A group of ectomycorrhizal fungi restricts organic matter accumulation in boreal forest.

Boreal forest soils are important global carbon sinks, with significant storage in the organic topsoil. Decomposition of these stocks requires oxidative enzymes, uniquely produced by fungi. Across Swedish boreal forests, we found that local carbon storage in the organic topsoil was 33% lower in the presence of a group of closely related species of ectomycorrhizal fungi - Cortinarius acutus s.l.. This observation challenges the prevailing view that ectomycorrhizal fungi generally act to increase carbon storage in soils but supports the idea that certain ectomycorrhizal fungi can complement free-living decomposers, maintaining organic matter turnover, nutrient cycling and tree productivity under nutrient-poor conditions. The indication that a narrow group of fungi may exert a major influence on carbon cycling questions the prevailing dogma of functional redundancy among microbial decomposers. Cortinarius acutus s.l. responds negatively to stand-replacing disturbance, and associated population declines are likely to increase soil carbon sequestration while impeding long-term nutrient cycling.

Keeping pace with forestry: Multi-scale conservation in a changing production forest matrix.

The multi-scale approach to conserving forest biodiversity has been used in Sweden since the 1980s, a period defined by increased reserve area and conservation actions within production forests. However, two thousand forest-associated species remain on Sweden's red-list, and Sweden's 2020 goals for sustainable forests are not being met. We argue that ongoing changes in the production forest matrix require more consideration, and that multi-scale conservation must be adapted to, and integrated with, production forest development. To make this case, we summarize trends in habitat provision by Sweden's protected and production forests, and the variety of ways silviculture can affect biodiversity. We discuss how different forestry trajectories affect the type and extent of conservation approaches needed to secure biodiversity, and suggest leverage points for aiding the adoption of diversified silviculture. Sweden's long-term experience with multi-scale conservation and intensive forestry provides insights for other countries trying to conserve species within production landscapes.

Correction to: Keeping pace with forestry: Multi-scale conservation in a changing production forest matrix.

In the original published article, the sentence "Nevertheless, semi-natural forest remnants continue to be harvested and fragmented (Svensson et al. 2018; Jonsson et al. 2019), and over 2000 forest-associated species (of 15 000 assessed) are listed as threatened on Sweden's red-list, largely represented by macro-fungi, beetles, lichens and butterflies (Sandström 2015)."under the section Introduction was incorrect. The correct version of the sentence is "Nevertheless, semi-natural forest remnants continue to be harvested and fragmented (Svensson et al. 2018; Jonsson et al. 2019), and approximately 2000 forest-associated species (of 15 000 assessed) are on Sweden's red-list, largely represented by macro-fungi, beetles, lichens and butterflies (Sandström 2015)."

The evolutionary species pool concept does not explain occurrence patterns of dead-wood-dependent organisms: implications for logging residue extraction.

Emulation of natural disturbances is often regarded as a key measure to make forestry biodiversity-oriented. Consequently, extraction of logging residues is assumed to have little negative effect in comparison to extraction of dead wood mainly formed at natural disturbances. This is consistent with the evolutionary species pool hypothesis, which suggests that most species are evolutionary adapted to the naturally most abundant habitats. We tested this hypothesis for dead-wood-dependent macrofungi, lichens, and beetles in a boreal forest landscape in central Sweden, assuming that species are adapted to conditions similar to today's unmanaged forest. No occurrence patterns, for the species groups which we investigated, were consistent with the hypothesis. Overall, stumps and snags had the highest habitat quality (measured as average population density with equal weight given to each species) and fine woody debris the lowest, which was unexpected, since stumps were the rarest dead-wood type in unmanaged forest. We conclude that the evolutionary species pool concept did not explain patterns of species' occurrences, and for two reasons, the concept is not reliable as a general rule of thumb: (1) what constitute habitats harbouring different species communities can only be understood from habitat-specific studies and (2) the suitability of habitats is affected by their biophysical characteristics. Thus, emulation of natural disturbances may promote biodiversity, but empirical studies are needed for each habitat to understand how natural disturbances should be emulated. We also conclude that stump extraction for bioenergy is associated with larger risks for biodiversity than fine woody debris extraction.

