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In September 2011, an international team of cryptogam experts visited seven national nature reserves in five mountain areas of Slovakia: Havesova and Stužica in the Poloniny Mts., Vihorlat in the Vihorlatske vrchyMts., Oblik in the Slanske vrchyMts., Dobrocský prales and Klenovský Vepor in the Veporske vrchy Mts. and Badinsky prales in the Kremnicke vrchy Mts. The reserves were selected to represent examples of the best protected old-growth beech forests in the country. The aim was to study the diversity of wood-inhabiting fungi on fallen beech logs and epiphytic lichens on standing beech trees. In total, 215 fungal species and 128 lichens were recorded on beech wood and bark, and 27 fungi and 26 lichens on additional substrates. The site of the highest conservation value is Stužica with 126 fungi and 79 lichens recorded on beech, of which 12 fungi and 19 lichens are indicators of high nature conservation value. Combined with historical records, a total of 19 non-lichenised fungal indicators are now reported from the site, making it the highest ranked natural beech forest in Europe. The second most important reserve for fungal diversity is Havesova with 121 species, including 14 indicator species recorded on beech wood. For lichens, the second most important reserve is Klenovský Vepor with 69 species including 18 lichen indicators recorded on beech. Nine fungus species are here reported as new to Slovakia: Asterostroma medium, Entoloma hispidulum, E. pseudoparasiticum, Gloeohypochnicium analogum, Hohenbuehelia valesiaca, Hymenochaete ulmicola, Hypocrea parmastoi, Melanomma spiniferum and Scutellinia colensoi. Lichen species Alyxoria ochrocheila is reported as new to Slovakia and Lecanographa amylacea, which was considered extinct in the Slovak Red list, was also recorded. This is the first list of wood-inhabiting fungi and epiphytic lichens of old-growth beech forests in Slovakia, and hence an important contribution to the exploration of biodiversity in Slovakia.

Among the factors that affect the conservation efficiency of protected areas, lack of connectivity is considered as one of the main problems. In this study, we assessed the influence of connectivity of European beech forest reserves on wood-inhabiting fungal communities, compared to the influence of local factors. To address this topic, we used a data matrix consisting of 344 fungal species on 1571 resource units (i.e. fallen beech logs, including their standing snags) sampled in 42 European beech forest reserves. Our results show that connectivity has significant effects on wood-inhabiting fungal communities in European beech forest reserves, and that the effectiveness of reserves for maintaining the wood-inhabiting fungal diversity is compromised by habitat fragmentation. Connectivity at small scales (measured as the area of the reserve) had a strong influence on the occurrence of indicator species and was also critical for the number of species at a resource. Connectivity at larger scales (connectivity to surrounding beech forests) seemed to be particularly critical for the community composition both at resource and reserve levels. In line with previous research, we found other covariates such as size of the resource units and annual temperature range to positively influence wood-inhabiting fungal species richness. The effects of habitat fragmentation were especially strong in western and northern European regions where the smallest and more isolated reserves were located. We propose that an effective conservation strategy for wood-inhabiting fungi should focus on increasing the areas of the present reserves as well as conserving new reserves in the proximity of the existing ones.

Currently we have only a limited understanding of the evolutionary and ecological significance of reproductive traits of fungi. We compared data on fruit body size, spore size and shape between saprotrophic and mutualistic (ectomycorrhizal) fungi in Northern and Central Europe. Lifestyle and reproductive traits showed strong phylogenetic signals. A phylogenetically informed analysis demonstrated that saprotrophs produce on average smaller fruit bodies than mutualistic species. The two guilds, however, do not differ in spore size. Overall this suggests that fruit bodies of ectomycorrhizal fungi produce on average more spores than saprotrophic fungi. We argue that this difference is related to resource availability: ectomycorrhizal fungi receive carbon from their hosts and, therefore, evolution favours large fruit bodies, whereas the fruit body size of saprotrophic fungi might have responded to resource availability and the distribution and size of resource patches.

Increasing species richness of light demanding species in forests may not be a conservation concern if we accept a macroecological and evolutionary baseline for biodiversity. Most of the current biodiversity in Europe has evolved in the Pleistocene or earlier, and in ecosystems markedly influenced by dynamic natural processes, including grazing. Many threatened species are associated with high-light forest environments such as forest glades and edges, as these have strongly declined at least partially due to the decline of large herbivores in European forests. Hence, moderate grazing in forests should be an ecological baseline and conservation target rather than a concern.

