluens), both of which rapidly dominate post-fire fungal communities (Bruns et al., 2020). ...
... Although their prevalence is relatively short-lived, Pyronema grow rapidly post-fire, producing abundant biomass in the form of ascocarps and mycelia (Seaver, 1909;Adamczyk et al., 2012;Bruns et al., 2020;Hughes et al., 2020). However, both Pyronema ascocarps and their DNA decline rapidly after they peak in abundance following fire (Bruns et al., 2020). ...
Wildfires represent a fundamental and profound disturbance in many ecosystems, and their frequency and severity are increasing in many regions of the world. Fire affects soil by removing carbon in the form of CO2 and transforming remaining surface carbon into pyrolyzed organic matter (PyOM). Fires also generate substantial necromass at depths where the heat kills soil organisms but does not catalyze the formation of PyOM. Pyronema species strongly dominate soil fungal communities within weeks to months after fire. However, the carbon pool (i.e., necromass or PyOM) that fuels their rise in abundance is unknown. We used a Pyronema domesticum isolate from the catastrophic 2013 Rim Fire (CA, United States) to ask whether P. domesticum is capable of metabolizing PyOM. Pyronema domesticum grew readily on agar media where the sole carbon source was PyOM (specifically, pine wood PyOM produced at 750°C). Using RNAseq, we investigated the response of P. domesticum to PyOM and observed a comprehensive induction of genes involved in the metabolism and mineralization of aromatic compounds, typical of those found in PyOM. Lastly, we used 13C-labeled 750°C PyOM to demonstrate that P. domesticum is capable of mineralizing PyOM to CO2. Collectively, our results indicate a robust potential for P. domesticum to liberate carbon from PyOM in post-fire ecosystems and return it to the bioavailable carbon pool.
... Post-fire fungi are categorized into two main categories based on their response to fruiting in burnt habitats [5]: (1) fruiting occurs only after a fire, and (2) fruiting can occur at any time, but fruiting is enhanced after a fire. Anthracobia spp., Ascobolus spp., Morchella exuberans Clowez, Hugh Sm. and S. Sm., Peziza echinospora P. Karst., Plicaria spp., Psathyrella Life 2021, 11, 518 2 of 10 pennata (Fr.) A. Pearson and Dennis, Pholiota carbonaria (Fr.) ...
... Gray, Sphaerosporella spp., and Thelephora terrestris Ehrh. [5]. ...
... There is an apparent lack of knowledge concerning the colonization of bryophytes on burnt wood, which is an essential topic when planning forest restoration treatments that aim to increase biodiversity. In relation to P. carbonaria, basidiomata of this species start to appear about five months after a fire event and continue fruiting for at least 18 months [5], placing both fungal spores and maturing bryophytes in close temporal and spatial proximity. ...
Wildfires play a critical role in maintaining biodiversity and shaping ecosystem structure in fire-prone regions, and successional patterns involving numerous plant and fungal species in post-fire events have been elucidated. Evidence is growing to support the idea that some post-fire fungi can form endophytic/endolichenic relationships with plants and lichens. However, no direct observations of fire-associated fungal–moss interactions have been visualized to date. Therefore, physical interactions between a post-fire fungus, Pholiota carbonaria, and a moss, Polytrichum commune, were visually examined under laboratory conditions. Fungal appressoria were visualized on germinating spores and living protonemata within two weeks of inoculation in most growth chambers. Appressoria were pigmented, reddish gold to braun, and with a penetration peg. Pigmented, reddish gold to braun fungal hyphae were associated with living tissue, and numerous mature rhizoids contained fungal hyphae at six months. Inter-rhizoidal hyphae were pigmented and reddish gold to braun, but no structures were visualized on mature gametophyte leaf or stem tissues. Based on our visual evidence and previous work, we provide additional support for P. carbonaria having multiple strategies in how it obtains nutrients from the environment, and provide the first visual documentation of these structures in vitro.
... There were, however, pyrophilous taxa that we did not detect in our study. For example, we did not detect Pyronema spp. or Pholiota carbonaria that have been found to produce fruiting bodies following wildfire 43,75,76 . We also did not detect any pyrophilous insects. ...
... Species were retained if the bootstrap values were at least 0.80 for assignments that were at least 70% correct based on leave one sequence out testing using a reference sequence database based on UNITE and assuming that query taxa are present in the reference set 103 . ITS SVs were screened for the presence of known pyrophilous (fire-loving) fungi based on studies that used sporocarp sampling at the genus and species ranks 43,47,76 . Fungal ITS sequences were then assigned using the FungalTraits database by importing their reference set into R 119 . ...
Wildfire is a natural disturbance in boreal forest systems that has been predicted to increase in frequency, intensity, and extent due to climate change. Most studies tend to assess the recovery of one component of the community at a time but here we use DNA metabarcoding to simultaneously monitor soil bacteria, fungi, and arthropods along an 85-year chronosequence following wildfire in jack pine-dominated ecosites. We describe soil successional and community assembly processes to better inform sustainable forest management practices. Soil taxa showed different recovery trajectories following wildfire. Bacteria shared a large core community across stand development stages (~ 95–97% of their unique sequences) and appeared to recover relatively quickly by crown closure. By comparison fungi and arthropods shared smaller core communities (64–77% and 68–69%, respectively) and each stage appeared to support unique biodiversity. We show the importance of maintaining a mosaic ecosystem that represents each stand development stage to maintain the full suite of biodiversity in soils following wildfire, especially for fungi and arthropods. These results will provide a useful baseline for comparison when assessing the effects of human disturbance such as harvest or for assessing the effects of more frequent wildfire events due to climate change.
... The Scutellinia OTU7 matched closely Anthracobia sp. in Pyronemataceae (100% query coverage and 100% sequence similarity to GenBank accession MK840934.1)-a likely pyrophilous fungus (Claridge, Trappe and Hansen 2009;Hughes et al. 2020). To focus on fire effects, we also analysed samples from within and outside the shrub islands together. ...
... Any combination of these factors may alter relative competitiveness in the resident communities and stimulate organisms that respond positively and proliferate across fire-impacted landscapes. High-severity fires may be unique in that they may open niches for opportunistic taxa that a lowseverity fire would not ( Figure S6, Supporting Information), and these taxa may thrive in post-fire grasslands (Yang et al. 2020) or forests (Fujimura et al. 2005;Hughes et al. 2020). ...
Fire can impact terrestrial ecosystems by changing abiotic and biotic conditions. Short fire intervals maintain grasslands and communities adapted to frequent, low-severity fires. Shrub encroachment that follows longer fire intervals accumulates fuel and can increase fire severity. This patchily distributed biomass creates mosaics of burn severities in the landscape—pyrodiversity. Afforded by a scheduled burn of a watershed protected from fires for 27 years, we investigated effects of woody encroachment and burn severity on soil chemistry and soil-inhabiting bacteria and fungi. We compared soils before and after fire within the fire-protected, shrub-encroached watershed and soils in an adjacent, annually burned, non-encroached watershed. Organic matter and nutrients accumulated in the fire-protected watershed but responded less to woody encroachment within the encroached watershed. Bioavailable nitrogen and phosphorus and fungal and bacterial communities responded to high severity burn regardless of encroachment. Low severity fire effects on soil nutrients differed, increased bacterial but decreased fungal diversity, and effects of woody encroachment within the encroached watershed were minimal. High severity burns in the fire-protected watershed led to a novel soil system state distinct from non-encroached and encroached soil systems. We conclude that severe fires may open grassland restoration opportunities to manipulate soil chemistry and microbial communities in shrub-encroached habitats.
... Pyrophilous (fire-loving) fungi are a well-known, ecological guild of fungi that are restricted to post-fire environments and fruit abundantly in months immediately following fire (Petersen 1970, Hughes et al., 2020. Because of their predicable occurrence after fire, they almost certainly need to be able to interact with post-fire soil chemistry. ...
... It is important to note that pyrophilous fungi fruits are found only in burned habitats and are abundant in the first weeks, months or years after fire (Bruns et al., 2020, Hughes et al., 2020, Petersen, 1970, not necessarily right after the event. At first glance, the causal relationship between fire events and fruiting body development seems circumstantial. ...
