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Fungal traits can provide a mechanistic understanding of how wood‐inhabiting fungi interact with their environment and how that influences community assembly in deadwood. However, fungal trait exploration is relatively new and almost no studies measure fungal traits in their environment.In this study we tested species‐ and trait‐environment relationships in reproducing fungal communities inhabiting 571 Norway spruce (Picea abies) logs in 55 isolated forest patches (0.1–9.9 ha) of different naturalness types, located in Northern boreal Sweden. The studied patches were (1) semi‐natural set‐aside patches within highly managed landscapes, or (2) old‐growth natural patches located in an unmanaged landscape. We tested species and trait relationships to deadwood substrate and forest patch variables. We measured mean fruit body size and density for each of the 19 species within communities.Traits assembled in relation to log decay stage and forest patch naturalness, illustrating the important role of deterministic environmental filtering in shaping reproducing wood‐inhabiting fungal communities. Early decay stage communities had larger, less dense, annual fruiting bodies of half‐resupinate type and were more often white‐rot fungi. Species rich mid‐decay stage communities had mixed trait assemblages with more long lived perennial fruit bodies of intermediate size, and both brown‐ and white‐rot fungi equally represented. Finally, late decay stage communities had smaller, denser and perennial fruit bodies, more often of the brown‐rot type.The relationships between the studied traits and decay stages were similar in both set‐aside and natural patches. However, set‐aside semi‐natural patches in highly managed landscapes more frequently supported species with smaller, perennial and resupinate fruit bodies compared to natural patches in an unmanaged landscape.Synthesis. We found that log decay stage was the primary driver of fungal community assembly of species and traits in isolated forest patches. Our results suggest that decay stage filters four reproduction traits (fruit body density, size, lifespan and type) and one resource‐use trait (white or brown rot). Our results highlights, for the first time, that communities with diverse fungal reproductive traits are maintained foremost across all deadwood decay stages under different forest naturalness conditions. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Extreme droughts are globally increasing in frequency and severity. Most research on drought in forests focuses on the response of trees, while less is known about the impacts of drought on forest understory species and how these effects are moderated by the local environment. We assessed the impacts of a 45‐day experimental summer drought on the performance of six boreal forest understory plants, using a transplant experiment with rainout shelters replicated across 25 sites. We recorded growth, vitality and reproduction immediately, 2 months, and 1 year after the simulated drought, and examined how differences in ambient soil moisture and canopy cover among sites influenced the effects of drought on the performance of each species. Drought negatively affected the growth and/or vitality of all species, but the effects were stronger and more persistent in the bryophytes than in the vascular plants. The two species associated with older forests, the moss Hylocomiastrum umbratum and the orchid Goodyera repens, suffered larger effects than the more generalist species included in the experiment. The drought reduced reproductive output in the moss Hylocomium splendens in the next growing season, but increased reproduction in the graminoid Luzula pilosa. Higher ambient soil moisture reduced some negative effects of drought on vascular plants. Both denser canopy cover and higher soil moisture alleviated drought effects on bryophytes, likely through alleviating cellular damage. Our experiment shows that boreal understory species can be adversely affected by drought and that effects might be stronger for bryophytes and species associated with older forests. Our results indicate that the effects of drought can vary over small spatial scales and that forest landscapes can be actively managed to alleviate drought effects on boreal forest biodiversity. For example, by managing the tree canopy and protecting hydrological networks. [ABSTRACT FROM AUTHOR]

Restoration of degraded habitat is frequently used in ecological compensation. However, ecological restoration suffers from innate problems of long delivery times of features shown to be good proxies for biodiversity, e.g., large dead trees. We tested a possible way to circumvent this problem; the translocation of hard-to-come deadwood substrates from an impact area to a compensation area. Following translocation, deadwood density in the compensation area was locally equivalent to the impact area, around 20 m3 ha−1, a threshold for supporting high biodiversity of rare and red-listed species. However, deadwood composition differed between the impact and compensation area, showing a need to include more deadwood types, e.g., late decomposition deadwood, in the translocation scheme. To guide future compensation efforts, the cost for translocation at different spatial scales was calculated. We conclude that translocation of deadwood could provide a cost-efficient new tool for ecological compensation/restoration but that the method needs refinement. [ABSTRACT FROM AUTHOR]

