Fire as a driver of fungal diversity - A synthesis of current knowledge.

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.

Systematic analysis of Russula in the North American Rocky Mountain alpine zone.

Russula (Russulales) is an important ectomycorrhizal fungal genus in Arctic and alpine regions where it occurs with Salix, Betula, Dryas, and Polygonum, yet a complex phylogenetic analysis of the genus in these habitats is lacking. This research compared collections of Russula from the Rocky Mountain alpine (Colorado, Montana, Wyoming) with reference specimens from Arctic and alpine habitats, mostly in Europe, using an in-depth morphological study and a phylogenetic analysis of the nuc rDNA internal transcribed spacer region ITS1-5.8S-ITS2 (ITS barcode) and the second largest subunit of the RNA polymerase II gene (rpb2). One hundred thirty-nine Russula collections were sequenced, including type material. Ten species are reported from alpine or treeline habitats in the Rocky Mountains. This is the first formal report of R. cf. altaica, R. saliceticola, and R. subrubens from the Rocky Mountains and of R. purpureofusca in North America. Russula laevis is reported for the first time under this name with a voucher, and not as an environmental sample. Previous reports of R. nana and R. laccata are molecularly confirmed. Two species are reported from subalpine habitats at treeline: R. montana with conifers and R. cf. altaica with Betula. In this study, R. laccata, R. subrubens, and R. laevis were collected in alpine habitats but have been reported below treeline in Europe; these species may also be present at lower elevations in North America. Most species have an intercontinental distribution and have been reported in other alpine or Arctic habitats. Two unidentified and potentially new species were only found in North America and are discussed. A key to the alpine Russulas of North America is provided.

Nodulose-spored Inocybe from the Rocky Mountain alpine zone molecularly linked to European and type specimens.

Inocybe (Inocybaceae) is one of the most diverse ectomycorrhizal genera in arctic and alpine habitats where the primary hosts are Salix, Betula, and Dryas. Subgenus Inocybe is common in these habitats and typically characterized by the presence of thick-walled pleurocystidia. Here, we focus on species that have angular or nodulose spores. Historically, over 30 taxa from this group have been reported from arctic and alpine habitats. Many names have been synonymized, whereas molecular analysis has revealed new species. Nuc rDNA internal transcribed spacer ITS1-5.8S-ITS2 (ITS) sequence data of 26 type specimens in this group now allow for further taxonomic clarification and comparison across continents of disjunct populations. Here, we compare ITS sequence data and the D1-D2 portion of nuc 28S rDNA (28S) from Rocky Mountain specimens with those of types and European reference material. We report 10 species from the Rocky Mountain alpine zone, all of which are conspecific with known European boreal, montane, or alpine species, and four are described as new; all have intercontinental distributions. Nodulose-spored Inocybe taxa that occur in the Rocky Mountain alpine zone include I. alpinomarginata, sp. nov., I. arctica, I. giacomi, I. leonina, I. murina, sp. nov., I. occulta, I. paragiacomi, sp. nov., I. phaeocystidiosa, I. purpureobadia, and I. subgiacomi, sp. nov. Remarkably, these species occur at elevations up to 4000 m and at latitudes as low as 36°N, hundreds of miles from the Arctic, the European alpine, and original type localities. Distributions are explained in part by host distributions and historical glaciation patterns. A key and full descriptions for Rocky mountain species are provided to promote species recognition.

The genus Hebeloma in the Rocky Mountain Alpine Zone.

