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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.

• Demographic studies measure drivers of plant fecundity including seed production and survival, but few address both abiotic and biotic drivers of germination such as variation in climate among sites, population density, maternal plants, seed type and fungal pathogen abundance. • We examined germination and microbial communities of seeds of Danthonia californica, which are either chasmogamous (external, wind-pollinated) or cleistogamous (internal, self-fertilized) and Festuca roemeri, which are solely chasmogamous. Seed populations were sourced across environmental gradients. We tested germination and used high-throughput sequencing to characterize seed fungal community structure. • For F. roemeri, maternal plants significantly influenced germination as did climate and pathogens; germination increased from wetter, cooler sites. For D. californica, the main drivers of germination were maternal plant, seed type and pathogens; on average, more chasmogamous seeds germinated. Fungal communities depended largely on seed type, with fewer fungi associated with cleistogamous seeds, but the communities also depended on site factors such as vapor pressure deficit, plant density and whether the seeds had germinated. • Putative pathogens that were negatively correlated with germination were more abundant for both D. californica and F. roemeri chasmogamous seeds than D. californica cleistogamous seeds. In D. californica, cleistogamous and chasmogamous seeds contain vastly different fungal communities.

Species of Hygrocybe (waxcaps) are mostly colorful mushrooms, which are characteristic of undisturbed grasslands. These fungi are endangered in many places worldwide, but their biology remains a mystery: while isotopic signatures indicate that waxcaps are neither mycorrhizal nor saprotrophic, they were recently observed in plant roots and molecularly detected in aboveground tissues. We aimed to establish a model system of Plantago lanceolata plants colonized by H . coccinea for future detailed studies of the plant–fungus association, and species-specific primers were designed to control infection success and screen environmental samples for waxcaps. The experimentally treated plants grown from surface-sterilized seeds were indeed colonized by waxcaps after 22 weeks of incubation. However, the fungal infection was independent from the experimental treatment and apparently resulted from infected seeds. Screening of field material confirmed that at least one species, i.e., H . virginea , is a maternally transmitted endophytic fungus associated with P . lanceolata . In the experiments, it obviously expanded to the roots during or after seed germination. The endophytic growth is also consistent with the carbon isotopic signature of Hygrocybe , which deviates less from the host plants’ signature than known from ectomycorrhizal associations. However, waxcaps obviously acquire nitrogen (N) from a source outside the plant, like mycorrhizal fungi do. The extensive root system of P . lanceolata is hypothesized to facilitate reaching of nitrogen sources for Hygrocybe which are enriched in the heavier 15  N isotope.

This study describes four gray or brown species of Cuphophyllus (Hygrophoraceae, Agaricales), two of them new species, restricted to arctic-alpine and northern boreal zones of North America, and relates them morphologically and phylogenetically using multigene and nuc rDNA internal transcribed spacer ITS1-5.8S-ITS (ITS barcode) analyses to their similar, known counterparts. Cuphophyllus cinerellus, epitypified here, is shown to be a pan-palearctic species with sequence-confirmed collections from Fennoscandia and easternmost Asia. Occupying a similar habitat in the Nearctic is its sister species, the morphologically similar but novel C. esteriae, so far known only from eastern North America, including Greenland. Sister to the C. cinerellus-C. esteriae lineage, and known only from boreal raised Sphagnum bogs in Newfoundland, is a new medium-sized light cinereous brown species, C. lamarum. It has a yellow stipe but is phylogenetically distant from the yellow-stiped European C. flavipes and its North American sister species, Hygrophorus pseudopallidus. As cryptic speciation was discovered within C. flavipes, we lecto- and epitypify the name and transfer H. pseudopallidus to Cuphophyllus based on ITS analysis of the holotype. We also transfer the small European Hygrocybe comosa to Cuphophyllus based on morphology. Cuphophyllus hygrocyboides is reported from North America with the first sequence-confirmed collections from arctic-alpine British Columbia and Greenland. In addition, sequencing the holotype of C. subviolaceus identifies it as the sister species to the putative C. lacmus. Both species seem to have an intercontinental distribution. In total, we add new sequences to GenBank from 37 Cuphophyllus collections, including the holotypes of C. hygrocyboides and C. subviolaceus, the two new epitypes, and the two novel species.