Sexual spores are important for the dispersal and population dynamics of fungi. They show remarkable morphological diversity, but the underlying forces driving spore evolution are poorly known. We investigated whether trophic status and substrate associations are associated with morphology in 787 macrofungal genera. We show that both spore size and ornamentation are associated with trophic specialization, so that large and ornamented spores are more probable in ectomycorrhizal than in saprotrophic genera. This suggests that spore ornamentation facilitates attachment to arthropod vectors, which ectomycorrhizal species may need to reach lower soil layers. Elongated spore shapes are more common in saprotrophic taxa, and genera associated with above ground substrates are more likely to have allantoid (curved elongated) spores, probably to lower the risk of wash out by precipitation. Overall, our results suggest that safe arrival on specific substrates is a more important driver of evolution in spore morphology than dispersal per se.

Aim: Aphyllophoroid fungi are associated with plants, either using plants as a resource (as parasites or decomposers) or as symbionts (as mycorrhizal partners). In spite of their strong association with plants, it is unknown how much plant distributions determine their biogeographical patterns compared with environmental factors such as climate and human land use. In this study, our aims are to (1) describe the spatial diversity patterns of aphyllophoroid fungi in Europe and (2) identify the factors shaping these patterns. Location: Europe, as well as the adjacent Subarctic to Arctic islands (Greenland, Faroe Islands, Iceland, Svalbard), Palestine and the south-east coast of the Caspian Sea. Methods: We compiled a dataset consisting of 14,030 fruitbody occurrences of 1,491 aphyllophoroid fungal species from 39 geographical areas (17 countries) belonging to eight biogeographical regions. We assessed the differences in fungal species richness and overall diversity and its nestedness and turnover components across biogeographical regions of Europe, as well as between southern and northern Europe (based on geographical latitude of 50 degrees as threshold). We used cluster and ordination analyses to classify the European aphyllophoroid communities biogeographically and evaluated the importance of climate, host-tree species, topography and human land-use intensity in explaining biogeographical variation. Results: The importance of biogeographical regions in determining European aphyllophoroid fungal communities varies for different diversity components. Species richness and nestedness are best explained by plant-based biogeographical regions, whereas overall beta diversity and species turnover are driven mostly by variation in climate, and nestedness mostly by tree species occupancy. Beta-diversity patterns of aphyllophoroid fungi do not differ between southern and northern Europe. Main conclusions: At the continental scale, aphyllophoroid fungi are less shaped by historical legacies than vascular plant and animal communities, and trends of overall beta diversity in southern and northern Europe are similar to patterns found for bryophytes.

Aim Fungi are drivers of wood decay in forested ecosystem, while bryophytes use dead wood as a platform for their autotrophic lifestyle. We tested the hypothesis that fungal communities on beech logs are mainly structured by substrate quality, while bryophyte communities are structured by climatic gradients. In addition, we tested whether community structure in both organism groups is altered along a gradient from nearly pristine forest to forests heavily affected by management and human disturbance in the past. Location Europe. Methods We surveyed 1207 fallen beech logs in 26 of the best-preserved forest stands across six European countries, representing a gradient in overall naturalness of the forest landscape. Recorded species were classified into ecological guilds. Indirect ordination and variation partitioning was used to analyse the relationship between species composition and environmental variables, recorded at log or site level. Results In total, 10,367 bryophyte and 15,575 fungal records were made, representing 157 and 272 species, respectively. Fungal communities were more clearly structured by substrate quality than were bryophyte communities. In both groups a distinct turnover in species composition was evident along a longitudinal gradient from Central to Western Europe. Fungi specialized in trunk rot and specialized epixylic bryophytes were scarcely represented in Atlantic regions, and partly replaced by species belonging to less specialized guilds. Variables related to climate and forest conditions were confounded along this main geographical gradient in community composition. Main conclusions We found that bryophyte and fungal communities co-occurring on fallen beech logs in European beech forest reserves differed in their responses to biogeographical drivers and local-scale habitat filters. Both groups responded to major gradients in climate and forest conditions, but the loss of specialist guilds in degraded forest landscapes points to a functionally important effect of forest landscape degradation at the European continental scale.

Intensive forest management creates habitat degradation by reducing the variation of forest stands in general, and by removing old trees and dead wood in particular. Non-intervention forest reserves are commonly believed to be the most efficient tool to counteract the negative effects on biodiversity, but actual knowledge of the conservation efficiency is limited, especially for recent reserves. The structure of ecological communities is often described with measures of nestedness, beta diversity and similarity between communities. We studied whether these measures differ among forest reserves with different management histories. For this purpose, we used a large data set of wood-inhabiting fungi collected from dead beech trees in European beech-dominated forest reserves. The structure of fungal assemblages showed high beta diversity, while nestedness and similarity was low. During the decomposition process of trees beta diversity between the communities occupying different trees increased in natural, but not in previously managed sites. Effects of management and decay process on nestedness were complex. We argue that the detected differences most likely reflect historical effects which have extirpated specialized species from the local species pools in managed sites, and resulted in more homogeneous communities in managed sites. It is alarming that community structure is affected the most in the latest decay stages where the decay process turns the dead wood into litter, and which is thus the interface between the wood decay and the litter-decaying ecosystem. The effects of simplified communities in late decay stages on soil biodiversity should be studied.

