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Despite host-fungal symbiotic interactions being ubiquitous in all ecosystems, understanding how symbiosis has shaped the ecology and evolution of fungal spores that are involved in dispersal and colonization of their hosts has been ignored in life-history studies. We assembled a spore morphology database covering over 26,000 species of free-living to symbiotic fungi of plants, insects and humans and found more than eight orders of variation in spore size. Evolutionary transitions in symbiotic status correlated with shifts in spore size, but the strength of this effect varied widely among phyla. Symbiotic status explained more variation than climatic variables in the current distribution of spore sizes of plant-associated fungi at a global scale while the dispersal potential of their spores is more restricted compared to free-living fungi. Our work advances life-history theory by highlighting how the interaction between symbiosis and offspring morphology shapes the reproductive and dispersal strategies among living forms. Alexander von Humboldt-Stiftung, Grant/Award Number: Feodor-Lynen Fellowship; Australian Research Council, Grant/Award Number: DP190103714 and FT0100590; Bundesministerium für Bildung und Forschung, Grant/Award Number: 01LC1501A; Deutsche Forschungsgemeinschaft, Grant/Award Number: HE6183; Deutscher Akademischer Austauschdienst; Division of Environmental Biology, Grant/Award Number: 1623040 and 1655759; Grantová Agentura České Republiky, Grant/Award Number: 21-17749S; H2020 European Research Council, Grant/Award Number: 647038 and 694368; Universities Australia.

This version of the article has been accepted for publication, after peer review and is subject to Springer Nature’s AM terms of use, but is not the Version of Record and does not reflect post-acceptance improvements, or any corrections. The Version of Record is available online at https://doi.org/10.1007/s13225-021-00493-7. Fungi are highly important biotic components of terrestrial ecosystems, but we still have a very limited understanding about their diversity and distribution. This data article releases a global soil fungal dataset of the Global Soil Mycobiome consortium (GSMc) to boost further research in fungal diversity, biogeography and macroecology. The dataset comprises 722,682 fungal operational taxonomic units (OTUs) derived from PacBio sequencing of full-length ITS and 18S-V9 variable regions from 3200 plots in 108 countries on all continents. The plots are supplied with geographical and edaphic metadata. The OTUs are taxonomically and functionally assigned to guilds and other functional groups. The entire dataset has been corrected by excluding chimeras, index-switch artefacts and potential contamination. The dataset is more inclusive in terms of geographical breadth and phylogenetic diversity of fungi than previously published data. The GSMc dataset is available over the PlutoF repository.

A recent paper by Schall et al. (2020) concluded that beech forests managed in even-aged (EA) rotation systems were more efficient than unmanaged (UNM) forest and forest managed in uneven-aged (UEA) selective cutting systems in supporting landscape-scale biodiversity in Germany. The authors based their conclusion on a comprehensive multitaxon survey and a promising resampling model for assessing gamma diversity at landscape scale. Here, we challenge their conclusions and evaluate the importance of UNM forests for conservation of forest biodiversity. The average amount of dead wood reported from EA stands (27.8 m3/ha) was almost 30% higher than reported from UNM stands (21.6 m3/ha) in the study. Averages from long UNM temperate forests in Europe are typically six to seven times higher (131–157 m3/ha). We therefore conclude the UNM studied stands to reflect legacies of former management, and to be poorly representative of UNM forests. Data from our own studies, including long UNM beech stands in Denmark, demonstrate how this shortcoming seriously undermines the general validity of the presented results to conservation of forest biodiversity. Synthesis and applications. Preservation and restoration of intact forest ecosystems remains essential to biodiversity conservation. We show that the findings of Schall et al. (2020) do not contradict this important notion. Schall and colleagues identified UEA management systems as potentially inferior to more traditional EA management systems for conserving forest biodiversity at the landscape scale. The paper also provides insight into the limited short-term conservation value of simply abandoning forest management in intensively managed landscapes. Based on this, we call for discarding the current orthodox view of non-intervention when new forest reserves are created in temperate Europe. Active reinforcement of natural disturbance regimes and active habitat creation may lead to faster recovery of natural stand structure and forest biodiversity.