Intensive forestry practices have had a negative impact on boreal forest biodiversity; as a consequence, the need for restoration is pressing. Polypores (wood‐inhabiting fungi) are key decomposers of dead wood, but, due to a lack of coarse woody debris (CWD) in forest ecosystems, many species are under threat. Here, we study the long‐term effects on polypore diversity of two restoration treatments: creating CWD by felling whole trees and prescribed burning. This large‐scale experiment is located in spruce‐dominated boreal forests in southern Finland. The experiment has a factorial design (n = 3) including three levels of created CWD (5, 30, and 60 m3 ha−1) crossed with burning or no burning. In 2018, 16 years after launching the experiment, we inventoried polypores on 10 experimentally cut logs and 10 naturally fallen logs per stand. We found that overall polypore community composition differed between burned and unburned stands. However, only red‐listed species abundances and richness were positively affected by prescribed burning. We found no effects of CWD levels created mechanically by felling of trees. We show, for the first time, that prescribed burning is an effective measure for restoring polypore diversity in a late‐successional Norway spruce forest. Burning creates CWD with certain characteristics that differ from what is created by CWD restoration by felling trees. Prescribed burning promotes primarily red‐listed species, demonstrating its effectiveness as a restoration measure to promote diversity of threatened polypore species in boreal forests. However, because the CWD that the burning creates will decrease over time, to be functional, prescribed burns need to be applied regularly on the landscape scale. Large‐scale and long‐term experimental studies, such as this one, are invaluable for establishing evidence‐based restoration strategies. [ABSTRACT FROM AUTHOR]

Context: Both climatic extremes and land-use change constitute severe threats to biodiversity, but their interactive effects remain poorly understood. In forest ecosystems, the effects of climatic extremes can be exacerbated at forest edges. Objectives: We explored the hypothesis that an extreme summer drought reduced the richness and coverage of old-growth forest species, particularly in forest patches with high edge exposure. Methods: Using a high-resolution spatially explicit precipitation dataset, we could detect variability in drought intensity during the summer drought of 2018. We selected 60 old-growth boreal forest patches in central Sweden that differed in their level of drought intensity and amount of edge exposure. The year after the drought, we surveyed red-listed and old-growth forest indicator species of vascular plants, lichens and bryophytes. We assessed if species richness, composition, and coverage were related to drought intensity, edge exposure, and their interaction. Results: Species richness was negatively related to drought intensity in forest patches with a high edge exposure, but not in patches with less edge exposure. Patterns differed among organism groups and were strongest for cyanolichens, epiphytes associated with high-pH bark, and species occurring on convex substrates such as trees and logs. Conclusions: Our results show that the effects of an extreme climatic event on forest species can vary strongly across a landscape. Edge exposed old-growth forest patches are more at risk under extreme climatic events than those in continuous forests. This suggest that maintaining buffer zones around forest patches with high conservation values should be an important conservation measure. [ABSTRACT FROM AUTHOR]

Managed forest stands are typically younger and structurally less diverse than natural forests. Introduction of non-native tree species might increase the structural changes to managed forest stands, but detailed analyses of tree- and stand-structures of native and non-native managed forests are often lacking. Improved knowledge of non-native forest structure could help clarify their multiple values (e.g. habitat for native biodiversity, bioenergy opportunities). We studied the structural differences between the introduced, non-nativePinus contortaand the nativePinus sylvestrisandPicea abiesover young forest stand ages (13–34 years old) in managed forests in northern Sweden. We found thatP. contortastands had greater mean basal areas, tree heights, diameters at breast height, and surface area of living branches than the two native species in young stands. The surface area of dead attached branches was also greater inP. contortathanP. abies. Although this indicates greater habitat availability for branch-living organisms, it also contributes to the overall more shaded conditions in stands ofP. contorta. Only one older 87 years oldP. contortastand was available, and future studies will tell how structural differences betweenP. contortaand native tree species develop over the full forestry cycle. [ABSTRACT FROM AUTHOR]

[Display omitted] • Habitat heterogeneity and resource amount and quality were tested as indicators. • Habitat heterogeneity predicts the richness of species of conservation concern. • Dead wood and tree age predict the richness of red-listed species. • Thresholds for habitat heterogeneity score and deadwood were calculated. • Habitat heterogeneity score is useful tool in practical forestry and conservation. Reliable assessment measures are crucial for tracking changes in biodiversity and for evaluating the state of biodiversity. Two of the main drivers of biodiversity are habitat heterogeneity and resource amount. These drivers are used as proxies of biodiversity but assessing both is costly, limiting their practical use. To test which of the drivers best predicts the number and abundance of sessile species of conservation concern (including macrofungi, lichens, bryophytes, and vascular plants), we assessed forest stand heterogeneity using a method developed in Sweden ('Habitat Heterogeneity Score HHS'), and quantified the resource amount and quality of ecologically important structural variables (deadwood volume, basal area of living trees, proportion of broadleaved trees, and the age of the oldest tree in the stand). We conducted the assessments in 77 boreal conifer-dominated forest stands in two regions of Sweden. Despite some group-specific organism differences, HHS was the best predictor of both number and abundance of all species of conservation concern, regardless of the region. Further, HHS was the best predictor of red-listed species number and abundance in the southern region, while a model including the volume of deadwood and the age of the oldest tree performed best in the northern region. Deadwood (CWD) volume was the single best resource amount predictor of the number and abundance of species of conservation concern, emphasizing the critical role that dead trees have for biodiversity. In addition, we calculated threshold values for deadwood volume and HHS depicting the level above which the number of red-listed species is significantly higher, and found this value to be higher in the southern region (22.4 m3 ha−1 deadwood and a HSS value of 17) than in the north (20.0 m3 ha−1 and 16). These values can be used as guidance when identifying coniferous forests with high enough qualities to support red-listed species. To conclude, the method of assessing habitat heterogeneity presented in this study is a practical and reliable way to identify forests of high biological diversity, and can therefore be part of the toolbox for sustainable forestry in boreal forests. [ABSTRACT FROM AUTHOR]