Biodiversity surrogates are commonly used in conservation biology. Here we review how fungi have been used as such in forest conservation, emphasizing proposed surrogate roles and practical applications. We show that many fungal surrogates have been suggested based on field experience and loose concepts, rather than on rigorously collected scientific data. Yet, they have played an important role, not only in forest conservation, but also in inspiring research in fungal ecology and forest history. We argue that, even in times of ecosystem oriented conservation planning and molecular tools to analyze fungal communities, fruit bodies of macrofungi have potential as convenient conservation shortcuts and easy tools to communicate complex biodiversity for a broader audience. To improve the reliability of future fungal surrogates we propose a three step protocol for developing evidence based schemes for practical application in forest conservation. peerReviewed

Species occurrence observations are increasingly available for scientific analyses through citizen science projects and digitization of museum records, representing a largely untapped ecological resource. When combined with open-source data, there is unparalleled potential for understanding many aspects of the ecology and biogeography of organisms. Here we describe the process of assembling a pan-European mycological meta-database (ClimFun) and integrating it with open-source data to advance the fields of macroecology and biogeography against a backdrop of global change. Initially 7.3 million unique fungal species fruit body records, spanning nine countries, were processed and assembled into 6 million records of more than 10,000 species. This is an extraordinary amount of fungal data to address macro-ecological questions. We provide two examples of fungal species with different life histories, one ectomycorrhizal and one wood decaying, to demonstrate how such continental-scale meta-databases can offer unique insights into climate change effects on fungal phenology and fruiting patterns in recent decades.

Woody plants host diverse communities of associated organisms, including wood-inhabiting fungi. In this group, host effects on species richness and interaction network structure are not well understood, especially not at large geographical scales. We investigated ecological, historical and evolutionary determinants of fungal species richness and network modularity, that is, subcommunity structure, across woody hosts in Denmark, using a citizen science data set comprising > 80 000 records of > 1000 fungal species on 91 genera of woody plants. Fungal species richness was positively related to host size, wood pH, and the number of species in the host genus, with limited influence of host frequency and host history, that is, time since host establishment in the area. Modularity patterns were unaffected by host history, but largely reflected host phylogeny. Notably, fungal communities differed substantially between angiosperm and gymnosperm hosts. Host traits and evolutionary history appear to be more important than host frequency and recent history in structuring interactions between hosts and wood-inhabiting fungi. High wood acidity appears to act as a stress factor reducing fungal species richness, while large host size, providing increased niche diversity, enhances it. In some fungal groups that are known to interact with live host cells in the establishment phase, host selectivity is common, causing a modular community structure.

Summary In a recent paper in Fungal Ecology, Gange et al. (2011) concluded that the wood-inhabiting basidiomycote Auricularia auricula-judae has widened its host range and changed its phenology since 1980 in the study area in southern England, most likely as a response to global warming. At the same time the authors report, that the number of macrofungi fruiting on Sambucus nigra, the most common host of A. auricula-judae, has tripled, while another fungus preferring S. nigra, Hyphodontia sambuci, has remained constant. We challenge these conclusions and point out that the reported changes may just as well reflect a change in foraying quality over time. To support this statement we present an analysis of data from an ongoing mapping project in Denmark. We show that species accumulation curves and species richness estimators are highly context specific, and that random sampling by amateurs and specialists provide very different representations of actual patterns in nature. We show that the three species analysed by Gange et al. (2011) behave very differently in this context, and are hence not straight forward to compare over time based on data from amateur forays. We conclude that more rigorous standardising procedures are needed to reach sound scientific conclusion based on analyses of foray data over time.

Biodiversity information databases and platforms have seen considerable progress in recent years. They have a high potential in conservation science in general, but may be even more revolutionary in relation to poorly known species groups such as fungi, whose practical conservation work has been jeopardised by scattered and poorly controlled information. We review the tradition of collecting information on species occurrences in mycology and discuss the characteristics of the present fungal biodiversity information databases. With a special focus on population trend monitoring of fruit body producing macrofungi, we emphasise several unrealised opportunities of these databases and point out some relevant future directions for them. As especially important, we see the more effective utilisation of citizen science effort and combining the traditional database information with the one derived with modern molecular methods. Also, we emphasise the importance of information on collection effort, including the use of GPS based tracking data, along with the observations.