Forest fires generate a large amount of carbon that remains resident on the site as dead and partially 'pyrolysed' (i.e. burnt) material that has long residency times and constitutes a significant pool in fire-prone ecosystems. In addition, fire-induced hydrophobic soil layers, caused by condensation of pyrolysed waxes and lipids, increase post-fire erosion and can lead to long-term productivity losses. A small set of pyrophilous fungi dominate post-fire soils and are likely to be involved with the degradation of all these compounds, yet almost nothing is currently known about what these fungi do or the metabolic processes they employ. In this study, we sequenced and analysed genomes from fungi isolated after Rim fire near Yosemite National Park in 2013 and showed the enrichment/expansion of CAZymes and families known to be involved in fruiting body initiation when compared to other basidiomycete fungi. We found gene families potentially involved in the degradation of the hydrophobic layer and pyrolysed organic matter, such as hydrophobic surface binding proteins, laccases (AA1_1), xylanases (GH10, GH11), fatty acid desaturases and tannases. Thus, pyrophilous fungi are important actors to restate the soil's functional capabilities.
... Hřiland 1990, Orton 1993, Ludwig 2000, 2001, Holec 2005, 2012) is a small fungus with a pileus up to 30 mm in diameter, dirty yellow-brown, rusty brown to greyish brown, fibrillose-tomentose to tomentose-scaly. However, the delimitation of G. decipiens is still unclear (Hughes et al. 2020, Holec et al. 2022. ...
Available as Open Acces in Czech Mycology 76(1): 83–94 (http://www.czechmycology.org/_cmo/CM76106.pdf). A new record of the rare agaric species Gymnopilus stabilis from the Czech Republic is described morphologically and genetically. The basidiomata show good agreement with the recently published epitype and diagnostic characters of the species. While the robustness of the basidiomata, the presence of a pink hue, and a distinct sweetish aromatic smell are typical characters, though not
always present, the predominantly warm orange colour of adult pilei seems to be stable character. The combination of fleshy basidiomata and typical pileus colour distinguishes G. stabilis from
G. penetrans/hybridus and G. decipiens, which are taxa sometimes confused with G. stabilis. Comparison of the nearest ITS rDNA sequences from GenBank confirmed the identity of our record and
showed that the species is distributed not only in Europe and Siberia, but also in Pakistan and India. The ecological characterisation of G. stabilis is updated, showing that it is a saprotrophic species on dead wood of conifers, both Pinus and Picea, but also a facultative anthracophilous fungus able to
grow on burnt wood and ash.
... It is known that A. maurilabra occurs exclusively on burnt sites as early as 7 weeks after burning, but does not fruit later than 80 weeks after burning (Dix & Webster, 1995;Dougoud, 2001). Pyronema species occur in burnt areas worldwide (Petersen, 1970;Dougoud, 2001) and produce apothecia fairly early after fires (Moore & Korf, 1963;Bruns et al., 2020;Hughes et al., 2020). T. praecox occurs on limestone and burnt ground and fruits abundantly only after a fire event (Yang & Korf, 1985b;Dougoud, 2001;Raudabaugh et al., 2020) reported that this species always associates with burnt sites, probably mostly in the first or second year after a fire. ...
Three pyrophilous discomycetes species (Anthracobia maurilabra (Cooke) Boud., Pyronema domesticum (Sowerby) Sacc. and Tricharina praecox (P. Karst.) Dennis) have been cultivated and studied in vitro. Cultures were obtained from fresh apothecia сollected in the Holosiivskyi National Nature Park (Kyiv, Ukraine). The culture growth and morphological characteristics of the studied fungi on different media (beer wort agar, Czapek Dox agar, potato-dextrose agar) were analyzed. All investigated species can quite easily grow under laboratory conditions on different tested nutrient media. Potato-dextrose agar was the most suitable medium for the enhancement of radial growth and the best expresses all the phenotypes of the colony of studied fungi. Macro- and micromorphological descriptions of all fungal colonies and illustrations are provided. The micromorphological analysis showed that common to the mycelium of all studied species of fungi was the presence of numerous drops of oil in the hyphae, anastomoses like T. praecox, а net- like structure of A. maurilabra similar to nematode capture hook. Beside this, A. maurilabra and P. domesticum formed the sexual stage under experimental conditions. Forming fruiting bodies of A. maurilabra in culture has not been reported before. Moreover, P. domesticum was found to form abundant dark brown sclerotia on potato-dextrose agar and Czapek Dox agar. Possible pigment composition in the P. domesticum sclerotia is discussed based on the Raman spectroscopy study, performed on this genus for the first time. The established cultural characteristics can be useful for taxonomic identification of fungal species and for pure quality control of mycelial cultures during their introduction, preservation and future potential applications in biotechnological areas.
... In this study, the fungi enriched in burnt soils included both expected and unexpected taxa. Pyronema domesticum, a well-documented pyrophilous fungus (Seaver, 1909;Reazin et al., 2016;Bruns et al., 2020;Hughes et al., 2020;Pérez-Izquierdo et al., 2020), was the most abundant ASV encountered in the burned soils, but we also identified 18 ASVs that have not been widely characterized as pyrophilous. Several genera that were enriched in burnt microplots, namely Didymella, Dioszegia, Itersonilia, Naganishia, Spizellomyces, and Sporobolomyces, have not been documented as being heat resistant, but are extremophiles capable of tolerating cold and/or alkaline environments (Onofri, 1999;Białkowska et al., 2017;Buzzini et al., 2018). ...
... In Whitman et al., Mucoromycota increased at sites with higher burn severity [9]. Pezizales are known to have increased abundance of sporocarps after a burn [68,69], which would make them poised to colonize when the fire subsides. ...
Wildfires have continued to increase in frequency and severity in Southern California due in part to climate change. To gain a further understanding of microbial soil communities’ response to fire and functions that may enhance post-wildfire resilience, soil fungal and bacterial microbiomes were studied from different wildfire areas in the Gold Creek Preserve within the Angeles National Forest using 16S, FITS, 18S, 12S, PITS, and COI amplicon sequencing. Sequencing datasets from December 2020 and June 2021 samplings were analyzed using QIIME2, ranacapa, stats, vcd, EZBioCloud, and mixomics. Significant differences were found among bacterial and fungal taxa associated with different fire areas in the Gold Creek Preserve. There was evidence of seasonal shifts in the alpha diversity of the bacterial communities. In the sparse partial least squares analysis, there were strong associations (r > 0.8) between longitude, elevation, and a defined cluster of Amplicon Sequence Variants (ASVs). The Chi-square test revealed differences in fungi–bacteria (F:B) proportions between different trails (p = 2 × 10−16). sPLS results focused on a cluster of Green Trail samples with high elevation and longitude. Analysis revealed the cluster included the post-fire pioneer fungi Pyronema and Tremella. Chlorellales algae and possibly pathogenic Fusarium sequences were elevated. Bacterivorous Corallococcus, which secretes antimicrobials, and bacterivorous flagellate Spumella were associated with the cluster. There was functional redundancy in clusters that were differently composed but shared similar ecological functions. These results implied a set of traits for post-fire resiliency. These included photo-autotrophy, mineralization of pyrolyzed organic matter and aromatic/oily compounds, potential pathogenicity and parasitism, antimicrobials, and N-metabolism.
... odini, G. turficola) differ by only 2-3 bp in ITS region (see the "5" section), some morphological characters and ecological preferences (Høiland 1990;Moser et al. 2001;Holec 2012;Khan et al. 2017). However, the delimitation of G. decipiens is still unclear, which is documented, e.g., by difference in sequences of European and USA collections bearing this name (Hughes et al. 2020). There is also a sequenced collection of G. decipiens from Japan (Kasuya and Hosaka 2017). ...
Gymnopilus stabilis (Fungi, Agaricales), a very rare species of unclear identity described from Europe, was revised using morphological data and phylogenetic affiliations derived from ITS rDNA sequences. Collections from the Czech Republic, Germany, and Russia plus reference vouchers of related or similar species were studied. Gymnopilus stabilis sequences form a moderately supported but distinct clade in a lineage containing G. swaticus and a Gymnopilus collection of unknown identity from the USA. The G. stabilis lineage is sister to the G. sapineus clade, a group of species with low divergence in ITS rDNA region. This whole, well-supported group is clearly distinct from the lineage of G. penetrans, a species with which G. stabilis has sometimes been confused. Morphological characters of a recent, well-documented Czech collection of G. stabilis are described in detail and compared with the German and Russian ones. A literature review confirms that the taxonomic concept of G. stabilis has not substantially changed over time. In order to consolidate the taxonomy and nomenclature of this enigmatic species, lectotype and epitype are designated here for G. stabilis, and key diagnostic characters are highlighted. The taxonomic concepts of G. sapineus and G. penetrans are further specified based on well-documented, sequenced reference collections. Moreover, the position of G. hybridus is briefly discussed. The identification of several GenBank entries of G. stabilis, G. sapineus, and G. penetrans is revised in order to fix species concepts in this fascinating group of wood-decaying fungi.