Aim Geographic distributions of mushroom species remain poorly understood despite their importance for advancing our understanding of the habitat requirements, species interactions and ecosystem functions of this key group of organisms. Here, we estimate geographic range extents (maximum within‐species geographic distance) of genetically defined operational taxonomic units (OTUs). Location World‐wide, with emphasis on the American Pacific Northwest. Taxa Amanita, Agaricus, Cortinarius, Galerina, Hebeloma, Hydnum, Hygrocybe, Hygrophorus, Inocybe, Lepiota, Pholiota, and Russula + Lactarius; other genera in Agaricomycotina. Method We used publicly available OTUs from ribosomal internal transcribed spacer (ITS) sequences (n = 15,373) from 12 mushroom genera with worldwide distributions. For each of 2,324 ~ species‐level OTUs, we estimated the maximum within‐species range extent based on sample locality records. In parallel, we estimated range extents for species in four tree genera. Contrasting estimates from well‐studied trees allowed us to test for potential biases in our range estimates of less well inventoried mushrooms. Results The median range extents across the 2,324 mushroom OTUs varied from ~ 1,200 to 4,039 km, depending on assumptions. These extents were significantly lower than estimates from permuted or randomized data. Mushroom ranges were comparable to the median natural range extent of tree species (1,613 km). In contrast, the tree median species range increased to 16,581 km when anthropogenic range extensions were included. At least 10 mushroom species were similarly broadly distributed, eight of which have been associated with human activity. Main conclusions Overall, like tree species, mycorrhizal and saprotrophic fungi show evidence of biogeographic structure rather than global distributions. This reconstruction of geographic range extents drew upon investments into ITS barcoding of extensive herbarium collections. Large scale analyses such as ours can yield estimates of fungal geographic range extents that are a prerequisite to a deeper understanding of the diverse roles of fungi in ecosystems.

Agaric fungi of the southern Appalachian Mountains including Great Smoky Mountains National Park are often heterozygous for the rDNA internal transcribed spacer region (ITS) with >42% of collections showing some heterozygosity for indels and/or base-pair substitutions. For these collections, intra-individual haplotype divergence is typically less than 2%, but for 3% of these collections intra-individual haplotype divergence exceeds that figure. We hypothesize that high intra-individual haplotype divergence is due to hybridization between agaric fungi with divergent haplotypes, possibly migrants from geographically isolated glacial refugia. Four species with relatively high haplotype divergence were examined: Armillaria mellea, Amanita citrina f. lavendula, Gymnopus dichrous and the Hygrocybe flavescens/chlorophana complex. The ITS region was sequenced, haplotypes of heterozygotes were resolved through cloning, and phylogenetic analyses were used to determine the outcome of hybridization events. Within Armillaria mellea and Amanita citrina f. lavendula, we found evidence of interbreeding and recombination. Within G. dichrous and H. flavescens/chlorophana, hybrids were identified but there was no evidence for F 2 or higher progeny in natural populations suggesting that the hybrid fruitbodies might be an evolutionary dead end and that the genetically divergent Mendelian populations from which they were derived are, in fact, different species. The association between ITS haplotype divergence of less than 5% (Armillaria mellea = 2.6% excluding gaps; Amanita citrina f. lavendula = 3.3%) with the presence of putative recombinants and greater than 5% (Gymnopus dichrous = 5.7%; Hygrocybe flavescens/chlorophana = 14.1%) with apparent failure of F 1 hybrids to produce F 2 or higher progeny in populations may suggest a correlation between genetic distance and reproductive isolation.