Diversity patterns and dynamics at forest edges are not well understood. We disentangle the relative importance of edge-effect variables on spatio-temporal patterns in species richness and occupancy of deadwood-dwelling fungi in fragmented old-growth forests. We related richness and log occupancy by 10 old-growth forest indicator fungi and by two common fungi to log conditions in natural and anthropogenic edge habitats of 31 old-growth Picea abies forest stands in central Sweden. We compared edge-to-interior gradients (100 m) to the forest interior (beyond 100 m), and we analyzed stand-level changes after 10 yr. Both richness and occupancy of logs by indicator species was negatively related to adjacent young clear-cut edges, but this effect decreased with increasing clear-cut age. The occupancy of logs by indicator species also increased with increasing distance to the natural edges. In contrast, the occupancy of logs by common species was positively related or unrelated to distance to clear-cut edges regardless of the edge age, and this was partly explained by fungal specificity to substrate quality. Stand-level mean richness and mean occupancy of logs did not change for indicator or common species over a decade. By illustrating the importance of spatial and temporal dimensions of edge effects, we extend the general understanding of the distribution and diversity of substrate-confined fungi in fragmented old-growth forests. Our results highlight the importance of longer forest rotation times adjacent to small protected areas and forest set-asides, where it may take more than 50 yr for indicator species richness levels to recover to occupancy levels observed in the forest interior. Also, non-simultaneous clear-cutting of surrounding productive forests in a way that reduces the edge effect over time (i.e., dynamic buffers) may increase the effective core area of small forest set-asides and improve their performance on protecting species of special concern for conservation. [ABSTRACT FROM AUTHOR]

Abstract: The loss of natural forest habitats due to forestry is the main reason for the decline in boreal forest biodiversity of the Nordic countries of Europe. Ecological rehabilitation may provide means to recover and sustain biodiversity. We analyzed the effects of controlled burning and dead-wood creation (DWC) on the diversity of pioneer wood-inhabiting fungi in managed Norway spruce (Picea abies) forests in southern Finland. Altogether 18 stands were first subjected to a partial cutting with ordinary logging residues in form of cut stumps and treetops left on site. The subsequent rehabilitation treatments consisted of a controlled burning applied in half of the stands and three levels of dead-wood creation (5, 30 and 60m3 ha−1). The DWC involved creation of logs; felling of whole trees to mimic downed logs formed by natural disturbance processes. Each treatment was replicated three times. Inside each stand, substrates were sampled in two different biotopes; one on mineral soil and one on mineral soil with a thin peat layer. We surveyed the fungal flora on the logs (n =364) and the ordinary residue stumps (n =1767) and tops (n =845) five years after the treatments. When comparing different stands, controlled burning had a significant effect on species composition; certain species were significantly more frequent on substrates in burned stands than in unburned stands, indicating that these species were favored by controlled burning. By contrast, we found no significant effects of DWC levels or biotope on species composition or richness. When comparing different substrates, 99% of the logs hosted at least one species and the occurrence probability of certain species was significantly higher on logs than on ordinary residue stumps and tops. Yet, volume-based rarefaction analyses showed that residues were more species dense than the logs, indicating that ordinary logging residues constitute important resources for many pioneer species. We conclude that controlled burning combined with DWC have strong effects on biodiversity; it modifies the composition of the pioneer wood-inhabiting fungal species found in managed forests and may thereby also influence the further succession and diversity of the secondary fungal flora. [Copyright &y& Elsevier]