... including the type (from Czechia) and, in addition to the above, sequence data from Pakistan. It appears that H. nanum most commonly occurs on sandy soil, in post-fire regenerating boreal and temperate, coniferous forests Hughes et al. 2020). ...
Here, we present the results of studies of Japanese Hebeloma collections. The four species described by Imai as Hebeloma ( H . fimicola , H . helvolescens , H . humosum , and H . tomoeae ) are not from the genus Hebeloma , but are members of Agrocybe , Homophron , or Pholiota . Recombinations are made. Hebeloma crustuliniforme f. microspermum , described by Hongo, is a synonym of H . nanum . Three species of Hebeloma are described as new to science, all currently known only from Japan. Two of these species, H . asperosporum and H . cinnamomeum , are members of H . sect. Denudata while the third species H . citrisporum belongs to H. sect. Velutipes . Japanese records of H . cavipes , H . eburneum , H . hygrophilum , H . subtortum , and H . velutipes are validated. In total, fifteen species of Hebeloma are confirmed from Japan; this is compared with previous checklists.
... After manual editing in BioEdit (Hall, 1999), assembled sequences were submitted to GenBank. For the phylogenetic analysis (ITS rDNA), various Thelephora sequences were retrieved from GenBank and UNITE databases (Kõljalg et al., 2020), based on taxonomic papers on the genus (Sha et al., 2008;Tedersoo et al., 2014;Ramírez-López et al., 2015;Vizzini et al., 2016;Wang et al., 2017a;Li et al., 2020a) and other works (Smith et al., 2007;Kõljalg et al., 2000;Hughes et al., 2020). Sequences of insufficient length were omitted. ...
Fruit-bodies of six Thelephora species (Fungi, Basidiomycota, Thelephoraceae) were analyzed for their trace element concentrations. In Thelephora penicillata, extremely high concentrations of Cd and As were found, followed by highly elevated concentrations of Cu and Zn. The highest accumulation ability was found for Cd with a mean concentration of 1.17 ± 0.37 g kg⁻¹ (dry mass) in fruit-bodies collected from 20 unpolluted sites; the mean As concentration was 0.878 ± 0.242 g kg⁻¹. Furthermore, striking accumulation of Se (923 ± 28 mg kg⁻¹) was found in one sample of T. vialis and elevated concentrations of S were detected in T. palmata (19.6 ± 5.9 g kg⁻¹). The analyzed Thelephora species were sequenced and, based on the Maximum Likelihood phylogenetic analysis (ITS rDNA) of the genus, possible other Thelephora (hyper)accumulators were predicted on the basis of their phylogenetic relationship with the discovered (hyper)accumulators. The striking ability of T. penicillata to accumulate simultaneously Cd, As, Cu, and Zn has no parallel in the Fungal Kingdom and raises the question of a biological importance of metal(loid) hyperaccumulation in mushrooms.
... In contrast to our findings for AM fungi, bulk soil fungal and bacterial communities as a whole shifted in both diversity and composition following the Chimney Tops 2 fire (same fire, different sites; Brown et al., 2019). Similarly, pyrophilus fungal taxa (e.g., Sphaerosporella spp.) appeared within 6 months following the GSMNP fire and persisted for at least the next 2 years (Hughes et al., 2020a(Hughes et al., , 2020b. Shifts in microbial composition belowground can have functional consequences and even feedback to future fires, when decomposition slows (Hopkins et al., 2020). ...
Wildfires are increasing in frequency and intensity as drier and warmer climates increase plant detrital fuel loads. At the same time, increases in urbanization position 9% of fire-prone land within the United States at the wildland–urban interface. While rarely studied, the compounded effects of urbanization and wildfires may have unknown synergistically negative effects on ecosystems. Previous studies at the wildland–urban interface often focus on aboveground plant communities, but belowground ecosystems may also be affected by this double disturbance. In particular, it is unclear how much fire and urbanization independently or interactively affect nutritional symbioses such as those between arbuscular mycorrhizal (AM) fungi and the majority of terrestrial plants. In November 2016, extreme drought conditions and long-term fire suppression combined to create a wildfire within the Great Smoky Mountains National Park and the neighboring exurban city of Gatlinburg, TN. To understand how the double disturbance of urbanization and fire affected AM fungal communities, we collected fine roots from the 5 dominant understory species in September 2018 at each of 18 sites spanning 3 burn severities in both exurban and natural sites. Despite large variation in burn severity, plant species identity had the largest influence on AM fungi. AM fungal colonization, richness, and composition all varied most among plant species. Fire and urbanization did influence some AM fungal metrics; colonization was lower in burned sites and composition was more variable among exurban locations. There were no interactions among burn severity and urbanization on AM fungi. Our results point to the large influence of plant species identity structuring this obligate nutritional symbiosis regardless of disturbance regime. Therefore, the majority of AM fungal taxa may be buffered from fire-induced ecosystem changes if plant community composition largely remains intact, plant species life history traits allow for AM fungal persistence after fire disturbance, and/or nearby undisturbed habitat can act as an inoculum source for recolonization following fires. Thus, it is critical to maintain natural, undisturbed habitats interspersed within the wildland–urban interface.
... Forest communities within southern Appalachia are largely structured by elevational gradients (Day and Monk 1974;Lorimer 1980;White et al., 2018) and plant community changes with elevation are often mirrored by patterns in fungal diversity (Veach et al., 2018), lichen diversity (Allen and Lendemer, 2016), and biogeochemical patterns of nitrogen and carbon cycling (Knoepp and Swank 1998;Knoepp et al., 2008Knoepp et al., , 2018. Fungal communities are generally considered hyperdiverse in these mountains as evidenced by numerous investigations using molecular and morphological datasets (Hesler 1937;Walker et al. 2005Walker et al. , 2008Baird et al., 2014;Veach et al., 2018;Barney et al., 2020;Hughes et al., 2020). Despite numerous investigations into fungal diversity in southern Appalachia, basic ecological knowledge is often lacking. ...
The southern Appalachian Mountains in the United States are heavily forested with diverse forest types and vegetation structures and is generally considered a biodiversity hotspot. Fungal sporocarp investigations in this region are not new, but multi-year interseasonal investigations into sporocarp community patterns are rare. Using a 4-year (2014-2017) repeated monthly macrofungal sampling dataset (May-October) across an eleva-tional gradient (associated with vegetation structure), we queried community, seasonal, and guild community ecological patterns. In doing so, we (1) demonstrate a temporal disconnect between saprobic and ectomycor-rhizal taxa, (2) describe increased community variability across the growing season, and (3) explore individual taxa occurrences across seasons and examine co-occurrence patterns between taxa and guilds. Further, we explore congruence between sporocarp and metabarcoding fungal datasets and advocate the utility of both sampling schemes.
... Burn morels are a non-timber product with high economic value, particularly in western North America (Alexander et al., 2002;Wurtz et al., 2005;McFarlane et al., 2005). However, a whole cadre of pyrophilous fungi has been documented on burned soil and charred wood after forest fires (Petersen, 1970;Claridge et al., 2009;Hughes et al., 2020). Many, such as Pyronema, Anthracobia and Geopyxis species, are cup fungi (Ascomycota) that fruit prolifically and consistently after fire (Greene et al., 2010;Bruns et al., 2020;Raudabaugh et al., 2020). ...
Forest fire is an important occurrence in western landscapes where it helps drive ecosystem processes, and prescribed fire is a common forest management strategy. An initial consequence of fire is burned ground with reduced biodiversity. Numerous studies have documented how forests recover after fire in terms of plant regeneration and animal colonization, but little is known of the ecological roles fungi play in this process. Pyrophilous (burn-loving) fungi are documented to reliably produce large fruitings and copious mycelium on burns after fire in North America, Europe, Asia, and Australia. We hypothesize that pyrophilous fungi help bind and stabilize soil after forest fires via their extensive mycelial network. Three pyrophilous fungi, Geopyxis carbonaria, Pyronema omphalodes, and Morchella septimelata were tested for their ability to aggregate burned soil. The fungi were isolated from burn sites, grown in vitro, and inoculated onto sterilized soil from a natural burn. The ability of each species to aggregate soil in comparison to non-inoculated controls was assessed after 10, 20, 30, and 40 days, using a wet sieve aggregate stability test. All three fungi increased soil aggregation after 10 days, and this increase was maintained for the 40-day period. The burned soil was up to 30% more aggregated when a fungus was present; results provide the first direct evidence that pyrophilous fungi aggregate burn soil. This further implies that these fungi play a role in reducing soil erosion and enhancing soil moisture soon after fire in burned forests. Pyrophilous fungi also decompose charred material, sequester carbon, and capture transient nitrogen pulses after fire. This overlooked group of fungi may be critical in enhancing conditions for plant regeneration after forest fire at an early stage in recovery. Consideration should be given to avoiding or delaying restoration activities that disturb this natural process, especially those that contribute to soil compaction, during early post-fire recovery when these fungi are proliferating.
