Klaus Høiland

High diversity of root associated fungi in both alpine and arctic Dryas octopetala.

BackgroundDryas octopetala is a widespread dwarf shrub in alpine and arctic regions that forms ectomycorrhizal (ECM) symbiotic relationships with fungi. In this study we investigated the fungal communities associated with roots of D. octopetala in alpine sites in Norway and in the High Arctic on Svalbard, where we aimed to reveal whether the fungal diversity and species composition varied across the Alpine and Arctic regions. The internal transcribed spacer (ITS) region of nuclear ribosomal DNA was used to identify the fungal communities from bulk root samples obtained from 24 plants.ResultsA total of 137 operational taxonomic units (OTUs) were detected (using 97% similarity cut off during sequence clustering) and well-known ECM genera such as Cenococcum, Cortinarius, Hebeloma, Inocybe and Tomentella occurred frequently. There was no decrease in fungal diversity with increasing latitude. The overall spatial heterogeneity was high, but a weak geographical structuring of the composition of OTUs in the root systems was observed. Calculated species accumulation curves did not level off.ConclusionsThis study indicates that the diversity of fungi associated with D. octopetala does not decrease in high latitude arctic regions, which contrasts observations made in a wide spectrum of other organism groups. A high degree of patchiness was observed across root systems, but the fungal communities were nevertheless weakly spatially structured. Non-asymptotical species accumulation curves and the occurrence of a high number of singletons indicated that only a small fraction of the fungal diversity was detected.

Warming-induced shift in European mushroom fruiting phenology

In terrestrial ecosystems, fungi are the major agents of decomposition processes and nutrient cycling and of plant nutrient uptake. Hence, they have a vital impact on ecosystem processes and the terrestrial carbon cycle. Changes in productivity and phenology of fungal fruit bodies can give clues to changes in fungal activity, but understanding these changes in relation to a changing climate is a pending challenge among ecologists. Here we report on phenological changes in fungal fruiting in Europe over the past four decades. Analyses of 746,297 dated and geo-referenced mushroom records of 486 autumnal fruiting species from Austria, Norway, Switzerland, and the United Kingdom revealed a widening of the annual fruiting season in all countries during the period 1970–2007. The mean annual day of fruiting has become later in all countries. However, the interspecific variation in phenological responses was high. Most species moved toward a later ending of their annual fruiting period, a trend that was particularly strong in the United Kingdom, which may reflect regional variation in climate change and its effects. Fruiting of both saprotrophic and mycorrhizal fungi now continues later in the year, but mycorrhizal fungi generally have a more compressed season than saprotrophs. This difference is probably due to the fruiting of mycorrhizal fungi partly depending on cues from the host plant. Extension of the European fungal fruiting season parallels an extended vegetation season in Europe. Changes in fruiting phenology imply changes in mycelia activity, with implications for ecosystem function.

Arctic root-associated fungal community composition reflects environmental filtering.

There is growing evidence that root‐associated fungi have important roles in Arctic ecosystems. Here, we assess the diversity of fungal communities associated with roots of the ectomycorrhizal perennial herb Bistorta vivipara on the Arctic archipelago of Svalbard and investigate whether spatial separation and bioclimatic variation are important structuring factors of fungal community composition. We sampled 160 plants of B. vivipara from 32 localities across Svalbard. DNA was extracted from entire root systems, and 454 pyrosequencing of ITS1 amplicons was used to profile the fungal communities. The fungal communities were predominantly composed of Basidiomycota (55% of reads) and Ascomycota (35%), with the orders Thelephorales (24%), Agaricales (13.8%), Pezizales (12.6%) and Sebacinales (11.3%) accounting for most of the reads. Plants from the same site or region had more similar fungal communities to one another than plants from other sites or regions, and sites clustered together along a weak latitudinal gradient. Furthermore, a decrease in per‐plant OTU richness with increasing latitude was observed. However, no statistically significant spatial autocorrelation between sites was detected, suggesting that environmental filtering, not dispersal limitation, causes the observed patterns. Our analyses suggest that while latitudinal patterns in community composition and richness might reflect bioclimatic influences at global spatial scales, at the smaller spatial scale of the Svalbard archipelago, these changes more likely reflect varied bedrock composition and associated edaphic factors. The need for further studies focusing on identifying those specific bioclimatic and edaphic factors structuring root‐associated fungal community composition at both global and local scales is emphasized.