Aim: In this study, we assessed the importance of local- to landscape-scale effects of land cover and land use on flying insect biomass. Location: Denmark and parts of Germany. Methods: We used rooftop-mounted car nets in a citizen science project (“InsectMobile”) to allow for large-scale geographic sampling of flying insects. Volunteers sampled insects along 278 five-km routes in urban, farmland, grassland, wetland and forest landscapes in the summer of 2018. The bulk insect samples were dried overnight to obtain the sample biomass. We extracted proportional land use variables in buffers between 50 and 1,000 m along the routes and compiled them into land cover categories to examine the effect of each land cover, and specific land use types, on insect biomass. Results: We found a negative association between urban cover and flying insect biomass (1% increase in urban cover = 1% [95% CI: −3.0 to 0.0] decrease in biomass in Denmark, and a 3% [95% CI: −3.0 to 0.0] decrease in Germany) at a landscape scale (1,000-m buffer). In Denmark, we also found positive effects of semi-natural land cover types, that is protected grassland (largest at the landscape scale, 1000 m) and forests (largest at intermediate scales, 250 m). Protected grassland cover had a stronger positive effect on insect biomass than forest cover did. For farmland cover, the positive association with insect biomass was not clearly modified by any variable associated with farmland use intensity. The negative association between insect biomass and urban land cover appeared to be reduced by increased urban green space. Main conclusions: Our results show that land cover has an impact on flying insect biomass with the magnitude of this effect varying across spatial scales. However, the vast expanse of grey space in urbanized areas has a direct negative impact on flying insect biomass across all spatial scales examined.

Identifying the spatial scales at which community assembly processes operate is fundamental for gaining a mechanistic understanding of the drivers shaping ecological communities. In this study, we examined whether and how traits and phylogenetic relationships structure fungal community assembly across spatial scales. We applied joint species distribution modelling to a European-scale dataset on 215 wood-inhabiting fungal species, which includes data on traits, phylogeny and environmental variables measured at the local (log-level) and regional (site-level) scales. At the local scale, wood-inhabiting fungal communities were mostly structured by deadwood decay stage, and the trait and phylogenetic patterns along this environmental gradient suggested the lack of diversifying selection. At regional scales, fungal communities and their trait distributions were influenced by climatic and connectivity-related variables. The fungal climatic niches were not phylogenetically structured, suggesting that diversifying selection or stabilizing selection for climatic niches has played a strong role in wood-inhabiting communities. In contrast, we found a strong phylogenetic signal in the responses to connectivity-related variables, revealing phylogenetic homogenization in small and isolated forests. Synthesis. Altogether, our results show that species-level traits and phylogenies modulate the responses of wood-inhabiting fungi to environmental processes acting at different scales. This result suggests that the evolutionary histories of fungal traits diverge along different environmental axes.

SummaryFungi play pivotal roles in ecosystem functioning, but little is known about their global patterns of diversity, endemicity, vulnerability to global change drivers and conservation priority areas. We applied the high-resolution PacBio sequencing technique to identify fungi based on a long DNA marker that revealed a high proportion of hitherto unknown fungal taxa. We used a Global Soil Mycobiome consortium dataset to test relative performance of various sequencing depth standardization methods (calculation of residuals, exclusion of singletons, traditional and SRS rarefaction, use of Shannon index of diversity) to find optimal protocols for statistical analyses. Altogether, we used six global surveys to infer these patterns for soil-inhabiting fungi and their functional groups. We found that residuals of log-transformed richness (including singletons) against log-transformed sequencing depth yields significantly better model estimates compared with most other standardization methods. With respect to global patterns, fungal functional groups differed in the patterns of diversity, endemicity and vulnerability to main global change predictors. Unlike α-diversity, endemicity and global-change vulnerability of fungi and most functional groups were greatest in the tropics. Fungi are vulnerable mostly to drought, heat, and land cover change. Fungal conservation areas of highest priority include wetlands and moist tropical ecosystems.

Fungal and arthropod consumers constitute the vast majority of global terrestrial biodiversity. Yet, the link from richness and composition of producer (plant) communities to the richness of consumer communities is poorly understood. Fungal and arthropod species richness could be a simple function of producer species richness at a site. Alternatively, it could be a complex function of chemical and structural properties of the producer species making up communities. We used databases on plant–fungus and plant–arthropod trophic links to derive the richness of consumer biota per associated plant species (coined link score). We assessed how well link scores could be predicted by simple attributes of plant species. Next, we used a multi-taxon inventory of 130 sites, representing all major habitat types in a country (Denmark), to investigate whether link scores summed over plant species in communities (coined link sum) could outperform simple plant species richness as predictor of fungal and arthropod richness at the sites. We found plant species' link scores for both fungi and arthropods to be positively related to plant size, regional occupancy, nativeness and ectomycorrhizal status. Link-based indices generally improved the prediction of richness of fungal and arthropod communities. For fungal communities, both observed link sum (from databases) and predicted link sum (from plant attributes) had high predictive power, while plant richness alone had none. For arthropod communities, predictive performance varied between functional groups. For both fungi and arthropods, richness predictions were further improved by considering abiotic habitat conditions. Our results underline the importance of plants as niche space for the megadiverse groups of arthropods and fungi. The plant–attribute approach holds promise for predicting local and regional consumer richness in areas of the world lacking detailed plant–consumer databases.