... Similar to previous studies (Bruns et al., 2019;Cairney and Bastias, 2007;Fujimura et al., 2005;Reazin et al., 2016;Smith et al., 2004;Visser, 1995;Yang et al., 2020), we observed a shift in dominance in the burned sites, from Basidiomycete-to Ascomycete-dominated. The emergence and dominance of Chromelosporium and Pustularia, both rare in the unburned sites, supports previous findings that cryptic and rare taxa tend to dominate the burned environment (Bruns et al., 2019;Fujimura et al., 2005;Horton and Bruns, 1998;Hughes et al., 2020;Smith et al., 2004;Visser, 1995). The most dominant species in both the unburned and burned environments was W. rehmii, a well-known symbiont of P. ponderosa (Fujimura et al., 2005;Glassman et al., 2015;Smith et al., 2005) that is likely to be resilient to high-severity fires. ...
Ponderosa pine (Pinus ponderosa) forests are increasingly experiencing high-severity, stand-replacing fires. Whereas alterations to aboveground ecosystems have been extensively studied, little is known about soil fungal responses in fire-adapted ecosystems. We implement a chronosequence of four different fires that varied in time since fire, 2 years (2015) to 11 years (2006) and contained stands of high severity burned P. ponderosa in eastern Washington and compared their soil fungal communities to adjacent unburned plots. Using Illumina Miseq (ITS1), we examined changes in soil nutrients, drivers of species richness for ectomycorrhizal (plant symbionts) and saprobic (decomposers) fungi, community shifts, and post-fire fungal succession in burned and unburned plots. Ectomycorrhizal richness was 43.4% and saprobic richness 12.2% lower in the burned plots, leading to long-term alterations to the fungal communities that did not return to unburned levels, even after 11 years. Differences in the post-fire fungal community were driven by pyrophilous, “fire-loving” fungi, including the ectomycorrhizal Ascomycete genera Pustularia and Wilcoxina, and the saprobic Basidiomycete genus Geminibasidium. Ectomycorrhizal and saprobic fungi were intimately linked to the soil environment: depth of the organic matter, total carbon, total nitrogen, and their interaction with fire predicted ectomycorrhizal richness. Whereas total carbon, time since fire, treatment, and the interaction between time since fire and treatment predicted saprobic richness. We conclude that high-severity wildfires lead to lower ectomycorrhizal richness and significantly altered ectomycorrhizal and saprobic communities in fire-adapted ecosystems, selecting resilient and fire-adapted species, such as W. rehmii and Geminibasidium sp., thus initiating post-fire succession.
... Recent wildfires have shown that fire tolerant taxa are already present in long unburned ecosystems and increase in abundance following fire Smith et al. 2016;Hughes et al. 2020). Improving our understanding between microbial community structure, seasonality, and function in fire recurrent ecosystems may therefore provide a more generalizable model for predicting future changes with more frequent fires. ...
Fire alters microbial community composition, and is expected to increase in frequency due to climate change. Testing whether microbes in different ecosystems will respond similarly to increased fire disturbance is difficult though, because fires are often unpredictable and hard to manage. Fire recurrent or pyrophilic ecosystems, however, may be useful models for testing the effects of frequent disturbance on microbes. We hypothesized that across pyrophilic ecosystems, fire would drive similar alterations to fungal communities including altering seasonal community dynamics. We tested fire's effects on fungal communities in two pyrophilic ecosystems, a Longleaf pine savanna and tallgrass prairie. Fire caused similar fungal community shifts including a) driving immediate changes that favored taxa able to survive fire and take advantage of post-fire environments, and b) altering seasonal trajectories due to fire-associated changes to soil nutrient availability. This suggests that fire has predictable effects on fungal community structure and intra-annual community dynamics in pyrophilic ecosystems, and that these changes could significantly alter fungal function. Parallel fire responses in these key microbes may also suggest that recurrent fires drive convergent changes across ecosystems, including less fire frequented systems that may start burning more often.
A root disease in plantations of Pinus radiata and Pinus pinaster , where trees died in distinct patches, was present in the Western Cape province of South Africa during the 1970s and 1980s. Phytophthora cinnamomi was initially believed to be the cause, but the disease was later ascribed to the insect-associated fungus Leptographium serpens , a fungal species residing in the Ophiostomatales. Doubt regarding the cause of the disease was raised in a later study due to the fact that most Leptographium spp., particularly those that colonise ray parenchyma tissues, which is the case for L. serpens , are not typically primary disease agents. In this study, cultures of an unidentified sterile fungus collected from the dying trees were revived and identified using DNA sequencing methods, which were not available when the disease was first studied. These cultures were identified as the pyrophillic pathogen Rhizina undulata , well-known to cause patch death of conifers in South Africa and elsewhere in the world. While the patches of dying trees no longer exist and the disease cannot be newly studied, it is most likely that the tree death originally thought to be caused by L. serpens was due primarily to R. undulata . The study provides a vivid example of the value of preserving cultures of fungi for later study and the power of modern techniques to identify fungal pathogens.
Prescribed fire is a critical strategy for mitigating the effects of catastrophic wildfires. While the above-ground response to fire has been well-documented, fewer studies have addressed the effect of prescribed fire on soil microorganisms. To understand how soil microbial communities respond to prescribed fire, we sampled four plots at a high temporal resolution (two burned, two controls), for 17 months, in a mixed conifer forest in northern California, USA. Using amplicon sequencing, we found that prescribed fire significantly altered both fungal and bacterial community structure. We found that most differentially abundant fungal taxa had a positive fold-change, while differentially abundant bacterial taxa generally had a negative fold-change. We tested the null hypothesis that these communities assembled due to neutral processes (i.e., drift and/or dispersal), finding that >90% of taxa fit this neutral prediction. However, a dynamic sub-community composed of burn-associated indicator taxa that were positively differentially abundant was enriched for non-neutral amplicon sequence variants, suggesting assembly via deterministic processes. In synthesizing these results, we identified 15 pyrophilous taxa with a significant and positive response to prescribed burns. Together, these results lay the foundation for building a process-driven understanding of microbial community assembly in the context of the classical disturbance regime of fire.
The rise in wildfire frequency and severity across the globe has increased interest in secondary succession. However, despite the role of soil microbial communities in controlling biogeochemical cycling and their role in the regeneration of post-fire vegetation, the lack of measurements immediately post-fire and at high temporal resolution has limited understanding of microbial secondary succession. To fill this knowledge gap, we sampled soils at 17, 25, 34, 67, 95, 131, 187, 286, and 376 days after a southern California wildfire in fire-adapted chaparral shrublands. We assessed bacterial and fungal biomass with qPCR of 16S and 18S and richness and composition with Illumina MiSeq sequencing of 16S and ITS2 amplicons. Fire severely reduced bacterial biomass by 47%, bacterial richness by 46%, fungal biomass by 86%, and fungal richness by 68%. The burned bacterial and fungal communities experienced rapid succession, with 5-6 compositional turnover periods. Analogous to plants, turnover was driven by "fire-loving" pyrophilous microbes, many of which have been previously found in forests worldwide, and changed markedly in abundance over time. Fungal secondary succession was initiated by the Basidiomycete yeast Geminibasidium, which traded off against the filamentous Ascomycetes Pyronema, Aspergillus, and Penicillium. For bacteria, the Proteobacteria Massilia dominated all year, but the Firmicute Bacillus and Proteobacteria Noviherbaspirillum increased in abundance over time. Our high-resolution temporal sampling allowed us to capture post-fire microbial secondary successional dynamics and suggest that putative tradeoffs in thermotolerance, colonization, and competition among dominant pyrophilous microbes control microbial succession with possible implications for ecosystem function.