We review the state of fungal conservation in Europe. Despite a large public interest in fungi, they are often insufficiently considered in the conservation initiatives of most countries and not adequately heeded in international biodiversity agreements. We attribute this to the generally low profile of fungi among conservation stakeholders and decision makers together with limited efforts of mycological scientists to put mycological knowledge into a conservation context. Recent advances in mycological knowledge, taxonomy, distribution, ecology and threats now categorically enable fungi to be included within national and European conservation agendas. 33 European countries have produced fungal Red-Lists reporting the status of macrofungi and these are official in 20 countries. These lists indicate that at least 10 % of European larger fungi are threatened, mainly due to changing land use and nitrogen deposition. Fungal biodiversity may benefit from many general conservation efforts, but many specific fungus values are also overlooked. We advocate increased interaction between scientists and conservation coordinators and practitioners, greater promotion of fungi and their conservation and ecosystem service values by mycologists, the production of a European fungal Red-List and the need to integrate fungi with animals and plants in conservation issues.

Identification of trees with key microhabitats as well as knowledge of their ecological formation is important for the conservation of epiphytic bryophytes and lichens on beech. Based on the hypothesis that certain types of stem damage are crucial for the occurrence of epiphytes of conservation concern, we surveyed 145 beech trees (57–280 years) for different types of stem damage and analysed their relationship to other tree characteristics, epiphyte species and wood-inhabiting fungi in a forest landscape on acid soils. Three main types of stem damage were identified; canker, rot hole and surface rot. The incidence of rot holes was highest on trees with a low growth rate during the last 50 years, but was unrelated to tree size or age per se. Bark pH was significantly higher below than above rot holes, whereas no such relationship was found for cankers. Wood mould from rot holes had a very high pH, explaining the higher bark pH below this type of damage. The number of epiphyte species of conservation concern was strongly positively related to rot holes and high bark pH. Cankers had a weaker, yet significant positive effect. The fungus Psathyrella cernua was associated with the rot holes and is suggested to be a key species involved in creating this microhabitat. We conclude that slow-growing trees with rot holes are important for the conservation of epiphytes and should be selected as retention trees in managed forests. The study also highlights a potential for habitat restoration by inducing artificial damage and inoculation of decay fungi in beech trees.

Patterns of biodiversity, environment and human impact were studied in 57 sample plots in an 1,178 ha forest area in a rural mountain area of Nepal that is administrated by the Annapurna Conservation Area Project. Alpha-, beta- and gamma-diversity was measured or estimated for six groups of organisms: trees, shrubs, climbers, herbs, polypores and mycorrhizal fungi, and the recorded patterns were correlated with a set of environmental variables. Human impact in terms of fuelwood collection, selective cutting and grazing was found to influence species diversity patterns in all organism groups. Species richness of trees, climbers and polypores at plot level (alpha-diversity) generally responded negatively to human impact, whereas species richness of herbs and shrubs showed a positive relation. Species turnover (beta-diversity), measured as length of the DCA first axis, was significantly correlated to distance to village for all species groups. This indicates that the human impact is very important for the biodiversity patterns in the study area, and that biodiversity connected to undisturbed forest habitats are potentially threatened in the area. The results are discussed in the context of practical conservation. A proposal for future management zones addressing protection of biodiversity without limiting the local use of the forest resource is put forward.

Tricholoma scalpturatum is an ectomycorrhizal fungus that forms symbioses with roots of diverse trees and shrubs. It is commonly encountered in a wide range of habitats, across temperate ecosystems. A previous study has revealed a high genetic diversity at a local scale, and ruderal abilities. To examine genetic structure at a large geographical scale, a total of 164 basidiocarps were collected from 30 populations located in Western Europe, from Spain to Scandinavia. These samples were analyzed by three molecular methods with different levels of resolution: inter-simple sequence repeats (ISSRs), restriction fragment length polymorphisms (RFLPs) in the rDNA internal transcribed spacer (ITS), and ITS sequence analysis. Considerable genetic variation was found, and the morphospecies was separated into two genetic groups that were distinct from each other. The ISSR data and the relatively low percentage value (96%) of shared sequence polymorphisms in the ITS between isolates from the two groups, strongly suggest cryptic species and long-lasting separation. No geographical exclusion was detected for these two widely distributed taxa. However, high estimates of population differentiation were observed in each group, including between populations less than a few kilometers apart. This result provides evidence for limited gene flow and/or founding effects. It also indicates that T. scalpturatum does not constitute a random mating population, and the hypothesis of endemism cannot be excluded for this cosmopolitan wind-dispersed fungus.

