Linking substrate and habitat requirements of wood‐inhabiting fungi to their regional extinction vulnerability

Loss of old‐growth forests and greatly reduced volumes of coarse dead wood in managed forests are the main reasons for the decline of many wood‐inhabiting species in Europe and elsewhere. To assess the habitat requirements and extinction vulnerability of 13 polypore species associated mainly with spruce, their occurrences were recorded on 96 521 dead‐wood objects in 331 stands along a regional gradient of forest utilization history across southern‐middle boreal Finland. The substrates studied included a variety of tree species and dead‐wood qualities investigated in both unmanaged and managed stands at different successional stages. Hierarchical logistic regression models were constructed to analyze the relationships between the occurrence probability of individual species and variables at the substrate, stand and regional scales. The substrate preferences of the polypore species studied overlapped, since most of them favored large‐diameter spruce logs in mid‐decay stages. However, only a few species were restricted to this substrate. Other species were able to use a wider range of host tree species and qualities of dead wood, including man‐made substrates that are abundant in managed forests (logging residues and stumps). Species confined to logs had a significantly lower occurrence probability in regions with the longest and most intensive forest use history. Species less specialized in their resource use showed no decline or the opposite trend. Loss of threatened species is likely if the preservation of old‐growth forests is not combined with conservation measures in managed forests. Increasing extraction of logging residues and stumps for biofuel may cause non‐threatened species to decline by reducing substrate qualities utilized by them. The hierarchical models predicted a considerable part of the variation in Species' occurrence probabilities, and therefore provide powerful tools for setting quantitative targets for management.     

Mimicking natural disturbances of boreal forests: the effects of controlled burning and creating dead wood on beetle diversity

The young successional stages of boreal forests are an important habitat for many saproxylic species. These habitats are formed by disturbances such as forest fires and they are characterized by large volumes of dead wood and sun-exposed conditions. Today, young successional stages of natural origin are very rare in Fennoscandia and there is need for restoration. We constructed a large-scale field experiment in which we studied the effects of two restoration practices on beetle diversity: controlled burning and partial harvesting with creating different volumes of dead wood. We sampled beetles with flight-intercept traps recording a total of 56,031 individuals and 755 species. The species richness and abundance of both saproxylic and non-saproxylic beetles were increased by burning and harvesting but the volume of dead wood created on harvested sites had no short-term effect on species richness or abundance. Rare species, especially saproxylic ones, preferred burned sites and a similar trend was observed among red-listed and pyrophilous species. Burning and harvesting also resulted in different species assemblages and there were some additional differences according to the volume of dead wood. We conclude that fire can be successfully used in restoration of managed boreal forests to increase species diversity and to facilitate the recovery of declined species. However, long-term monitoring is needed to clarify the effects of the restoration practices, in particular those of creating dead wood without using fire.     

Fallen retention aspen trees on clear-cuts can be important habitats for red-listed polypores: a case study in Finland

Green-tree retention is a relatively new forestry application, which aims at decreasing the negative effects of clear-cut logging on forest biodiversity. In this study, the value of retained aspens in maintaining diverse assemblages of wood-decaying fungi (polypores; Basidiomycota) on clear-cuts was investigated, after the retention trees had died, fallen and started to decay. A total of 110 fallen aspen trunks were investigated on clear-cuts and within old-growth forests in eastern Finland, southern boreal zone; and 499 records of polypores belonging to 46 species were made. The intermediately decayed trunks on a clear-cut area hosted more species and more red-listed species than did trunks within forests. Most of the polypore species with more than two records were found in both habitats. These results suggest that many aspen-associated polypores are able to live and reproduce in sun-exposed habitats, if the quality and quantity of dead wood fulfill the species-specific requirements. This unexpected result, however, may be partly due to the exceptionally great abundance of aspen in the study area. Furthermore, in the long term, the local benefits of fallen retention trees can be limited, unless the local continuity of large aspens, both living and dead, is ensured.     