Understanding which fungi exhibit certain ecological traits, such as habitat, host, or substrate associations, and knowing how these traits change across space and time can provide invaluable insight into the roles fungi play in their respective ecosystems. Archived sporocarp data, such as the collection and observation records accessible through the Mycology Collections Portal (MyCoPortal), are well suited for trait investigations, since these records circumvent the need for field work, are geographically and temporally diverse, and often have detailed and trait-relevant environmental metadata. However, there are inefficiencies and inadequacies in the MyCoPortal online interface that affect data set generation and trait searching, and many of the available records have outdated or misspelled taxon names as well as misspelled location names. Thus, we created the r package fungarium, which enables the efficient download of complete MyCoPortal data sets from within the R environment, enhances the identification of trait-relevant records, confirms or updates taxon names while also accounting for spelling errors, and fixes misspelled location names. Utilizing this package and MyCoPortal data, we demonstrated methods for analyzing taxonomic, geographic, and temporal patterns in ecological traits, using fire association as an example. We found that fire association, which was quantified via fire-associated enrichment factors (fire-associated records/total records), differed substantially between taxa, and these differences were qualitatively supported by existing literature, as hypothesized. Sampling bias within the MyCoPortal data and limitations of the burned acreage data set used (i.e., Monitoring Trends in Burn Severity) were identified as confounding factors in our geographic and temporal analyses, as evidenced by the unexpected lack of correlation between fire association and burned acreage on county and year bases. However, both confounding factors likely depend on the trait analyzed and external data set used. Overall, the fungarium package and associated methods presented here effectively enable the use of archived sporocarp data for future ecological trait studies.
Wildfires represent a fundamental and profound disturbance in many ecosystems, and their frequency and severity are increasing in many regions of the world. Fire affects soil by removing carbon in the form of CO 2 and transforming remaining surface carbon into pyrolyzed organic material (PyOM). Fires also generate substantial necromass at depths where the heat kills soil organisms but does not catalyze the formation of PyOM. Pyronema species strongly dominate soil fungal communities within weeks to months after fire. However, the carbon pool (i.e. necromass or PyOM) that fuels their rise in abundance is unknown. We used a Pyronema domesticum isolate from the catastrophic 2013 Rim Fire (CA, USA) to ask if P. domesticum is capable of metabolizing PyOM. P. domesticum grew readily on agar media where the sole carbon source was PyOM (specifically, pine wood PyOM produced at 750 °C). Using RNAseq, we investigated the response of P. domesticum to PyOM and observed a comprehensive induction of genes involved in the metabolism and mineralization of aromatic compounds, typical of those found in PyOM. Lastly, we used ¹³ C-labeled 750 °C PyOM to demonstrate that P. domesticum is capable of mineralizing PyOM to CO 2 . Collectively, our results indicate a robust potential for P. domesticum to liberate carbon from PyOM in post-fire ecosystems and return it to the bioavailable carbon pool.
IMPORTANCE
Fires are increasing in frequency and severity in many regions across the world. Thus, it’s critically important to understand how our ecosystems respond to inform restoration and recovery efforts. Fire transforms the soil, removing many nutrients while leaving behind both nutritious necromass and complex pyrolyzed organic matter, which is often recalcitrant. Filamentous fungi of the genus Pyronema strongly dominate soil fungal communities soon after fire. While Pyronema are key pioneer species in post-fire environments, the nutrient source that fuels their rise in abundance is unknown. In this manuscript, we used a P. domesticum isolate from the catastrophic 2013 Rim Fire (CA, USA) to demonstrate that P. domesticum metabolizes pyrolyzed organic material, effectively liberating this complex pyrolyzed carbon and returning it to the bioavailable carbon pool. The success of Pyronema in post-fire ecosystems has the potential to kick-start growth of other organisms and influence the entire trajectory of post-fire recovery.
The genus Gymnopilus (Fungi, Agaricales, Cortinariaceae) was monographically studied. The study is based on a thorough observation of macro- and microcharacters using numerous personal and herbarium collections from the Czech Republic. Selected
collections from Austria, Bulgaria, Croatia, Finland, Germany, Italy, Poland, Romania, Russia, Slovakia, Sweden, and Ukraine were also used for a better understanding of the species concept. In each species, macro- and microcharacters, fructification period, substrates, relation to vegetation and altitude, and distribution are evaluated in detail. Taxonomic remarks on all species are added and contemporary literature on Gymnopilus is summarised. A key for the identification of species growing in the Czech Republic is provided. The following species were recognised in this area: Gymnopilus spectabilis (taxon with robust fruitbodies), G. bellulus, G. josserandii, G. flavus, G. fulgens, G. decipiens, G. penetrans (= G. hybridus), G. sapineus and G. picreus. The following names commonly used in contemporary literature are either not recognised in the Czech
Republic or considered doubtful or hardly interpretable taxa: Gymnopilus stabilis, G. junonius s. str. (taxon with slender fruitbodies), and G. liquiritiae. Line drawings and photographs of important microcharacters are provided for each species. Colour photographs of 6 species are added.
Diverse plant‐associated fungi are thought to have symbiotrophic and saprotrophic states because they can be isolated from both dead and living plant tissues. However, such tissues often are separated in time and space, and fungal activity at various stages of plant senescence is rarely assessed directly in fungal community studies.
We used fungal ribosomal RNA metatranscriptomics to detect active fungal communities across a natural senescence gradient within wild‐collected gametophytes of Dicranum scoparium (Bryophyta) to understand the distribution of active fungal communities in adjacent living, senescing and dead tissues.
Ascomycota were active in all tissues across the senescence gradient. By contrast, Basidiomycota were prevalent and active in senescing and dead tissues. Several fungi were detected as active in living and dead tissues, suggesting their capacity for multi‐trophy. Differences in community assembly detected by metatranscriptomics were echoed by amplicon sequencing of cDNA and compared to culture‐based inferences and observation of fungal fruit bodies in the field.
The combination of amplicon sequencing of cDNA and metatranscriptomics is promising for studying symbiotic systems with complex microbial diversity, allowing for the simultaneous detection of their presence and activity.
Two species of Laccaria discovered in relicts of Fagus grandifolia var. mexicana forests in eastern Mexico are described based on the macro- and micromorphological features, and their identity supported by molecular analysis of the internal transcribed spacer (ITS) and large subunit (LSU) of the ribosomal RNA gene. The phylogeny obtained here showed that one of the Mexican species is nested in an exclusive clade which in combination with its striking morphological features, infers that it represents a new species, while the other species is placed as a member in the Laccaria trichodermophora clade. This is the first report in Mexico of Laccaria with Fagus grandifolia var. mexicana trees, with which the reported species may form ectomycorrhizal association. Descriptions are accompanied with illustrations of macro- and micromorphological characters and a discussion of related taxa are presented.
The paper presents 32 species of macrofungi new to the Kampinos National Park, found during the studies on fire-damaged areas after the forest fires in 2015. Three species new to Poland were described and illustrated ( Calycellina leucella , Exobasidium juelianum , and Gymnopilus decipiens ). Four species from Polish red list of macrofungi have been recorded in the Kampinos National Park for the first time: Botryobasidium vagum (R), Geastrum coronatum (V), Helicogloea farinacea (E), Inonotus cuticularis (R). During the current studies 17 pyrophilous species new to the Kampinos National Park were found.
The aim of the present work was to investigate the potential for territorial and dispersive clonality in natural populations of the postfire root rot ascomycete Rhizina undulata. Population studies based on vegetative compatibility tests were done with strains isolated from individual sporocarps at five burned sites in three different localities (separated by 20–40 km) in the Curronian Spit of western Lithuania. Among a total of 103 strains, the tests identified 14 distinct vegetative compatibility groups (VCGs) of R. undulata, 13 of which were represented by 2–48 strains and three were encountered at 2–4 different sites. Occurrence on spatially separated sites of the same VCG of the fungus indicated a presence of dispersive clonality in R. undulata populations. On a local scale clusters of vegetative compatible sporocarps usually occupied discrete territories, implying territorial clonality. The two largest VCGs covered areas up to 7 and 3 m across. The results show that both dispersive and territorial clones are characteristics of natural populations of the fungus.
The importance of fire in shaping Appalachian vegetation has become
increasingly apparent over the last 25 years. This period has seen declines in
oak (Quercus) and pine (Pinus) forests and other fire-dependent ecosystems,
which in the near-exclusion of fire are being replaced by fire-sensitive
mesophytic vegetation. These vegetation changes imply that Appalachian
vegetation had developed under a history of burning before the fire-exclusion
era, a possibility that has motivated investigations of Appalachian fire history
using proxy evidence. Here we synthesize those investigations to obtain an
up-to-date portrayal of Appalachian fire history. We organize the report by data
type, beginning with studies of high-resolution data on recent fires to provide
a context for interpreting the lower-resolution proxy data. Each proxy is
addressed in a subsequent chapter, beginning with witness trees and continuing
to fire-scarred trees, stand age structure, and soil and sediment charcoal. Taken
together, these proxies portray frequent burning in the past. Fires had occurred
at short intervals (a few years) for centuries before the fire-exclusion era.