Bryophyte and fungal communities were investigated on fallen trees representing seven deciduous tree species in a mixed near natural nemoral forest. Bryophytes were represented by 41 taxa, including several very frequent species. Of the 296 fungal species, most were recorded with very low frequency and the share of high frequent species was much lower than among the bryophytes. Species turnover was bigger in the fungal communities, compared to the bryophyte communities, and related to a higher extent to measured differences in environmental conditions. Tree species diversity was found to be an important factor for fungal species composition, while only small differences in bryophyte species composition were found between the different tree species. On the other hand bryophyte species richness showed distinct relations to tree species and microclimatic variables, a tendency which was not evident for fungal diversity. It is concluded that the two organism groups to some extent differ in their conservation demands. Thus, conservation of wood-inhabiting bryophytes requires prioritising of large, coherent forest stands in which a stable humid microclimate and a reasonable supply of dead wood is secured. Successful conservation of fungi requires that substantial amounts of dead wood are left for natural decay in a variety of natural forest environments representing different tree species, so that heterogeneity in dead wood types is secured.

Macrofungal diversity was investigated on 281 decaying beech trees distributed across 14 forests in Denmark, based on sporocarp inventories. Two aspects of diversity were considered, i.e., species richness per fallen tree and the incidence of red-listed species occurrence per tree. For both diversity measures the effects of both tree and site variables were tested. In total, 319 fungal species were identified, including 28 red-listed. Decay stage and wood volume were identified as key variables influencing species richness as well as red-listed species incidence. Red-listed species, however, showed a preference for more decayed trees than non red-listed species. Further, red-listed species incidence was found to be significantly higher on broken trees, compared to fallen trees with a distinct root-plate, indicating tree death cause to be important for some red-listed species. The relations between diversity measures and site variables were conflicting. Species richness per tree decreased with increasing maximum tree age and dead wood continuity, possibly a consequence of competitive exclusion of unspecialised opportunistic species in old-grown stands. For red-listed species the opposite trend was evident, and it is concluded that forest history may have fundamental effects on the community structure of wood-inhabiting fungi. Accordingly, simple species richness may be a misleading conservation measure if the aim is to conserve the most threatened aspects of forest biodiversity.

The effects of exudates from uncolonized and from partly decayed beech wood on the extension rates of 16 later stage decay fungi were investigated. The partly decayed wood had been colonized by the pyrenomycete Eutypa spinosa, or the basidiomycetes Fomes fomentarius, Stereum hirsutum, and Trametes versicolor, all known as common early decay agents in European beech forests. Sterilized wood pieces were placed onto 0.5% malt agar, opposite to small agar plugs containing the test fungi. The latter showed very variable and species-specific growth responses to the various wood types. The presence of uncolonized wood stimulated extension rates in many species, whereas the four previously decayed wood types had variable stimulatory or inhibitory effects. Wood decayed by S. hirsutum resulted in reduced extension rate, delayed growth, or total inhibition in the majority of species, thus it is suggested that this species uses secondary metabolites in a defensive strategy. A single species was, however, stimulated in the presence of S. hirsutum-decayed wood. In contrast, the presence of wood decayed by F. fomentarius was stimulatory to 45% of the species. The other previously decayed wood types generally resulted in more variable responses, depending upon species. The results are discussed in an ecological context and it is suggested that the exudates from the partly decayed wood that are responsible for the reported effects may function as infochemicals, structuring microbial communities in wood.

The importance of different fractions of coarse wood debris (cwd) for species diversity of wood-inhabiting fungi was investigated in near-natural Danish beech stands. Species number per tree increased significantly with increasing tree size, pointing out large trees to be most valuable for fungal diversity if single samples are compared. Rarefaction curves, evaluating the importance of different cwd fractions in a cumulative space, revealed a different and more complex picture. Rarefaction curves based on wood volume showed small trees and branches to host more species per volume unit than larger trees and logs, respectively, while snags were the most species-poor fraction. Surface-based curves showed species density to be rather similar among cwd types, though species density still decreased slightly with tree size. These results are interpreted to reflect a combination of two factors: firstly, small diameter cwd represent a larger surface area per volume and hence more space for fungal sporocarps, than large diameter cwd. This ‘surface area factor’ explains the high degree of similarity of the surface-based rarefaction curves. Secondly, a collection of small diameter cwd involves more separate units than an equal volume of large diameter cwd, and represents thereby more individual cases of fungal infection and, most likely, more variation in environmental conditions. The effect of this ‘number of item factor’ is reflected in the slightly increasing species density per surface area with decreasing tree size. Richness patterns of red-listed and non-red-listed species were found to be strikingly similar across cwd types, and a general preference for large logs among red-listed species was hence absent. An individual look at the most frequent encountered red-listed species revealed substrate preference patterns to occur in three of six species, of which one, the heart-rot agent Ischnoderma resinosum , preferred large logs. Based on these results, it would be obvious to conclude that local fungal species diversity is most efficiently increased in managed forests if small diameter cwd is prioritised for natural decay. However, small diameter wood appear to be unable to support heart-rot agents and other species depending on a long and diverse infection history and thus the integrity of saproxylic communities may be seriously undermined if only small diameter cwd is left for decay. Therefore, we strongly recommend that whole, naturally dead trees, representing the full range of cwd habitats, are prioritised for natural decay in managed forests whenever possible.