Caio Graco-Roza was funded by the Coordination for the Improvement of Higher Education Personnel (CAPES), the Carlos Chagas Filho Research Support Foundation (FAPERJ) and the Ella and Georg Erhnrooth Foundation; Jan Altman by research grants INTER-EXCELLENCE LTAUSA19137 provided by Czech Ministry of Education, Youth and Sports, 20-05840Y of the Czech Science Foundation, and long-term research development project no. RVO 67985939 of the Czech Academy of Sciences; Otso Ovaskainen was funded by Academy of Finland (grant no. 309581), Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme (223257), and the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement no. 856506; ERC-synergy project LIFEPLAN); and Jianjun Wang was funded by CAS Key Research Program of Frontier Sciences (QYZDB-SSW-DQC043) and National Natural Science Foundation of China (91851117). The "sPlot" project was initiated by sDiv, the Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (DFG FZT 118), and is now a platform of iDiv. The study was supported by the TRY initiative on plant traits (). We are also grateful to Jens Kattge and TRY database. TRY is hosted, developed and maintained at the Max Planck Institute for Biogeochemistry (MPI-BGC) in Jena, Germany, in collaboration with the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. The CESTES database of metacommunities is also an initiative of iDiv led by Alienor Jeliazkov. We thank sDiv for supporting the open science initiative., Aim: Understanding the variation in community composition and species abundances (i.e., β-diversity) is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments. Location: Global. Time period: 1990 to present. Major taxa studied: From diatoms to mammals. Method: We measured the strength of the decay using ranked Mantel tests (Mantel r) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features. Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances. Main conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments., Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) Fundacao Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio De Janeiro (FAPERJ), Ella and Georg Erhnrooth Foundation, Ministry of Education, Youth & Sports - Czech Republic LTAUSA19137 Grant Agency of the Czech Republic 20-05840Y Czech Academy of Sciences RVO 67985939, Academy of Finland 309581, Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme 223257, European Research Council (ERC) 856506, CAS Key Research Program of Frontier Sciences QYZDB-SSW-DQC043, National Natural Science Foundation of China (NSFC) 91851117, German Research Foundation (DFG) DFG FZT 118, TRY initiative on plant traits

15 páginas.- 6 figuras.- 93 referencias, Fungi are highly diverse organisms, which provide multiple ecosystem services. However, compared with charismatic animals and plants, the distribution patterns and conservation needs of fungi have been little explored. Here, we examined endemicity patterns, global change vulnerability and conservation priority areas for functional groups of soil fungi based on six global surveys using a high-resolution, long-read metabarcoding approach. We found that the endemicity of all fungi and most functional groups peaks in tropical habitats, including Amazonia, Yucatan, West-Central Africa, Sri Lanka, and New Caledonia, with a negligible island effect compared with plants and animals. We also found that fungi are predominantly vulnerable to drought, heat and land-cover change, particularly in dry tropical regions with high human population density. Fungal conservation areas of highest priority include herbaceous wetlands, tropical forests, and woodlands. We stress that more attention should be focused on the conservation of fungi, especially root symbiotic arbuscular mycorrhizal and ectomycorrhizal fungi in tropical regions as well as unicellular early-diverging groups and macrofungi in general. Given the low overlap between the endemicity of fungi and macroorganisms, but high conservation needs in both groups, detailed analyses on distribution and conservation requirements are warranted for other microorganisms and soil organisms., The bulk of the funding is derived from the Estonian Science Foundation (grants PRG632, PRG1170, PRG1615, MOBTP198), EEA Financial Mechanism Baltic Research Programme (EMP442), and Novo Nordisk Fonden (NNF20OC0059948).

The geographical distributions of species associated with European temperate broadleaf forests have been significantly influenced by glacial–interglacial cycles. During glacial periods, these species persisted in Mediterranean and extra-Mediterranean refugia and later, during interglacial periods, expanded northwards. The widespread saproxylic beetle Bolitophagus reticulatus depends closely on European temperate broadleaf forests. It usually develops in the tinder fungus Fomes fomentarius, a major decomposer of broadleaf-wood. We sampled B. reticulatus in sporocarps from European beech (Fagus sylvatica) and Oriental beech (Fagus orientalis) across Europe and the Caucasus region. We analysed mitochondrial gene sequences (cox1, cox2, cob) and 17 microsatellites to reconstruct the geographical distribution of glacial refugia and postglacial recolonization pathways. We found only marginal genetic differentiation of B. reticulatus, except for a significant split between populations of the Caucasus region and Europe. This indicates the existence of past refugia south of the Great Caucasus, and a contact zone with European populations in the Crimean region. Further potential refugia might have been located at the foothills of the Pyrenees and in the Balkan region. Our genetic data suggest a phalanx-wise recolonization of Europe, a reflection of the high mobility of B. reticulatus.