Hygrocybe ceracea (Sowerby) P. Kumm, the small and macroscopic fungus, with a yellow, greasy, and fragile appearance is reported for the first time on Funaria hygrometrica in Tonekabon, west of Mazandaran province, North of Iran. This fungus appears among the stones and cobblestones with a little humic in high humidity of winter, after stopping rain. The stipe is white, 2 mm in average diameter and 1 -2 cm in length. The cap diameter is 0.5 cm in average. Spores are lacrymoid, 4 X 2.35 pm in their dimensions. The average length of basidium is 10 pm. Tetraspores often remain attached together after separation from the basidium. The cap is initially convex, but it turns to the conic shape with passing time and aging. The middle part becomes turgid, the sugarloaf shape and the margins become flat, and its colour turns from yellow to brownish orange. Its companion species in Iran are Funaria, lichen and grass. The relationship between H. ceracea and Funaria hygrometrica is parasitic type. Funaria leaf spots are caused by infection with this fungus. It is not a poisonous fungus, hence domestic birds and crows feed on it with great interest. Regrowth and grazing resistance were evident in function of the fungus.

The nutritional modes of genera in Hygrophoraceae (Basidiomycota: Agaricales), apart from the ectomycorrhizal Hygrophorus and lichen-forming taxa, are uncertain. New δ 15 N and δ 13 C values were obtained from 15 taxa under Hygrophoraceae collected in central Massachusetts and combined with isotopic datasets from five prior studies including a further 12 species using a data standardization method to allow cross-site comparison. Based on these data, we inferred the probable nutritional modes for species of Hygrophorus, Hygrocybe, Humidicutis, Cuphophyllus and Gliophorus. A phylogeny of Hygrophoraceae was constructed by maximum likelihood analysis of nuclear ribosomal 28S and 5.8S sequences and standardized δ 15 N and δ 13 C values were used for parsimony optimization on this phylogeny. Our results supported a mode of biotrophy in Hygrocybe, Humidicutis, Cuphophyllus and Gliophorus quantitatively unlike that in more than 450 other fungal taxa sampled in the present and prior studies. Parsimony optimization of stable isotope data suggests moderate conservation of nutritional strategies in Hygrophoraceae and a single switch to a predominantly ectomycorrhizal life strategy in the lineage leading to Hygrophorus. We conclude that Hygrophoraceae of previously unknown nutritional status are unlikely to be saprotrophs and are probably in symbiosis with bryophytes or other understory plants.

Hygrocybe aurantiomagnifica (Agaricales), which we describe here as a new species, is characterised by gregarious basidiomata, yellow to orange pileus with an almost plane, smooth surface and undulate or lobed margin; distant, whitish lamellae becoming pinkish-violet at the edge; absence of cheilocystidia and an enterocutis pileipellis. Morphological along with molecular phylogenetic analyses of the internal transcribed spacer (nrITS) and the nuclear large subunit (nrLSU), confirm H. aurantiomagnifica as a new species of subgenus Pseudohygrocybe section Firmae. A comprehensive morphological description, illustrations, comparisons with related species and a key including similar Brazilian species are provided.

In the Chęciny district, xerothermic grasslands developed on deforested slopes of limestone hills, truncated folds, and mounds. Their origin is directly connected with agricultural and pastoral farming of man. Xerothermic grassland belongs to the class Festuco-Brometea, and the alliance Cirsio-Brachypodion. The plant association Thalictro-Salvietum pratensis is the most widespread in this area. The xerothermic grasslands have their own characteristic biota of macromycetes. The following steppe, xerothermic, and thermophilous fungi deserve special attention: Agaricus bernardii, Camarophyllus virgineus, Conocybe sienophylla, Entoloma incanum, Hygrocybe konradii, H. persistens, H. reae, Lepiota alba, and Leucopaxillus lepistoides. As a result of the xerothermic swards being progressively overgrown by shrubs and trees, among others, by Pinus sylvestris, one can find fungi species accompanying this tree: Suillus collinitus, also fungi producing underground fruit bodices Rhizopogon obtextus and Rh. roseolus. The main factors threatening xerothermic grasslands of the Chęciny district are, among others, the devastation and disappearance of natural habitats, often as a result of inappropriate human land management as well as the pollution of air, water, and soil. The effective protection of rare, threatened, and also legally protected fungi species is only possible by protecting their natural habitats as a whole.