A national strategy for "Natural Forests and Other Forest Types of High Conservation Value in Denmark" was launched in 1992 and by the year 2000 6500 ha of forest was protected as strict natural forest reserves. Descriptions of potential sites, emphasizing forest composition and history, served as the main decision basis in the design of the reserve network. According to the strategy the area of untouched or specially managed forests will be increased considerably during the next few decades, but even though one of the most important purposes of the strategy is the protection and restoration of forest biodiversity no measurable goals are given in this respect. This study pays direct attention to biodiversity as reflected by the presence/absence of indicator species. It focuses on wood-inhabiting fungi since this group of organisms is especially dependent on low-intervention forests and may indicate potential reserve areas with high diversity of other saproxylic organisms. Subject to a budget constraint, a...

An analysis of factors influencing the diversity of macrofungi fruiting on decaying beech logs at site level is presented. Variables related to log size and shape and decay stage were found to explain up to 56% of the variation in total species richness and 42% of the variation in the richness of threatened (red-listed) species. Inclusion of variables relating to the vernal flora and the degree of soil contact further increased the explained variation in total species richness to 71%, but these variables were non-significant with respect to red-listed species. However, inclusion of the variable log type, distinguishing uprooted logs, logs broken at root neck and logs broken 1–7 and 8–15 m above ground, increased the amount of explained variation in richness of red-listed species to 50%. Among the log size and shape variables, the number of bole forks was superior in describing the variation in both total and red-listed species richness. Accordingly, forked trees should preferably be selected for decay in order to improve biodiversity, since they support comparably higher species diversity than unforked logs and have limited economical value. The importance of log type for the richness of threatened species appears to be connected with the occurrence of certain non-dominant primary decayers, causing heart rot, subsequently allowing the establishment of red-listed species. Accordingly, it is suggested that a high diversity of primary decayers may be a key to the conservation of wood decaying fungi. Microclimatic variables were found to have a limited effect on fruit body diversity on the studied logs; however, the microclimatic regime is discussed as an important factor in relation to management of dead wood for fungal biodiversity.

The occurrence of fungi and slime moulds on 70 decaying beech logs was surveyed based on the presence/absence of sporocarps. In total 277 species of fungi and 25 of slime moulds were recorded and summarised in a log/species datamatrix. The structure of the datamatrix was analysed using detrended correspondance analysis (DCA). The ecological nature of the gradients expressed by the first three DCA-axes was then investigated by environmental and log related variables. The first and strongest gradient corresponded to changes in the community development during the decay process. The second gradient was complex and corresponded to both decay rate and microclimatic stress. The third, rather weak, gradient was influenced by soil conditions. The gradients are discussed in a context of fungal ecological strategy theories. A model generalised from the community development on the studied logs is proposed.

Nature-based silviculture is a promising approach to meet the criteria for sustainable forestry. This brings the natural forest into focus as a basic reference for forest management. The present study focuses on the structural dynamics of a near-natural temperate deciduous forest in Denmark as a reference for forestry. The study was conducted in Suserup Skov (55° 22′N, 11° 34′E, 19.2 ha), an ancient woodland with a long history of low human impact. It is a mixed deciduous forest dominated by Fagus sylvatica L. and Fraxinus excelsior L. with some Quercus robur L. and Ulmus glabra Huds. The dynamics and structures of Suserup Skov can be described in time and space according to the mosaic-cycle concept. A specific model of the forest cycle in Suserup Skov was developed, including five sequential phases. The typical duration of each phase was determined: innovation (14 years), aggradation (56 years), early biostatic (96 years), late biostatic (108 years) and degradation (10 years) phases, equalling some 284 years for the full cycle. In principle, the forest cycle takes place at any given patch of the forest, asynchronous from patch to patch, resulting in a shifting mosaic of the constituent phases. The mosaic, mapped in a 10.65-ha plot in 1992, had an average patch size of 839 m2, ranging from 100 to 12 730 m2. The patches of the shorter phases (innovation, degradation) were on average considerably smaller than the patches of the longer phases, which can be explained partly by fusion of neighbouring patches. The aggregate areas of each phase were almost directly proportional to their respective duration, suggesting that the shifting mosaic was close to the steady state. The disturbance regime and disturbance history of Suserup Skov is discussed. It is concluded that several disturbances of smaller scale had occurred during the past centuries and that the most influencing recent disturbance has been the long period of human influence by rural, woodland management (pannage, pasture and coppicing) before the enclosure of the forest in 1807. The Suserup Skov example shows that a temperate deciduous forest under a relatively calm disturbance regime can develop into a very fine-grained mosaic, apparently approaching the shifting mosaic steady state within a rather small area. The structural steady state in Suserup Skov occurs, even though the overall species composition is still under successional change. Further, `climax microsuccession' from Fraxinus to Fagus occurs as an integral part of the forest cycle in Suserup Skov. Cyclic and directional processes are intermingled in Suserup Skov, highlighting the complex relationships between the concepts of succession and climax. The paper finally exemplifies how natural forests can be used as a reference for silviculture, nature conservation and landscape planning.