Structures determining bryophyte species richness in a managed forest landscape in boreo-nemoral Europe

Forest patches with high biological value are protected as woodland key habitats (WKH), which are identified by the presence of forest structures and indicator species. However, management for conservation needs to consider also managed forests as habitats for species. In this respect, there is a need to set quantitative targets for species and structures at different landscape scales. Due to non-intensive methods of forest management used prior to 1940 in Latvia, it might be expected that large areas of forest have developed structures that can support many species characteristic of natural forests. The aim of the study was to create a model that best described the richness of bryophyte species that are characteristic of natural forests, using forest structures as explanatory factors. The structures and bryophyte communities on living trees and coarse woody debris (CWD) were described in plots along transects blindly placed in areas dominated by State forests under commercial management. Explanatory variables related to tree species composition and tree size explained 54% of the variation in WKH indicator species richness on living trees. The best explanatory factors were maximum diameter of deciduous tree species and CWD. Low richness of total bryophyte and indicator species was found on dead wood, and the amount of variation in bryophyte species richness on CWD explained by explanatory variables was low. The study indicates the importance of deciduous tree substrate in managed forests in maintaining the spatial continuity of epiphytic species diversity. However, the forests in the managed forest landscape did not support high diversity of epixylic species, even in the WKHs, due to low diversity of suitable dead wood substrate.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

Large beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>) trees as ‘lifeboats’ for lichen diversity in central European forests

The forest management practices used in central Europe in the last several centuries have led to loss of lichen diversity that may be largely attributed to a loss of substrate variability and quantity. In an attempt to obtain information enabling us to mitigate this process, we surveyed affinity of lichen species to the substrates they currently occupy in six forest areas in the Czech Republic, located between 200 and 1000 m a.s.l. Tree bases and stems represented the most important substrate for lichen species, and especially so for threatened (i.e. red-listed) species. Lichen species richness per individual tree generally increased with stem diameter, especially for beech. Stems and tree bases of large-diameter beeches provide habitats that have enabled the survival of a crucial component of the red-listed lichen species in central Europe, far outweighing other tree species. The deciduous tree species that are commonly considered as favourable for lichen diversity (e.g. maples, ash, elms) were inhabited by only a few other lichen species additional to those associated with beech. This may be due to the low frequency of these tree species in most managed forests, and also some forest reserves, at the present time. Similarly, low incidence of dead wood in managed forests has likely limited its contribution to the lichen diversity, despite the high potential for lichen diversity associated with such substrates. It is thus apparent that bark of large-diameter live beech trees comprises a keystone habitat element in the provision of lichen diversity in central European forests.     

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.     

Fast but ephemeral effects of ecological restoration on forest beetle community

Many protected areas have a long history of human intervention before being protected. In protected forests, the past land use has reduced the amount of natural structures, which are crucial substrates for thousands of species. We evaluate the short-term ecological effect of forest restoration (dead wood creation) on conifer-associated saproxylic (dead-wood dependent) beetles. More specifically, we analyze the effect of dead wood creation on the number of beetle species and individuals 1 and 5 years after restoration in spruce and pine forests, using a large-scale monitoring network over Finland. The number of saproxylic beetle species and individuals was larger at restored than at control plots both 1 and 5 years after restoration in both spruce and pine forests. Community composition in restored plots was different from control plots 1 year after restoration, but had returned towards the control plot composition 5 years after restoration, while control plots remained largely unchanged. Both in spruce and pine forests, there were more red-listed and rare saproxylic beetles in restored than in control plots 1 and 5 years after restoration. Our results indicate that restoration has an overall positive influence on saproxylic beetle diversity immediately after dead wood creation, but this effect is rather short-lived. Long term monitoring of restored dead wood is crucial in investigating successional pathways as well as biotic communities in advanced decay stages, and in fully evaluating the ecological effect of dead wood creation as a forest restoration measure.     