Indeed, burning has played an important ecological role for millennia. Fires
were especially common and spatially extensive on landscapes with large
expanses of oak and pine forest, notably in the Ridge and Valley province
and the Blue Ridge Mountains. Burning favored oak and pine at the expense
of mesophytic competitors, but fire exclusion has enabled mesophytic plants
to expand from fire-sheltered sites onto dry slopes that formerly supported
pyrogenic vegetation. These changes underscore the need to restore fire-dependent
ecosystems.
Background
Fungal endophytes inhabit symptomless, living tissues of all major plant lineages to form one of earth’s most prevalent groups of symbionts. Many reproduce from senesced and/or decomposing leaves and can produce extracellular leaf-degrading enzymes, blurring the line between symbiotrophy and saprotrophy. To better understand the endophyte–saprotroph continuum we compared fungal communities and functional traits of focal strains isolated from living leaves to those isolated from leaves after senescence and decomposition, with a focus on foliage of woody plants in five biogeographic provinces ranging from tundra to subtropical scrub forest.
Methods
We cultured fungi from the interior of surface-sterilized leaves that were living at the time of sampling (i.e., endophytes), leaves that were dead and were retained in plant canopies (dead leaf fungi, DLF), and fallen leaves (leaf litter fungi, LLF) from 3–4 species of woody plants in each of five sites in North America. Our sampling encompassed 18 plant species representing two families of Pinophyta and five families of Angiospermae. Diversity and composition of fungal communities within and among leaf life stages, hosts, and sites were compared using ITS-partial LSU rDNA data. We evaluated substrate use and enzyme activity by a subset of fungi isolated only from living tissues vs. fungi isolated only from non-living leaves.
Results
Across the diverse biomes and plant taxa surveyed here, culturable fungi from living leaves were isolated less frequently and were less diverse than those isolated from non-living leaves. Fungal communities in living leaves also differed detectably in composition from communities in dead leaves and leaf litter within focal sites and host taxa, regardless of differential weighting of rare and abundant fungi. All focal isolates grew on cellulose, lignin, and pectin as sole carbon sources, but none displayed ligninolytic or pectinolytic activity in vitro . Cellulolytic activity differed among fungal classes. Within Dothideomycetes, activity differed significantly between fungi from living vs. non-living leaves, but such differences were not observed in Sordariomycetes.
Discussion
Although some fungi with endophytic life stages clearly persist for periods of time in leaves after senescence and incorporation into leaf litter, our sampling across diverse biomes and host lineages detected consistent differences between fungal assemblages in living vs. non-living leaves, reflecting incursion by fungi from the leaf exterior after leaf death and as leaves begin to decompose. However, fungi found only in living leaves do not differ consistently in cellulolytic activity from those fungi detected thus far only in dead leaves. Future analyses should consider Basidiomycota in addition to the Ascomycota fungi evaluated here, and should explore more dimensions of functional traits and persistence to further define the endophytism-to-saprotrophy continuum.
Fire is the dominant factor affecting C and N losses from the semiarid forests of the eastern Sierra Nevada. As prescription fire becomes a best management practice, it is critical to develop an estimate of these fluxes. The objectives of this study were (i) to test and refine methods to estimate the volatilized C and N losses from the forest floor following fire, (ii) to investigate the interactions between O-horizon temperature and nutrient loss, and (iii) to assess measured N losses in the context of atmospheric N deposition, leaching, and N fixation. The quantities of C and N volatilized from the forest floor by prescription fire in the Sierra Nevada were measured using two different field-based methods: weight loss estimation and Calelement ratio determination. Three sites were included in the study: Marlene, Sawtooth and Spooner. The weight method indicated C losses of 6.12, 7.39, and 17.8 Mg C ha -1 at the Sawtooth, Marlene, and Spooner sites, respectively. The ratio method indicated comparable C losses from the Sawtooth (6 Mg C ha -1 ) site, but greater losses at Marlene (16 Mg C ha -1 ) and Spooner (24 Mg C ha -1 ) sites. The weight method indicated N losses of 56.2, 60.8, and 362 kg N ha -1 , at the Sawtooth, Marlene, and Spooner sites, respectively. The ratio method indicated comparable N losses of 59.9 kg N ha -1 at the Sawtooth site, but considerably greater losses at Marlene (243 kg N ha -1 ), and Spooner (524 kg N ha -1 ) sites. The Ca-element method was preferred because of minimal needs for preburn sampling. Regardless of method, the estimated losses were significant, particularly for N, compared with deposition and leaching rates. Volatilization will represent the major mechanism for N loss from forest ecosystems of this region subjected to prescribed fire.
After severe wildfires, pine recovery depends on ectomycorrhizal (ECM) fungal spores surviving and serving as partners for regenerating forest trees. We took advantage of a large, severe natural forest fire that burned our long-term study plots to test the response of ECM fungi to fire. We sampled the ECM spore bank using pine seedling bioassays and high-throughput sequencing before and after the California Rim Fire. We found that ECM spore bank fungi survived the fire and dominated the colonization of in situ and bioassay seedlings, but there were specific fire adapted fungi such as Rhizopogon olivaceotinctus that increased in abundance after the fire. The frequency of ECM fungal species colonizing pre-fire bioassay seedlings, post-fire bioassay seedlings and in situ seedlings were strongly positively correlated. However, fire reduced the ECM spore bank richness by eliminating some of the rare species, and the density of the spore bank was reduced as evidenced by a larger number of soil samples that yielded uncolonized seedlings. Our results show that although there is a reduction in ECM inoculum, the ECM spore bank community largely remains intact, even after a high-intensity fire. We used advanced techniques for data quality control with Illumina and found consistent results among varying methods. Furthermore, simple greenhouse bioassays can be used to determine which fungi will colonize after fires. Similar to plant seed banks, a specific suite of ruderal, spore bank fungi take advantage of open niche space after fires.The ISME Journal advance online publication, 16 October 2015; doi:10.1038/ismej.2015.182.
In this study, Geoglossum umbratile Sacc. and Peziza lobulata (Velen.) Svrček were reported for the first time
in Turkey. Their descriptions, morphological characteristics, and photographs are presented.
Forest fire is a common natural disturbance in forested ecosystems and has a large impact on the microbial communities in forest soils. The response of soil fungal communities to forest fire is poorly documented. Here, we investigated the fungal community structure and function across a 152-year boreal forest fire chronosequence using high-throughput sequencing of the Internal Transcribed Spacer (ITS2) region and a functional gene array (GeoChip). Our results demonstrate that the boreal forest soil fungal community was most diverse soon after the fire disturbance and declined over time. The differences among the fungal community were explained by changes in the abundance of Basidiomycetes and Ascomycetes. Ectomycorrhizal (ECM) fungi contributed to the increase of basidiomycete abundance over time with the OTUs representing the genera Cortinarius and Piloderma dominating. Hierarchical cluster analysis by using gene signal intensity revealed that the sites with different fire histories formed separate clusters suggesting differences in the potential to maintain essential biogeochemical soil processes. The site with the highest biological diversity had also the most diverse genes. The genes involved in the organic matter degradation in the mature forest, where ECM fungi were the most abundant were as common as in the youngest site where saprotrophic fungi had relatively higher abundance. This study provides insight into the impact of fire disturbance on soil fungal community dynamics.
Copyright © 2015, American Society for Microbiology. All Rights Reserved.
Based on traditional morphology, sequence data, and phylogenetic analyses, 18 new species are here described: Coprinellus christianopolitanus, Coprinopsis musae, C. udicola, Psathyrella arenosa, P. carminei, P. fennoscandica, P. ichnusae, P. lilliputana, P. lyckebodensis, P. madida, P. rybergii, P. sabuletorum, P. scanica, P. siccophila, P. stridvallii, P. sublatispora, P. vesterholtii, and Typhrasa nanispora. Psathyrella hololanigera and P. tenera are reported as new to Europe and P. parva as new to the Nordic countries. A four-gene dataset on Psathyrellaceae were analyzed by Maximum Parsimony, Maximum Likelihood, and Bayesian methods. Constraint analyses were performed to determine limits of /Psathyrella, and to evaluate whether the /Coprinellus, /cordisporus, and /gossypina clades could be regarded with confidence as monophyletic clades outside of the clade /Psathyrella. This was not unambiguously supported. Based on the phylogenetic results, Kauffmania is proposed as a monotypic genus for the species P. larga and Typhrasa for P. gossypina and the new described species T. nanispora. The genus Homophron is formally validated and three combinations are proposed: H. spadiceum, H. cernuum, and H. camptopodum. The genus Cystoagaricus Singer is emended and the following new combinations are proposed: C. hirtosquamulosus, C. squarrosiceps, C. olivaceogriseus, and C. silvestris. Neotypes have been selected for seven species described by Fries, and ITS sequence data for these were generated. The following new combinations are proposed: Coprinopsis canoceps, C. cineraria, C. melanthina, C. submicrospora, C. uliginicola, and Typhrasa gossypina. Brief comments are given to other species of interest. Psathyrella ornatispora were found not to belong in Psathyrellaceae. A key to 106 psathyrelloid species in Northern Europe is provided.