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.

Distribution patterns of fungi and other organisms are influenced by several factors over various scales in time and space. With their microscopic, often wind-dispersed spores, fungi are potentially able to disperse between continents, and many wood-inhabiting fungi with broad host ranges have been thought to have more or less global distribution patterns. With increased insight in fungal taxonomy outside Europe, and the use of molecular methods and mating experiments, it is becoming increasingly clear that many species, previously thought to have a wide distribution, actually circumscribe several biological taxa, each with a much more restricted distribution. Thus, continental drift, glaciations and other long-term geological and geographical factors have more impact on the current distribution patterns of fungi than believed earlier. At the continental scale, climate and host tree distribution patterns are important factors influencing the distribution of wood-inhabiting species, and climate change is likely to affect the distribution patterns of wood-inhabiting fungi considerably in the coming centuries. In the short time, man has had a strong impact on the abundance and distribution of dead wood habitat types, and this has clearly affected current distribution and frequency of many species. Most importantly, species strictly associated with large decaying logs have decreased in many parts of Europe, while common species associated with coniferous wood have expanded in many regions, due to widespread planting of coniferous trees, where such species are naturally absent or infrequent.

Conservation of fungi faces the challenge of high species diversity, limited knowledge and a general lack of public awareness. In practical conservation the high species diversity makes it necessary to focus on a limited number of indicator species. Indicator species schemes are burdened by shortcomings: some are experience based and flimsy in their definition of indicator goals, whereas others are scientific in their approach, but with disputable results or an irrelevant indicator goal. The IUCN criteria for red-listing organisms are not specifically designed for fungi, and red-listing fungi, that is calculating the risk of their extinction, is complicated by a limited knowledge on population sizes, lifespan and spatial dynamics in fungi. In this chapter both approaches are discussed from an overall perspective, and with respect to two groups of saprotrophic basidiomycetes which are decreasing in Europe—grassland and wood-inhabiting fungi.

The wide variety of dead wood habitats supports a wide variety of specialized fungi, which globally may exceed 100,000 species. Of these the majority of known taxa are Basidiomycota. They exhibit a wide variety of strategies to gain and hold territory within wood, defined by their mode of dispersal, establishment, competitive ability and adaptation to disturbance and stress factors. Many habitat factors affect community composition and development, both exogenous, e.g. microclimatic regime, and endogenous, e.g. interspecific interactions. Initial microenvironmental, factors—at one extreme high stress and at the other extreme absence of abiotic stress—are major determinants of the communities that establish. Following initial establishment, community development is influenced by four main driving forces: stress aggravation (worsening of abiotic environmental conditions), stress alleviation (improvement in abiotic conditions), disturbance and combat (interspecific competition for space rather than directly for nutrients). The ecological strategies adopted by wood decay Basidiomycota, habitat factors influencing community development and community development pathways are discussed in relation to angiosperm wood.