Diversity of wood-decaying fungi under different disturbance regimes—a case study from spruce mountain forests

Rapid destruction of forest habitats has led to the establishment of protected areas in formerly managed forests with the aim of restoring biodiversity. Conservation in spruce-dominated reserves is often contradicted by salvage logging after insect outbreaks. Here we study the community characteristics of wood decaying fungi in a high montane Norway Spruce forest with three different management types: (1) a formerly managed area disturbed by a large-scale bark beetle outbreak, (2) an area with continuous salvage logging, and (3) an old-growth forest. Bark beetle activity in the disturbed area resulted in downed wood amounts comparable to those of the old-growth forest. However, species accumulation curves for the disturbed forest were more similar to those of the logged forest than to those of the old-growth forest. This arose because of differences in the diversity of wood decay classes; wood decay in the disturbed forest was more homogeneous. Logs in the disturbed forest originated almost exclusively from bark-beetle-infested trees, but the causes of tree mortality in the old-growth forest were manifold. Although most red-listed species were clearly confined to old-growth forest, Antrodiella citrinella was most abundant in the disturbed forest. Our analysis furthermore showed that the between stand scale is the most effective unit for diversity wood-decaying fungi. We therefore suggest a conservation strategy for preserving old-growth forests and establishing protected forest stands to enhance structural heterogeneity in spruce-dominated forests. For this, a careful screening of protected areas throughout Europe is necessary to provide managers with guidelines for conservation.     

Dead trees left in clear-cuts benefit saproxylic Coleoptera adapted to natural disturbances in boreal forest

Forest management alters the pattern of forest dynamics from that in natural conditions in the boreal region. In order to examine how certain forestry measures matching natural dynamics affect forest insects, we compared assemblages of saproxylic Coleoptera on dead, standing birch trunks left behind in eight clear-cut areas with corresponding assemblages in seven mature forests in southern and eastern Finland. We used trunk-window traps for sampling. Distinct beetle assemblages were associated with the different habitats. Median numbers of species or specimens caught did not differ between closed forests and clear-cuts, but individual beetle species occurred unevenly among the habitats. Several beetle species associated with open forest habitat, e.g. burned forests or storm-damage areas, including species regarded as threatened in Finland, were found almost exclusively, in clear-cuts. Correspondingly, a number of beetle species occurring frequently in closed forests were not found in clear-cuts. We conclude that dead trunks left in the clear-cut areas may host not only generalist saproxylic species but also many beetle species specialized to warm, sun-exposed environments, and such species may not be able to survive in closed forests. Management measures matching suppressed natural disturbances are found useful in preserving diversity in managed forests.     

Importance of high quality early-successional habitats in managed forest landscapes to rare beetle species

Species adapted to early-successional forest habitats are in managed landscapes largely confined to clearcuts. To improve habitat quality on clearcuts, green tree and dead wood retention is widely applied in forestry; however, its effects on rare early-successional species have rarely been shown. We repeatedly surveyed two red-listed beetle species (Upis ceramboides and Platysoma minus) on clearcuts in a managed boreal forest landscape. We found that U. ceramboides decreased its occupancy over time while P. minus increased, indicating that red-listed species vary in their ability to successfully utilise managed habitats. We found no effect of connectivity on probability of occurrence, colonisation or extinction per clearcut. Trees retained alive improved habitat quality of clearcuts, since both species were more frequent in dead wood of such trees, in comparison to logging residues. We suggest that retention can be improved by protecting and creating dead wood as intact trees during harvesting. Rare specialist species require habitat of high quality, and consequently it is impossible to meet the requirements of these species on every clearcut. To preserve all early-successional species at a regional scale, we recommend focusing retention of green trees and dead wood to one or a few trees species on each clearcut and in each landscape.     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Influence of Forest Management on the Species Richness and Composition of Wood-inhabiting Basidiomycetes in Swiss Forests