Numerous taxa of Hebeloma have been reported in association with Salix, Dryas, and Betula in arctic-alpine habitats. However, species are notoriously difficult to delineate because morphological features overlap, and previously there was little reliable molecular data available. Recent progress in ITS-sequencing within the genus, coupled with an extensive database of parametrically described collections, now allows comparisons between species and their distributions. Here we report 16 species of Hebeloma from the Rocky Mountain alpine zone from some of the lowest latitudes (latitude 36°-45°N) and highest elevations (3000-4000 m) for arctic-alpine fungi in the northern hemisphere. Twelve of these species have been reported from arctic-alpine habitats in Europe and Greenland and are now molecularly confirmed from the Middle and Southern Rockies, greatly expanding their distribution. These are: Hebelomaalpinum, H.aurantioumbrinum, H.dunense, H.hiemale, H.marginatulum, H.mesophaeum, H.nigellum, H.oreophilum, H.subconcolor, H.spetsbergense, H.vaccinum, and H.velutipes. Hebelomahygrophilum is known from subalpine habitats in Europe, but was never recorded in arctic-alpine ecology. Three species recorded from the Rockies, but as yet not reported from Europe, are H.alpinicola, H.avellaneum, and H.excedens. The last two have never previously been reported from an arctic-alpine habitat. For all three of these species, the holotypes have been studied morphologically and molecularly, and have been incorporated into the analysis.

Systematics of the ectomycorrhizal genus Lactarius in the Rocky Mountain alpine zone.

Lactarius (Russulales) is an important component of ectomycorrhizal fungal communities in cold-dominated contiguous arctic and disjunct alpine habitats where it associates primarily with Betula, Dryas and Salix However, little is known of this genus in the central and southern Rocky Mountain alpine zone (3000-3900 m) of North America. Molecular phylogenetic analyses of nuc rDNA ITS1-5.8S-ITS2 (ITS barcode) and the second largest subunit of the RNA polymerase II gene (RPB2) partial sequences in conjunction with detailed morphological examination confirm at least six species occurring above treeline. Most have intercontinental distributions in North America and Eurasia according to molecular comparison with type material and collections from Europe, Fennoscandia, Svalbard and Alaska. Rocky Mountain collections of L. lanceolatus (subgenus Russularia), along with the type from Alaska are paraphyletic with respect to L. aurantiacus and North American taxa L. luculentus and L. luculentus v. laetus Rocky Mountain collections of L. nanus, L. glyciosmus, L. repraesentaneus and L. salicis-reticulatae (subgenus Piperites) all form clades with European material from type localities and other arctic-alpine habitats. The arctic-alpine L. pseudouvidus/L. brunneoviolaceus group appears to be a complex containing additional taxa. North American material originally described as part of this group is well-separated phylogenetically and is described here as L. pallidomarginatus sp. nov. Lactarius lanceolatus, L. nanus and L. salicis-reticulatae appear largely restricted to arctic-alpine habitats with Salix Lactarius glyciosmus and L. repraesentaneus occur in arctic-alpine, subalpine and boreal habitats with Betula and also Picea and possibly Salix for the latter. Species distributions are hypothesized to be shaped by host ranges, glaciation and long distance dispersal.

Morphological and molecular systematics of Rocky Mountain alpine Laccaria.

The alpine zone is comprised of habitats at elevations above treeline, and macromycetes play important ecological roles as decomposers and mycorrhizal symbionts here as elsewhere. Laccaria is an important group of ectomycorrhizal basidiomycetes widely used in experimental and applied research. A systematic study of alpine Laccaria species using morphological, cultural and molecular (ribosomal DNA internal transcribed spacer) data revealed five taxa in the Rocky Mountain alpine zone: L. laccata var. pallidifolia, L. nobilis (the first published report for arctic-alpine habitats), L. pumila, L. montana and L. pseudomontana (a newly described taxon similar to L. montana with more ellipsoidal, finely echinulate basidiospores). All occur in the southern Rocky Mountains of Colorado; however, only L. pumila and L. montana were found on the Beartooth Plateau in the northern Rocky Mountains of Montana and Wyoming. All are associated with dwarf and shrub Salix species, with L. laccata var. pallidifolia also associated with Dryas octopetala and Betula glandulosa. Maximum-parsimony phylogenetic analysis of rDNA-ITS sequences for 27 Laccaria accessions supports the morphological species delineations.