Lactarius lanceolatus O.K. Mill G. Laursen Lactarius lanceolatus O.K. Mill G. Laursen, 1973: 43. Holotype: Alaska, Beaufort Lagoon (VPI). A medium-sized Lactarius with white milk; and orange-brownish colours; very similar to L. aurantiacus but occurring in alpine and arctic habitats with dwarf Salix. DESCRIPTION: Cap 25-55 mm, convex to applanate with a decurved or inrolled margin, later depressed With expanded margin, sometimes with a small umbo or papilla, margin often finely crenulate; surface smooth, dry to sticky or glossy, often becoming mat and areolate-rimose in a concentric pattern, margin at first pruinose, dark brick to brick or orange-brown, darkest in the centre. Gills adnate to slightly decurrent, medium crowded, pinkish buff, later clay-pink to Isabella-coloured. Stem 20-30 x 5-11 mm, cylindric to subclavate; surface smooth, dry, faintly pruinose when young, clay-buff to cinnamon. Flesh fragile, soon hollow in the stem, pale cream to pale pinkish buff, outer part coloured like surface: smell faint or like L. quietus; taste mild or becoming slightly bitter. Milk White or watery white, unchanging; taste mild or slightly astringent. Spore deposit whitish. Spores 72-11.3 x 5.6-8.4 pm, av. 8.0-9.8 x 6.5-7.4 pm, subglobose to ellipsoid, Q = 1.05- 1.55, av. 1.21-138; ornamentation 0.3-0.9 pm high, of fine elongate warts, joined by fine lines and low ridges to form an incomplete reticulum, isolated warts numerous; plage inamyloid or with an amyloid distal spot. Basidia 35-60 x 10-13 pm, subclavate, 4-spored. Pleuromacrocystidia scattered to rather abundant, 50-135 x 6.5-10.5(-12) pm, narrowly fusiform to conical, often flexuose to moniliform, apex acute, subacute or mucronate. Gill edge heterogeneous; cheilomacrocystidia numerous, 20-60 x 4-10 pm, fusiform or conical, apex acute or moniliform. Pileipellis an intricate (ixo)trichoderm; terminal elements 15-50 x 2-5 pm, cylindric; subpellis very compact, of darker, slightly swollen hyphae, about 6 um broad. ECQLOGY AND DISTRIBUTION: In Europe L. lanceolatus is only known from the northernmost part of Fennoscandia. It associates with dwarf Salix and is widely distributed in Greenland, northern North America and probably Siberia. DISCUSSION: Lactarius lanceolatus is similar to, or maybe even conspecific with L. aurantiacus. Lactarius lanceolatus is often somewhat darker and more reddish than L. aurantiacus, and the cracking of the cap is usually not seen in L. aurantiacus. These differences may be caused by the extreme environmental conditions. Lactarius lanceolatus often has longer macrocystidia and larger, more faintly ornamented spores than L. aurantiacus, but intermediates occur. For further discussion, see Gulden et al. (1988)., Published as part of Jacob Heilmann-Clausen, Annemieke Verbeken & Jan Vesterholt, 1998, The Genus Lactarius, Copenhagen :Danish Mycological Society on page 180

Lactarius dryadophilus Kühner Lactarius dryadophilus Kühner, 1975a: 68. Holotype: Norway, Hardanger (LY). SYNONYM: Lactarius dryadophilus var. saliceticola Bon and Jamoni in Jamoni and Bon, 1992b: 21. A medium-sized to large, lilac staining Lactarius with cream-coloured, viscid cap with a bearded margin; growing in arctic and alpine areas. DESCRIPTION: Cap 30-100(-150) mm, at first convex with a slightly depressed centre and decurved or inrolled margin, sometimes slightly umbonate; surface sticky, viscid, at the margin bearded and tomentose with up to 2 mm long hairs, whitish chrome to pale cream or warm buff but more brownish in the centre, azonate or slightly zonate. Gills adnate, medium broad, crowded, often forked, whitish to pale cream, later pinkish buff. Stem 20-30 x 15-20 mm, typically curved and tapering downwards; surface whitish to pale cream, sometimes yellowish at the base, very finely pitted or with ochraceous spots, especially towards the base. Flesh firm, becoming hollow in the stem, white, changing to lilac; smell fruity, sweetish; taste mild, like cedar wood to very slightly acrid. Milk white, changing to lilac in contact with the flesh. Spore deposit cream. Spores 9.3-11.8 x 7.2-9.2 um, av. 10.2-10.3 x 8.1-8.4 um, broadly ellipsoid to ellipsoid, Q = 1.15-1.40, av. 1.22-1.27; ornamentation up to 0.3 um high, of rather fine and narrow ridges, often aligned but not forming a reticulum or at most a very incomplete one; isolated warts often elongate; plage inamyloid. Basidia 60-70 x 12-15 um, subclavate, 4-spored. Pleuromacrocystidia moderately abundant, 80-120 x 10-15 um, fusiform, tapering to a mucronate apex. Gill edge sterile; cheilomacrocystidia 40-60 x 7-9 um, fusiform; paracystidia 10-25 x 4-6(-10) um; cylindric to tortuous or clavate. Pileipellis an ixocutis to an ixotrichoderm, 150-200 um thick; hyphae 1-5 um broad, very thin-walled and shrivelled, gelatinized, repent or ascending, some with conspicuous incrustations. ECOLOGY AND DISTRIBUTION: Found in arctic and alpine vegetation types on rich, calcareous soils, often in Dryas-rich grasslands. Rather rare and known from Fennoscandia, the Alps, the Pyrenees and Greenland, and occuring from August to mid September. DISCUSSION: The cap is tomentose only to a short distance from the margin, unlike in L. repraesentaneus, which also has more yellow colours, a longer stem and a different spore ornamentation. The original description of L. groenlandicus was partly based on this species, but recently lectotypified as a synonym of L. pubescens (Knudsen nda Lamoure, 1993).