We explored evolutionary relationships within the Lyophyllaceae by combining sequence data from six loci. The most likely phylogram led us to reconsider the Lyophyllaceae classification with the recognition of two new genera (Myochromella and Sagaranella) based on ecological and/or morphological distinctiveness. Lyophyllaceae are ecologically highly diversified and our phylogeny suggests that four to five ecological transitions from free-living to parasitic or mutualistic lifestyles have occurred within the family. Due to moderate phylogenetic support recovered for several relationships within that clade and due to the uncertainty about the ecological strategy adopted by five of the sampled species, three out of these transitions could be unequivocally reconstructed suggesting that saprotrophy is plesiomorphic for Lyophyllaceae. Significant differences in rates of molecular evolution were detected among taxa. These differences are not associated with ecological transitions throughout the Lyophyllaceae, however, within each of the major clades identified in the family, taxa of different ecological strategies show an overall tendency to evolve at different speeds at the molecular level.
In hot deserts, plants cope with aridity, high temperatures, and nutrient-poor soils with morphological and biochemical adaptations that encompass intimate microbial symbioses. Whereas the root microbiomes of arid-land plants have received increasing attention, factors influencing assemblages of symbionts in aboveground tissues have not been evaluated for many woody plants that flourish in desert environments. We evaluated the diversity, host affiliations, and distributions of endophytic fungi associated with photosynthetic tissues of desert trees and shrubs, focusing on nonsucculent woody plants in the species-rich Sonoran Desert. To inform our strength of inference, we evaluated the effects of two different nutrient media, incubation temperatures, and collection seasons on the apparent structure of endophyte assemblages. Analysis of >22,000 tissue segments revealed that endophytes were isolated four times more frequently from photosynthetic stems than leaves. Isolation frequency was lower than expected given the latitude of the study region and varied among species a function of sampling site and abiotic factors. However, endophytes were very species-rich and phylogenetically diverse, consistent with less arid sites of a similar latitudinal position. Community composition differed among host species, but not as a function of tissue type, sampling site, sampling month, or exposure. Estimates of abundance, diversity, and composition were not influenced by isolation medium or incubation temperature. Phylogenetic analyses of the most commonly isolated genus (Preussia) revealed multiple evolutionary origins of desert-plant endophytism and little phylogenetic structure with regard to seasonality, tissue preference, or optimal temperatures and nutrients for growth in vitro. Together, these results provide insight into endophytic symbioses in desert-plant communities and can be used to optimize strategies for capturing endophyte biodiversity at regional scales.
Forty-one isolates of post-fire Ascomycetes, representing 1 species of Sphaeriales and 25 species of Pezizales, were tested for ability to hydrolyze amylose, amylopectin, gelatin, xylan, lipid, pectin, chitin, cellulose, and lignin. Isolates were also assayed for the presence of phenol oxidase enzymes: laccase, tyrosinase and peroxidase. The majority of isolates were able to degrade amylose, amylopectin, gelatin, and xylan, and were unable to utilize chitin. Cellulose, lignin, and pectin utilization showed the least variation between isolates of the same species. Pectin hydrolysis and phenol oxidase production were strongly influenced by the pH of the medium. Phenol oxidase production was identified in both saprotrophs and biotrophs. However, lignin degradation was primarily observed in the saprotrophs. It is hypothesized that phenol oxidases in post-fire Pezizales function in lignin degradation in saprotrophs, but function primarily in detoxification of host-produced phenols in biotrophs. Tests were able to identify guilds of fungi associated with non-lignified substrates (e.g. litter, fine roots) and lignified substrates (e.g. wood, woody roots). Tests also helped distinguish between saprotrophic and biotrophic modes of nutrition. The majority of post-fire Pezizales appear to be opportunistic decomposers or facultative biotrophs adapted to the transient post-fire environment.
A systematic study of a rare and enigmatic European species, Galerina clavus Romagn., is presented. Phylogenetic analyses show it to be most closely related to Pachylepyrium carbonicola (A.H. Sm.) Singer and P. funariophilum (M.M. Moser) Singer (Strophariaceae). Investigation of additional species of Pachylepyrium suggests this genus is polyphyletic as the type species, P. fulvidula (Singer) Singer, is nested in the Tubariaceae Vizzini based on multigene phylogenetic analyses. Pachylepyrium nubicola Singer is allied with Pholiota (Fr.) P. Kumm. based on high ITS similarity, and P. carbonicola and P. funariophilum, together with G. clavus, form a clade among a consortium of Strophariaceae Singer & A.H. Sm. and Hymenogastraceae Vittad. As a result, we propose Romagnesiella gen. nov. to accommodate G. clavus, for which a taxonomic description is given and lectotype and epitype are designated. The genus Crassisporium gen. nov. is proposed to encompass Pachylepyrium funariophilum (of which P. carbonicola is considered a younger taxonomic synonym), P. chilense M.M. Moser, and P. squarrulosum Singer. Crassisporium is distinguished from Romagnesiella by its thick-walled basidiospores and occurrence in burnt habitats. The identities of the morphologically similar Tubaria umbonata S. Lundell, T. embolus (Fr.) Sacc. and T. minima J.E. Lange are also discussed.
The present study compares the macrofungi at three study sites in the lowland wet eucalypt forest of southern Tasmania, all in early stages of regeneration, two from silvicultural treatments and one from wildfire. Although not part of a designed experiment, the three areas provided an unbroken, partially overlapping time-line in which macrofungi were recorded during the first 38 months of their development after disturbance and/or fire. Two of the three regenerating units had adjacent mature forest, the macrofungi of which were markedly different from those in the regeneration, the latter being dominated by opportunistic, predominantly saprotrophic species and very low in the symbiotic basidiomycetous ectomycorrhizal species that are abundant in the soils of mature forests. Studies such as these assist in a growing understanding of the nature of the early successional mycota in the southern forests of Tasmania.
Pyrophilous fungi produce sporocarps after a fire but little is known about their ecology prior to or after a fire event. Recently, the body snatchers hypothesis was proposed that suggests some post-fire fungi form endophytic and/or endolichenic relationships with plants and lichens. To test the body snatchers hypothesis, bryophyte, lichen, club moss, and soil samples were collected from unburned and mixed-intensity burned areas 1–2 y after a 2016 wildfire in the Great Smoky Mountains National Park, and from unburned areas in four states outside the park. Samples were examined for the presence of pyrophilous fungi occurring as endophytes or in lichens using culture-dependent and culture-independent techniques. Culture-dependent methods isolated Pholiota highlandensis, a known pyrophilous fungus, from five bryophyte samples. Culture-independent methods identified 22 pyrophilous taxa from bryophyte, club moss, lichen, and soil samples across a range of geographical localities. The ‘body snatchers’ hypothesis is supported since many bryophyte, lichen, and club moss samples contained pyrophilous taxa suggesting that these fungi occur as endophytes and/or endolichenic fungi until a fire event triggers them to produce sporocarps.
The mating system of the postfire ascomycete Rhizina undulata was investigated. Five fruit-bodies of the fungus were collected from three different sites in western Lithuania. Pure cultures were obtained both from the fruit-bodies and from single ascospores. Mycelia were paired on Petri dishes in all possible combinations. Complete intermingling was noted in all pairings among single ascospore isolates and mycelium from the same fruit-body, and from three different fruit-bodies from the same locality. Demarcation zones were noted in pairings among single ascospore isolates and fruit-body mycelia from different localities. Amplified DNA patterns within ascospore families and among families within the same locality were identical or nearly identical. Single ascospore isolates of R. undulata formed fruit-bodies in vitro, although ascospore production was not observed. Thus, R. undulata is reported to be homothallic. The possibilities are discussed both for dispersive and territorial clonality in its natural populations.