In order to investigate the diversity of wood-inhabiting aphyllophoroid basidiomycetes in Swiss forests, 86 plots of 50 m ² were established. They harboured a total of 3339 samples of woody debris, classified according to three categories (coarse, fine, and very fine woody debris), yielding 238 species of wood-inhabiting fungi. The selected sites cover the main forest types of Switzerland and various degrees of management intensity. A multiple linear regression analysis showed that substrate variation, i.e. differences in the quality of dead wood, including volume, age, degree of decomposition and host tree species, are the most important factors influencing diversity of wood-inhabiting fungi. In addition, a Principle Coordinate Analysis highlighted differences in the fungal communities in the different forest types. The greatest fungal species richness is found on thermophilic deciduous tree and woody shrub species. Fine and very fine woody debris, even present in intensively managed forests, often serve as important refuges for many species. Forests with a recent management intervention were found to be either species poor or species rich. Possible reasons for these differences may lay in forest size and landscape fragmentation, the distance to the nearest species pool or microclimatic factors. In Switzerland intensively managed forests harbour significantly less wood-inhabiting, aphyllophoroid fungi than non-managed or extensively managed forests. This is the case in both deciduous forests and in conifer forests. However, occasionally intensively managed forest will also harbour rare and endangered species.     

Perennial polypores as indicators of annual and red-listed polypores

Many polypores are specialized in their requirements for substrate and environment, and they have been suggested to indicate the continuity of coarse woody debris or naturalness of a forest stand. However, the use of polypores as indicators of conservation value is restricted by the temporally limited appearance of annual fruit bodies. We studied whether the species richness of perennial polypores (perennials) can be used to predict the species richness of annual or annual red-listed polypores (annuals). Our data included 1471 separate datasets (sample plots or larger inventoried areas) in different parts of Finland and Russian Karelia, ranging from the southern to northern boreal zone. At the large scale (the whole area) the number of perennials explained about 70% of the variation in the number of annuals, and about 67% in the number of red-listed annuals. A minimum set of 40–60 perennial occurrences gave a reliable estimate on the species richness of annuals, and 60–80 occurrences on the species richness of red-listed annuals. The richness of perennials predicted the richness of annuals and, in particular, richness of red-listed annuals, better than the size of inventoried area. According to our results, perennial polypores can be used as a surrogate for overall polypore species richness in natural and seminatural boreal forests, but the predictive power is weaker in managed forests. In addition, the relationship between the perennial and annual species seems to differ in different vegetation zones, management types and forest types. Due to this variation direct application of the indicator values derived from different vegetation zones and management or forest types are not recommended. Since perennials are easier to identify than annuals, detectable throughout the year, and have much smaller year-to-year variation, their use as an indicator group seems to offer advantages regarding the timing and cost-efficiency of inventories.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Forests regenerating after clear-cutting function as habitat for bryophyte and lichen species of conservation concern

The majority of managed forests in Fennoscandia are younger than 70 years old but yet little is known about their potential to host rare and threatened species. In this study, we examined red-listed bryophytes and lichens in 19 young stands originating from clear-cutting (30-70 years old) in the boreal region, finding 19 red-listed species (six bryophytes and 13 lichens). We used adjoining old stands, which most likely never had been clear-cut, as reference. The old stands contained significantly more species, but when taking the amount of biological legacies (i.e., remaining deciduous trees and dead wood) from the previous forest generation into account, bryophyte species number did not differ between old and young stands, and lichen number was even higher in young stands. No dispersal effect could be detected from the old to the young stands. The amount of wetlands in the surroundings was important for bryophytes, as was the area of old forest for both lichens and bryophytes. A cardinal position of young stands to the north of old stands was beneficial to red-listed bryophytes as well as lichens. We conclude that young forest plantations may function as habitat for red-listed species, but that this depends on presence of structures from the previous forest generation, and also on qualities in the surrounding landscape. Nevertheless, at repeated clear-cuttings, a successive decrease in species populations in young production stands is likely, due to increased fragmentation and reduced substrate amounts. Retention of dead wood and deciduous trees might be efficient conservation measures. Although priority needs to be given to preservation of remnant old-growth forests, we argue that young forests rich in biological legacies and located in landscapes with high amounts of old forests may have a conservation value.     

The white-backed woodpecker: umbrella species for forest conservation planning?