Retention of seed trees fails to lifeboat ectomycorrhizal fungal diversity in harvested Scots pine forests.

Fennoscandian forestry has in the past decades changed from natural regeneration of forests towards replantation of clear-cuts, which negatively impacts ectomycorrhizal fungal (EMF) diversity. Retention of trees during harvesting enables EMF survival, and we therefore expected EMF communities to be more similar to those in old natural stands after forest regeneration using seed trees compared to full clear-cutting and replanting. We sequenced fungal internal transcribed spacer 2 (ITS2) amplicons to assess EMF communities in 10- to 60-year-old Scots pine stands regenerated either using seed trees or through replanting of clear-cuts with old natural stands as reference. We also investigated local EMF communities around retained old trees. We found that retention of seed trees failed to mitigate the impact of harvesting on EMF community composition and diversity. With increasing stand age, EMF communities became increasingly similar to those in old natural stands and permanently retained trees maintained EMF locally. From our observations, we conclude that EMF communities, at least common species, post-harvest are more influenced by environmental filtering, resulting from environmental changes induced by harvest, than by the continuity of trees. These results suggest that retention of intact forest patches is a more efficient way to conserve EMF diversity than retaining dispersed single trees.

Consequences of bioenergy wood extraction for landscape-level availability of habitat for dead wood-dependent organisms.

Stumps and slash resulting from forest clearcutting is used as a source of low-net-carbon energy, but there are concerns about the consequences of biofuel extraction on biodiversity. Logging residues constitute potentially important habitats, since a large part of forest biodiversity is dependent on dead wood. Here we used snapshot field data from a managed forest landscape (25 000 ha) to predict landscape scale population changes of dead wood dependent organisms after extraction of stumps and slash after clearcutting. We did this by estimating habitat availability for all observed dead wood-dependent beetles, macrofungi, and lichens (380 species) in the whole landscape. We found that 53% of species occurred in slash or stumps. For most species, population declines after moderate extraction (≤30%) were small (<10% decline) because they mainly occur on other dead wood types. However, some species were only recorded in slash and stumps. Red listed species were affected by slash and stump extraction (12 species), but less often than other species. Beetles and fungi were more affected by stump extraction, while lichens were more affected by slash extraction. For beetles and lichens, extraction of a combination of spruce, pine and birch resulted in larger negative effects than if only extracting spruce, while for fungi tree species had little effect. We conclude that extensive extraction decreases the amount of habitat to such extent that it may have negative consequences on species persistence at the landscape level. The negative consequences can be limited by extracting only slash, or only logging residues from spruce stands.

Diverse ecological roles within fungal communities in decomposing logs of Picea abies.

Fungal communities in Norway spruce (Picea abies) logs in two forests in Sweden were investigated by 454-sequence analyses and by examining the ecological roles of the detected taxa. We also investigated the relationship between fruit bodies and mycelia in wood and whether community assembly was affected by how the dead wood was formed. Fungal communities were highly variable in terms of phylogenetic composition and ecological roles: 1910 fungal operational taxonomic units (OTUs) were detected; 21% were identified to species level. In total, 58% of the OTUs were ascomycetes and 31% basidiomycetes. Of the 231 337 reads, 38% were ascomycetes and 60% basidiomycetes. Ecological roles were assigned to 35% of the OTUs, accounting for 62% of the reads. Wood-decaying fungi were the most common group; however, other saprotrophic, mycorrhizal, lichenized, parasitic and endophytic fungi were also common. Fungal communities in logs formed by stem breakage were different to those in logs originating from butt breakage or uprooting. DNA of specific species was detected in logs many years after the last recorded fungal fruiting. Combining taxonomic identification with knowledge of ecological roles may provide valuable insights into properties of fungal communities; however, precise ecological information about many fungal species is still lacking.

A fungal perspective on conservation biology.