A systematic reevaluation of North American pyrophilous or “burn-loving” species of Pholiota is presented based on molecular and morphological examination of type and historical collections. Confusion surrounds application of the names P. brunnescens, P. carbonaria, P. castanea, P. fulvozonata, P. highlandensis, P. molesta, and P. subsaponacea, with multiple names applied to a single species and multiple species described more than once. Molecular annotations using nuc rDNA ITS1-5.8S-ITS2 (internal transcribed spacer [ITS] barcode) and RPB2 (RNA polymerase II second largest subunit) are used to aid in application of these names in a phylogenetic context. Based on ITS molecular annotations of 13 types, the following heterotypic synonymies are proposed: P. highlandensis (syn. P. carbonaria and P. fulvozonata); P. molesta (syn. P. subsaponacea); and P. brunnescens (syn. P. luteobadia). In addition, we observed that the species P. castanea, known previously only from the type collection in Tennessee, is found commonly on burned sites near the Gulf Coast and other southeast regions of the United States. Overall, the pyrophilous trait is evolutionarily derived in Pholiota. Endophytic and endolichenic stages were deduced for P. highlandensis, the most widely distributed of the pyrophilous Pholiota. As a result, we introduce the “body snatchers” hypothesis that explains the maintenance of some pyrophilous fungi in ecosystems as endophytes and/or endolichenic fungi. Photographs, taxonomic descriptions, and a dichotomous key to pyrophilous species of Pholiota that occur in North America are presented.
Laboratory incubation of soil samples from burned prairie stands demonstrated the presence of both coprophilous and carbonicolous ascomycetes. These fungi did not appear on soils collected from unburned prairie stands or from spring-burned stands that were sampled in July. Evidence is presented which suggests that prairie fires bring about a reduction in microbial competition at the soil surface, thereby permitting the successful development of these post-fire fungal colonists. Several of the coprophilous ascomycetes appearing on burned soil were also found on deer and rabbit feces collected from unburned prairie stands. It is suggested that grassland fires provide an additional mechanism by which the coprophilous forms can complete their life cycle. Furthermore, these fires may provide the principle means of initiating growth and development of several noncoprophilous ascomycetes.
Forty-one isolates of post-fire Ascomycetes, representing 1 species of Sphaeriales and 25 species of Pezizales, were tested for ability to hydrolyze amylose, amylopectin, gelatin, xylan, lipid, pectin, chitin, cellulose, and lignin. Isolates were also assayed for the presence of phenol oxidase enzymes: lacease, tyrosinase and peroxidase. The majority of isolates were able to degrade amylose, amylopectin, gelatin, and xylan, and were unable to utilize chitin. Cellulose, lignin, and pectin utilization showed the least variation between isolates of the same species. Pectin hydrolysis and phenol oxidase production were strongly influenced by the pH of the medium. Phenol oxidase production was identified in both saprotrophs and biotrophs. However, lignin degradation was primarily observed in the saprotrophs. It is hypothesized that phenol oxidases in post-fire Pezizales function in lignin degradation in saprotrophs, but function primarily in detoxification of host-produced phenols in biotrophs. Tests were able to identify guilds of fungi associated with non-lignified substrates (e.g. litter, fine roots) and lignified substrates (e.g. wood, woody roots). Tests also helped distinguish between saprotrophic and biotrophic modes of nutrition. The majority of post-fire Pezizales appear to be opportunistic decomposers or facultative biotrophs adapted to the transient post-fire environment.
Reports of true morels (Morchella) fruiting on conifer burn sites are common in western North America where five different fire-adapted species of black morels (Elata Clade) have been documented based on multilocus phylogenetic analyses. Fruiting of post-fire morels in eastern North America, by comparison, are rare and limited to a report from Minnesota in 1977 and Eastern Ontario in 1991. Here, nuc rDNA internal transcribed spacer (ITS1-5.8S-ITS2 = ITS) sequences were used to identify the post-fire morel that fruited in great abundance the year following the 2012 Duck Lake Fire in the Upper Peninsula of Michigan and after the 2016 large-scale fire in the Great Smoky Mountains National Park in Tennessee as M. exuberans. A preliminary phylogenetic analysis suggests the collections from eastern North America may be more closely related to those from Europe than from western North America, Europe and China.
Species limits in the small genus Geopyxis are debatable because of problems with interpreting the few phenotypic features and poor documentation of types. To clarify species boundaries and diversity, we studied the morphology of 74 specimens of Geopyxis from the Northern Hemisphere, including five types, and sequenced four loci for 57 representatives: the nuc rDNA ITS1-5.8S-ITS2 (ITS), D1-D2 domains of nuc 28S rDNA (28S), translation elongation factor (tef1), and (or) part of the second largest subunit of the RNA polymerase II (rpb2) (5-7 region). Eight species are delimited. Six species are shown to be highly supported as reciprocally monophyletic: G. aleurioides sp. nov., G. alpina s. l., G. carbonaria, G. delectans, G. korfii, and G. majalis In addition, coalescent-based Bayesian species delimitation shows G. alpina s. l. constitutes three cryptic species: G. alpina s. str., G. deceptiva sp. nov., and G. rehmii ITS-28S sequences of type material show that G. vulcanalis and G. foetida are synonyms of G. carbonaria A lectotype is designated for Humaria delectans and the name is combined in Geopyxis Morphological characters that can be used to distinguish Geopyxis species are presence/absence of a long stipe, spore size and shape, and pigmented resinous exudates in medullary and ectal excipulum. Geopyxis carbonaria and G. delectans produce apothecia almost exclusively on burned ground. Bayesian analyses detected highly supported conflicts among different loci regarding generic delimitation and species relationships. Two hypogeous genera, Stephensia and Hydnocystis, are confirmed to nest within Geopyxis The relationships between species of Geopyxis and Tarzetta, Stephensia shanorii and Paurocotylis pila, are unresolved. Six out of eight species of Geopyxis recognized in this study have intercontinental disjunct distributions.
Nineteen species are recognized from the study area. Laccaria laccata is further divided into two varieties. Discussions regarding the distribution, presumed ectomycorrhizal hosts, and biology of these taxa are provided along with data on select extralimital taxa. -from Author
Morel mushrooms are globally distributed, socially and economically important reproductive structures produced by fungi of the genus Morchella. Morels are highly prized edible mushrooms and significant harvests are collected throughout their range, especially in the first year after fire, when some morel species fruit prolifically. Few studies have quantified post-fire morel mushroom abundance, despite their widespread human use. The purpose of this study is to provide the first ever estimate of post-fire morel mushroom abundance in Sierra Nevada mixed-conifer forest. Specifically, we estimate the abundance and spatial variability of morel mushrooms across an intensively mapped and measured forest research site during the first growing season following fire.
A specialized group of fungi (mostly ascomycetes and agarics) fruit amongst the ashes marking the sites of former fires. The term phoenicoid fungi, meaning ‘arising from the ashes’, has been proposed for them by Carpenter and Trappe (1985), but they are also referred to as pyrophilous, anthracophilous or carbonicolous.
The Ecological Importance of High-Severity Fires, presents information on the current paradigm shift in the way people think about wildfire and ecosystems. While much of the current forest management in fire-adapted ecosystems, especially forests, is focused on fire prevention and suppression, little has been reported on the ecological role of fire, and nothing has been presented on the importance of high-severity fire with regards to the maintenance of native biodiversity and fire-dependent ecosystems and species. This text fills that void, providing a comprehensive reference for documenting and synthesizing fires ecological role. Offers the first reference written on mixed- and high-severity fires and their relevance for biodiversity Contains a broad synthesis of the ecology of mixed- and high-severity fires covering such topics as vegetation, birds, mammals, insects, aquatics, and management actions Explores the conservation vs. public controversy issues around megafires in a rapidly warming world. © 2015 by Dominick A. DellaSala and Chad T. Hanson. Published by Elsevier Inc. All rights reserved.
A survey is given of macrofungal niche substrate groups and macrofungal communities on wood, fire places, dung, arthropods, mosses, and fruit-bodies of other fungi. The dependence of the fungal population on the kind of the substrate and on the origin of the substrate in different species of trees, mosses, animals, or fungi is reviewed in detail. Attention is payed to fungal succession on decaying wood, on fire places and on dung, to classification of lignicolous, carbophilous and coprophilous community types, to phenology of fruiting and to mycogeographical aspects.
Two common and easily confused species of this pyrophilous, operculate discomycete genus are characterized, P. domesticum and P. omphalodes, and various taxonomic characters are evaluated.