In northern Europe, a long history of land use has led to profound changes within forest ecosystems. The white-backed woodpecker (Dendrocopos leucotos) is one of several specialised forest species whose populations have declined. Conservation management directed at this species’ habitat has made it a de facto umbrella species for conservation of the biodiversity associated with forests rich in deciduous trees and dead wood. We assessed empirically the value of the white-backed woodpecker as an indicator and umbrella species in central Sweden. Occurrence of the woodpecker in breeding bird atlas squares (5 × 5 km²) indicated high species richness of forest birds, particularly species of special conservation concern, which included on average 13% more species in squares with than without the woodpecker. The number of red-listed cryptogam species expected to benefit from conservation actions directed at white-backed woodpecker habitats was higher in squares where the woodpecker bred compared to where is was absent. However, no such pattern was found for red-listed beetles, a group with very few records in the studied squares. White-backed woodpecker occurrence was positively associated with the current area of deciduous and mixed forest of high conservation value. Considering its indicator value, its specialised habitat requirements and its potential as a communication tool, using the white-backed woodpecker as an umbrella species may provide a coarse filter for the conservation of several other deciduous forest species. However, focusing solely on white-backed woodpecker habitat may not provide for the conservation of all such species, which stresses the need for a suite of complementary planning approaches.     

Large proportion of wood dependent lichens in boreal pine forest are confined to old hard wood

Intensive forest management has led to a population decline in many species, including those dependent on dead wood. Many lichens are known to depend on dead wood, but their habitat requirements have been little studied. In this study we investigated the habitat requirements of wood dependent lichens on coarse dead wood (diameter >10 cm) of Scots pine Pinus sylvestris in managed boreal forests in central Sweden. Twenty-one wood dependent lichen species were recorded, of which eleven were confined to old (estimated to be >120 years old) and hard dead wood. Almost all of this wood has emanated from kelo trees, i.e. decorticated and resin-impregnated standing pine trees that died long time ago. We found four red-listed species, of which two were exclusive and two highly associated with old and hard wood. Lichen species composition differed significantly among dead wood types (low stumps, snags, logs), wood hardness, wood age and occurrence of fire scars. Snags had higher number of species per dead wood area than logs and low stumps, and old snags had higher number of species per dead wood area than young ones. Since wood from kelo trees harbours a specialized lichen flora, conservation of wood dependent lichens requires management strategies ensuring the future presence of this wood type. Besides preserving available kelo wood, the formation of this substratum should be supported by setting aside P. sylvestris forests and subject these to prescribed burnings as well as to allow wild fires in some of these forests.     

Community Turnover of Wood-Inhabiting Fungi across Hierarchical Spatial Scales

For efficient use of conservation resources it is important to determine how species diversity changes across spatial scales. In many poorly known species groups little is known about at which spatial scales the conservation efforts should be focused. Here we examined how the community turnover of wood-inhabiting fungi is realised at three hierarchical levels, and how much of community variation is explained by variation in resource composition and spatial proximity. The hierarchical study design consisted of management type (fixed factor), forest site (random factor, nested within management type) and study plots (randomly placed plots within each study site). To examine how species richness varied across the three hierarchical scales, randomized species accumulation curves and additive partitioning of species richness were applied. To analyse variation in wood-inhabiting species and dead wood composition at each scale, linear and Permanova modelling approaches were used. Wood-inhabiting fungal communities were dominated by rare and infrequent species. The similarity of fungal communities was higher within sites and within management categories than among sites or between the two management categories, and it decreased with increasing distance among the sampling plots and with decreasing similarity of dead wood resources. However, only a small part of community variation could be explained by these factors. The species present in managed forests were in a large extent a subset of those species present in natural forests. Our results suggest that in particular the protection of rare species requires a large total area. As managed forests have only little additional value complementing the diversity of natural forests, the conservation of natural forests is the key to ecologically effective conservation. As the dissimilarity of fungal communities increases with distance, the conserved natural forest sites should be broadly distributed in space, yet the individual conserved areas should be large enough to ensure local persistence.