Mushroom's spore size and time of fruiting are strongly related: is moisture important?

Most basidiomycete fungi produce annual short-lived sexual fruit bodies from which billions of microscopic spores are spread into the air during a short time period. However, little is known about the selective forces that have resulted in some species fruiting early and others later in the fruiting season. This study of relationships between morphological and ecological characteristics, climate factors and time of fruiting are based upon thorough statistical analyses of 66 520 mapped records from Norway, representing 271 species of autumnal fruiting mushroom species. We found a strong relationship between spore size and time of fruiting; on average, a doubling of spore size (volume) corresponded to 3 days earlier fruiting. Small-spored species dominate in the oceanic parts of Norway, whereas large-spored species are typical of more continental parts. In separate analyses, significant relationships were observed between spore size and climate factors. We hypothesize that these relationships are owing to water balance optimization, driven by water storage in spores as a critical factor for successful germination of primary mycelia in the drier micro-environments found earlier in the fruiting season and/or in continental climates.

Population genetics and spatial structure of the fairy ring fungus Marasmius oreades in a Norwegian sand dune ecosystem

The population genetics and spatial structure of the fairy ring fungus Marasmius oreades (Bolt. : Fr.) Fr. was studied by DNA amplification fingerprinting (DAF). Basidiocarp samples were collected from fairy rings from two separate sand dune systems of about 560 m2 and 1750 m2, respectively, on the Lista Peninsula in southwestern Norway in 1996. Samples were collected after a careful mapping of fairy rings and a vegetation survey of the composition and spatial structure of vascular plants, bryophytes and lichens. DAF with standard arbitrary oligonucleotide primers was used to examine the genetic relationship between basidiocarp samples. The study showed that the fungal population contained a high number of genotypes and that about 90% of the fairy rings represented a separate genet. Both cluster and phylogenetic analyses of DAF amplification products established relationships between fairy rings and showed that genetically similar basidiocarps were found close to each other. Overall results showed a weak correspondence between genotype and spatial distribution and no correspondence between genotype and composition of the surrounding vegetation. Furthermore, the occurrence of the four dominant sand dune grass species was randomly distributed among the localities housing the various fungal genotypes, indicating that the fungus did not exhibit genotypic specialization to the various grass species that could host it as a pathogen. Results show that establishment of new individuals generally was mediated by basidiospore dispersal and not by fragmenting dikaryotic, vegetative mycelium, as previously proposed.

Links between Genetic Groups, Indole Alkaloid Profiles and Ecology within the Grass-Parasitic Claviceps purpurea Species Complex

The grass parasitic fungus Claviceps purpurea sensu lato produces sclerotia with toxic indole alkaloids. It constitutes several genetic groups with divergent habitat preferences that recently were delimited into separate proposed species. We aimed to 1) analyze genetic variation of C. purpurea sensu lato in Norway, 2) characterize the associated indole alkaloid profiles, and 3) explore relationships between genetics, alkaloid chemistry and ecology. Approximately 600 sclerotia from 14 different grass species were subjected to various analyses including DNA sequencing and HPLC-MS. Molecular results, supported by chemical and ecological data, revealed one new genetic group (G4) in addition to two of the three known; G1 (C. purpurea sensu stricto) and G2 (C. humidiphila). G3 (C. spartinae) was not found. G4, which was apparently con-specific with the recently described C. arundinis sp. nov, was predominantly found in very wet habitats on Molinia caerulea and infrequently in saline habitats on Leymus arenarius. Its indole-diterpene profile resembled G2, while its ergot alkaloid profile differed from G2 in high amounts of ergosedmam. In contrast to G1, indole-diterpenes were consistently present in G2 and G4. Our study supports and complements the newly proposed species delimitation of the C. purpurea complex, but challenges some species characteristics including host spectrum, habitat preferences and sclerotial floating ability.