• We investigated the thermal buffering of forest retention practices. • T max was lower in retention forests than in clear-cuts but higher than in old forests. • Basal area and canopy cover were identified as key factors impacting microclimates. • Preserving tree diversity and standing deadwood enhance stands' thermal buffering. • Lying deadwood related negatively to canopy cover, likely lowering thermal buffering. Retention forestry is increasingly adopted as an alternative to clearcutting practices and involves retaining structural and compositional complexity (e.g., living and dead trees) from preharvest to postharvest. Past studies have examined the role of retention forestry in supporting various ecosystem functions and biodiversity, whilst its microclimate buffering capacity has been largely neglected. We investigated the microclimates and the underlying mechanisms of retention forests relative to clearcuts and old forests in a boreal forest landscape in central Sweden. We found that both air temperature and vapour pressure deficit (VPD) differed significantly between the forest types. Old forests consistently exhibited the most buffered forest microclimates, followed by retention forests, while clearcuts displayed the lowest. Basal area and canopy cover were identified as the key determinants influencing air temperature and VPD across the forest types. Retention practices can also impact a stand's microclimates. Specifically, maintaining diverse tree species had the potential to lower the stand's maximum temperature, given its positive association with canopy cover. Large volumes of lying deadwood were found to be negatively correlated with both basal area and canopy cover, likely contributing to increased maximum temperatures. Furthermore, standing deadwood directly lowered the maximum temperature within forest stands. Finally, edge effects were observed in the retention forests, with south-facing edges experiencing significantly higher maximum temperature and VPD compared to north-facing edges and forest interiors. These south-facing edge effects were positively associated with the difference in lying deadwood volumes between forest edges and interiors. Our findings support the positive influence of retention practices on a stand's microclimate buffering, achieved through preserving diverse tree species, standing deadwood, and implementing measures to prevent severe wind-induced tree mortality, particularly in south-facing edges (e.g. creating south-facing buffer zones). Forest managers and policy makers can utilize these results to minimize the climate-change impacts on below-canopy biodiversity and functioning. [ABSTRACT FROM AUTHOR]

The biodiversity crisis calls for immediate restoration of deteriorated and rare habitat. Due to fire suppression and intensive forest management, boreal pine forests of high conservation value are exceptionally rare. Despite decades of restoration research in boreal forests, relatively few studies have evaluated multi-taxon biodiversity response of restoration measures in pine forests. In a Scots pine experiment, we investigated biodiversity patterns of wood-inhabiting fungi and beetles a decade after restoration (prescribed burning and deadwood creation) and forest management (harvest with varying retention). We found that fungi and beetles develop differently and have distinct preferences in deadwood originating from restoration. Standing deadwood supported more species for beetles and lying deadwood for fungi and for both taxa, standing and lying deadwood harboured different species assemblages. Burned deadwood displayed less variable assemblages than unburned deadwood for both organism groups. We found that, after a decade, deadwood type and not harvest with different retention levels better explained diversity patterns of wood-inhabiting beetles and fungi in pine forests. Pine forests are naturally prone to recurring disturbances creating open light conditions. Pine-associated species are therefore likely resistant to disturbance as long as a variety of deadwood resources are present. To accommodate multiple taxa, a variety of substrate and environment types is required. Beetles benefit from standing deadwood while fungi benefit from lying deadwood. To support assemblages with both rapid and slow turnover rates, a combination of recurring restoration and leaving restored stands in the adjacent landscapes is required. • Substrate type was more influential than retention level for saproxylic biodiversity. • Logs host more fungal species than snags. • Snags host more beetle species than logs. • Deadwood types hosted different saproxylic assemblages. • Different deadwood types and decay stages are needed to support saproxylic biodiversity. [ABSTRACT FROM AUTHOR]

Setting aside small remnant patches of high biodiversity forest within managed forest landscapes is often used as conservation measure to provide a refuge and future source population of forest biodiversity, including wood-inhabiting fungal communities. Yet little is known about the long-term fungal community assembly, how these small, isolated patches change through time and how forest management in the surrounding landscape impacts traits and community functionality housed within. We applied a joint species distribution model to compare how fungal traits and communities changed over two survey periods undertaken ~20 years apart in boreal forest set-aside and natural patches. Natural patches in naturally fragmented landscapes were considered reference forests for small, remnant, near-natural forest patches in intensively managed forest landscapes. We found the majority of fungal traits converged over time between set-aside and natural patches, without changes in overall species richness. Red-listed species occurrence was initially lower in set-aside patches, but reached a comparable level of natural patches over time as a result of opposing changes in both patch types. Functional trait changes were larger in set-aside patches, but convergence was also related to opposing changes in natural patches. This is the first study to directly measure and test wood fungal community trait-environment relationships over time in small, high-conservation value forest patches. The long-term functional trait and red-listed species values of set-asides, coupled with their capacity for old-growth recovery, make them valuable focal areas for conservation. [ABSTRACT FROM AUTHOR]