Hitherto fungi have rarely been considered in conservation biology, but this is changing as the field moves from addressing single species issues to an integrative ecosystem-based approach. The current emphasis on biodiversity as a provider of ecosystem services throws the spotlight on the vast diversity of fungi, their crucial roles in terrestrial ecosystems, and the benefits of considering fungi in concert with animals and plants. We reviewed the role of fungi in ecosystems and composed an overview of the current state of conservation of fungi. There are 5 areas in which fungi can be readily integrated into conservation: as providers of habitats and processes important for other organisms; as indicators of desired or undesired trends in ecosystem functioning; as indicators of habitats of conservation value; as providers of powerful links between human societies and the natural world because of their value as food, medicine, and biotechnological tools; and as sources of novel tools and approaches for conservation of megadiverse organism groups. We hope conservation professionals will value the potential of fungi, engage mycologists in their work, and appreciate the crucial role of fungi in nature.

Carbon sequestration is related to mycorrhizal fungal community shifts during long-term succession in boreal forests.

Boreal forest soils store a major proportion of the global terrestrial carbon (C) and below-ground inputs contribute as much as above-ground plant litter to the total C stored in the soil. A better understanding of the dynamics and drivers of root-associated fungal communities is essential to predict long-term soil C storage and climate feedbacks in northern ecosystems. We used 454-pyrosequencing to identify fungal communities across fine-scaled soil profiles in a 5000 yr fire-driven boreal forest chronosequence, with the aim of pinpointing shifts in fungal community composition that may underlie variation in below-ground C sequestration. In early successional-stage forests, higher abundance of cord-forming ectomycorrhizal fungi (such as Cortinarius and Suillus species) was linked to rapid turnover of mycelial biomass and necromass, efficient nitrogen (N) mobilization and low C sequestration. In late successional-stage forests, cord formers declined, while ericoid mycorrhizal ascomycetes continued to dominate, potentially facilitating long-term humus build-up through production of melanized hyphae that resist decomposition. Our results suggest that cord-forming ectomycorrhizal fungi and ericoid mycorrhizal fungi play opposing roles in below-ground C storage. We postulate that, by affecting turnover and decomposition of fungal tissues, mycorrhizal fungal identity and growth form are critical determinants of C and N sequestration in boreal forests.

Occurrence patterns of lichens on stumps in young managed forests.

The increasing demand for forest-derived bio-fuel may decrease the amount of dead wood and hence also the amount of available substrate for saproxylic ( = dead-wood dependent) organisms. Cut stumps constitute a large portion of dead wood in managed boreal forests. The lichen flora of such stumps has received little interest. Therefore, we investigated which lichens that occur on stumps in young (4-19 years), managed forests and analyzed how species richness and occurrence of individual species were related to stump and stand characteristics. We performed lichen inventories of 576 Norway spruce stumps in 48 forest stands in two study areas in Central Sweden, recording in total 77 lichen species. Of these, 14 were obligately lignicolous, while the remaining were generalists that also grow on bark, soil or rocks. We tested the effect of characteristics reflecting successional stage, microclimate, substrate patch size, and the species pool in the surrounding area on (1) total lichen species richness, (2) species richness of obligately lignicolous lichens and (3) the occurrence of four obligately lignicolous lichen species. The most important variables were stump age, with more species on old stumps, and study area, with similar total species richness but differences in occupancy for individual species. Responses for total lichen species richness and species richness of obligately lignicolous lichens were overall similar, indicating similar ecological requirements of these two groups. Our results indicate that species richness measurements serve as poor proxies for the responses of individual, obligately lignicolous lichen species.

Evidence-based knowledge versus negotiated indicators for assessment of ecological sustainability: the Swedish Forest Stewardship Council standard as a case study.