Molecular characterization and evaluation of plant litter-associated fungi from the spring ‘grazing corridor’ of a sheep herd vulnerable to alveld disease

This study sample and identify species of fungi on withered vegetation in the spring ‘grazing corridor’ from infields to Narthecium bogs for a sheep herd almost chronically vulnerable to phototoxic disease. Hepatogen photosensitizing disorders like alveld attack grazing sheep, especially lambs, in various parts of the world. It has been hypothesized that saponin metabolites in the monocotyledonous plant Nartheciumossifragum causes the disorder in Norway, however, this has not been verified. Thus, the search for other causal agents or saponin cofactors (the cofactor hypothesis) has been intensified, and endophytic poisonous fungi associated with dead N. ossifragum leaves and grasses are among the prime suspects. The fungal diversity was targeted by obtaining axenic cultures from surface-sterilized plant material, with subsequent DNA isolation, PCR, and sequencing of the ITS nrDNA region. The taxonomic affinities of the obtained sequences were thereafter explored by similarity searches against the public access sequence database EMBL/GenBank/DDBJ. Among the spectrum of identified taxa were representatives of Cladosporium, Fusarium, Penicillium, and Trichoderma, all of which are well known as potential producers of toxins. A possible involvement of these toxic species in the etiology of alveld is evaluated and discussed.

Fine-scale spatiotemporal dynamics of fungal fruiting : Prevalence, amplitude, range and continuity

Despite the critical importance of fungi as symbionts with plants, resources for animals, and drivers of ecosystem function, the spatiotemporal distributions of fungi remain poorly understood. The belowground life cycle of fungi makes it difficult to assess spatial patterns and dynamic processes even with recent molecular techniques. Here we offer an explicit spatiotemporal Bayesian inference of the drivers behind spatial distributions from investigation of a Swiss inventory of fungal fruit bodies. The unique inventory includes three temperate forest sites in which a total of 73 952 fungal fruit bodies were recorded systematically in a spatially explicit design between 1992 and 2006. Our motivation is to understand how broad-scale climate factors may influence spatiotemporal dynamics of fungal fruiting within forests, and if any such effects vary between two functional groups, ectomycorrhizal (ECM) and saprotrophic fungi. For both groups we asked: 1) how consistent are the locations of fruiting patches, the sizes of patches, the quantities of fruit bodies, and of prevalence (occupancy)? 2) Do the annual spatial characteristics of fungal fruiting change systematically over time? 3) Are spatial characteristics of fungal fruiting driven by climatic variation? We found high inter-annual continuity in fruiting for both functional groups. The saprotrophic species were characterised by small patches with variable fruit body counts. In contrast, ECM species were present in larger, but more distinctly delimited patches. The spatial characteristics of the fungal community were only indirectly influenced by climate. However, climate variability influenced overall yields and prevalence, which again links to spatial structure of fruit bodies. Both yield and prevalence were correlated with the amplitudes of occurrence and of fruit body counts, but only prevalence influenced the spatial range. Summarizing, climatic variability affects forest-stand fungal distributions via its influence on yield (amount) and prevalence (occupancy), whereas fungal life-history strategies dictate fine-scale spatial characteristics.

Reply to Gange et al.: Climate-driven changes in the fungal fruiting season in the United Kingdom.

The first comprehensive study of phenological changes in wild fungi (1) revealed a significantly earlier start and later end of the fungal fruiting season than nowadays, based on a dataset collected in a small area (30-mile radius) in southern England during 1950–2005. To determine whether these interesting results were reflected on a larger scale, we analyzed nationwide datasets from Austria, Norway, Switzerland, and the United Kingdom (2). Our analyses confirmed that, for the United Kingdom, the fruiting season has widened over the last half century. Like Gange et al. (1), we hypothesized that these changes are mainly driven by climatic variation. We are, therefore, surprised …

Mycorrhizal fungal communities in coastal sand dunes and heaths investigated by pyrosequencing analyses

Maritime sand dunes and coastal ericaceous heaths are unstable and dynamic habitats for mycorrhizal fungi. Creeping willow (Salix repens) is an important host plant in these habitats in parts of Europe. In this study, we wanted to assess which mycorrhizal fungi are associated with S. repens in four different coastal vegetation types in Southern Norway, three types from sand dunes and one from heaths. Moreover, we investigated which ecological factors are important for the fungal community structure in these vegetation types. Mycorrhizal fungi on S. repens root samples were identified by 454 pyrosequencing of tag-encoded internal transcribed spacer 1 (ITS1) amplicons. Significantly higher fungal richness was observed in hummock dunes and dune slacks compared to eroded dune vegetation. The compositional variation was mainly accounted for by location (plot) and vegetation type and was significantly correlated to content of carbon, nitrogen and phosphorus in soil. The investigated maritime sand dunes and coastal ericaceous heaths hosted mycorrhizal taxa mainly associated with Helotiales, Sebacinales, Thelephorales and Agaricales.