Assessing ecological sustainability involves monitoring of indicators and comparison of their states with performance targets that are deemed sustainable. First, a normative model was developed centered on evidence-based knowledge about (a) forest composition, structure, and function at multiple scales, and (b) performance targets derived by quantifying the habitat amount in naturally dynamic forests, and as required for presence of populations of specialized focal species. Second, we compared the Forest Stewardship Council (FSC) certification standards' ecological indicators from 1998 and 2010 in Sweden to the normative model using a Specific, Measurable, Accurate, Realistic, and Timebound (SMART) indicator approach. Indicator variables and targets for riparian and aquatic ecosystems were clearly under-represented compared to terrestrial ones. FSC's ecological indicators expanded over time from composition and structure towards function, and from finer to coarser spatial scales. However, SMART indicators were few. Moreover, they poorly reflected quantitative evidence-based knowledge, a consequence of the fact that forest certification mirrors the outcome of a complex social negotiation process.

Wood-inhabiting fungal communities in woody debris of Norway spruce (Picea abies (L.) Karst.), as reflected by sporocarps, mycelial isolations and T-RFLP identification.

Wood-inhabiting fungi play a key role in forest ecosystems and constitute an essential part of forest biodiversity. We therefore examined the composition and abundance of wood-inhabiting fungi by three methods: sporocarp counts, mycelial culturing and direct amplification of internal transcribed spacer terminal restriction fragment length polymorphism from wood combined with sequencing of reference rDNA. Seven-year-old slash piles left after a thinning were analyzed in a 50-year-old Norway spruce plantation. Fifty-eight fungal species were detected from the piled branches and treetops. More species were revealed by sporocarp counts and cultured mycelia than by direct amplification from wood. In principle, sporocarp monitoring may reveal all fruiting taxa, but it poorly reflects their relative abundance in the wood. In contrast, terminal restriction fragment length polymorphism will record the most frequent fungal taxa in the wood, but it may overlook uncommon taxa. Culturing mycelia from wood gives a bias towards species favoured by the cultural medium. The results demonstrate the advantage and the limitations of these methods to be considered in analyses of fungal communities in wood.

Size, distribution and biomass of genets in populations of Suillus bovinus (L.: Fr.) Roussel revealed by somatic incompatibility.

The spatial distribution of genets in the ectomycorrhizal fungus Suillus bovinus (L.: Fr.) Roussel were studied in somatic incompatibility pairings of isolates from five Scots pine (Pinus sylvestris L.) stands differing in forest history and age, With increasing forest age, the size of genets increased while the number of genets and production of sporocarps per unit area decreased. There was an estimated 700-5700 genets ha-1 in younger forests and 30-120 ha-1 in older ones. The maximum size of genets was 1.7-5.3 m in the younger forest and to 17.5 m in the older ones. The production of sporocarps per unit area decreased with increasing forest age. Furthermore, production of sporocarps increased nonlinearly with the size of the genet, suggesting that genets become fragmented. Ergosterol measurements indicated that the fungal biomass of one genet consisted of 20-45% sporocarps and 55-80% mycorrhiza, not including extramatrical mycelia. Mycorrhizal aggregations in soil were mapped in two 5 m2 areas and, based on somatic incompatibility tests, all were round CO belong to the same genet as sporocarps present above ground. Production of spores per sporocarp was estimated to be 1.1-12.8 × 108 . The observations are discussed in terms of population ecology.

Population structure and dynamics in Suillus bovinus as indicated by spatial distribution of fungal clones.

The spatial distribution of clones in the ectomycorrhizal fungus Suillus bovinus (L. ex Fr.) O. Kuntze was studied from somatic incompatibility pairings of isolates in four Scots pine (Pinus sylvestris L.) stands with different forest history. A pattern of increasing clonal size, decreasing clonal number, and decreasing fruiting per unit area, was found with increased forest age and relatively little disturbance. Establishment by spores was important on disturbed sites, while mycelial spread increased in importance with decreased disturbance. The number of clones was found to be 800 ha-1 in younger forest stands and decreased to between 25-130 ha-1 in mature stands. At the same time the diameter of the area occupied by a single clone increased from 1-3 m to 30 m. Implications of the observations are discussed in terms of ecological strategies and population dynamics.