Understanding the variation in community composition and species abundances, i.e., β-diversity, is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional turnover in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distances. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 149 datasets comprising different types of organisms and environments. We modelled an exponential distance decay for each dataset using generalized linear models and extracted r2 and slope to analyse the strength and the rate of the decay. We studied whether taxonomic or functional similarity has stronger decay across the spatial and environmental distances. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm, and organismal features. Taxonomic distance decay was stronger along spatial and environmental distances compared with functional distance decay. The rate of taxonomic spatial distance decay was the fastest in the datasets from mid-latitudes while the rate of functional decay increased with latitude. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distances but a higher rate of decay along environmental distances. Marine ecosystems had the slowest rate of decay. This synthesis is an important step towards a more holistic understanding of patterns and drivers of taxonomic and functional β-diversity.

Monitoring insects across space and time is challenging, due to their vast taxonomic and functional diversity. This study demonstrates how nets mounted on rooftops of cars (car nets) and DNA metabarcoding can be applied to sample flying insect richness and diversity across large spatial scales within a limited time period. During June 2018, 365 car net samples were collected by 151 volunteers during two daily time intervals on 218 routes in Denmark. Insect bulk samples were processed with a DNA metabarcoding protocol to estimate taxonomic composition, and the results were compared to known flying insect richness and occurrence data. Insect and hoverfly richness and diversity were assessed across biogeographic regions and dominant land cover types. We detected 15 out of 19 flying insect orders present in Denmark, with high proportions of especially Diptera compared to Danish estimates, and lower insect richness and diversity in urbanized areas. We detected 319 species not known for Denmark and 174 species assessed in the Danish Red List. Our results indicate that the methodology can assess the flying insect fauna at large spatial scales to a wide extent, but may be, like other methods, biased towards certain insect orders.

Consumers constitute the vast majority of global terrestrial biodiversity. Yet, local consumer richness is poorly understood. Plant species richness offers a simple hypothesis to how the diversification of carbon substrates may promote the diversity of arthropods and fungi. We took this one step further and used databases on plant-consumer interaction links to derive the richness of associated biota per plant species (link score). Using a species inventory of 130 sites we investigated 1) how well the link score could be predicted by plant attributes and 2) if the sum of plant species’ observed or predicted link scores could predict site richness of arthropods and macrofungi better than plant species richness alone. We found plant link scores to be positively related to plant size, abundance, nativeness and ectomycorrhizal status. Link based indices generally improved prediction of richness, stressing the importance of plants as niche space for the megadiverse groups of insects and fungi.

The Danish Fungal Atlas is a citizen science project launched in 2009 in collaboration among the University of Copenhagen, Mycokey and the Danish Mycological Society. The associated database now holds almost 1 million fungal records, contributed by more than 3000 recorders. The records represent more than 8000 fungal species, of which several hundred have been recorded as new to Denmark during the project. In addition several species have been described as new to science. Data are syncronized with the Global Biodiversity Information Facility (GBIF) on a weekly basis, and is hence freely available for research and nature conservation. Data have been used for systematic conservation planning in Denmark, and several research papers have used data to explore subjects such as host selection in wood-inhabiting fungi (Heilmann‐Clausen et al. 2016), recording bias in citizen science (Geldmann et al. 2016), fungal traits (Krah et al. 2019), biodiversity patterns (e.g. Andrew et al. 2018), and species discovery (Heilmann-Clausen et al. 2019). The project database is designed to faciliate direct interactions and communication among volunteers. The validation of submitted records is interactive and combines species-specific smart filters, user credibility, and expert tools to secure the highest possible data credibility. In 2019, an AI (artificial intelligence) trained species identification tool was launched along with a new mobile app, enabling users to identify and record species directly in the field (Sulc et al. 2020). At the same time, DNA sequencing was tested as an option to test difficult identifications, and in 2021 a high-throughput sequencing facility was developed to allow DNA sequencing of hundreds of fungal collections at a low cost. The presentation will give details on data validation, data use and how we have worked with cultivation of volunteers to provide a truly coherent model for collaboration on mushroom citizen science.

Insect diversity and abundance are increasingly reported as being under pressure. However, monitoring insects across space and time is challenging, due to their vast taxonomic and functional diversity. This study demonstrates how nets mounted on rooftops of cars (car nets) and DNA metabarcoding can be applied to sample flying insect diversity across a large spatial scale within a limited time period.During June 2018, 365 car net samples were collected by 151 volunteers during two daily time intervals on 218 routes in Denmark. Insect bulk samples were processed with a DNA metabarcoding protocol to estimate taxonomic composition, and the results were compared to known flying insect diversity and occurrence data.We detected 15 out of 19 flying insect orders present in Denmark. Diptera. Psocoptera and Thysanoptera were overrepresented, while Hymenoptera, Coleoptera, Lepidoptera, Trichoptera, Odonata, Neuroptera and Plecoptera were underrepresented, compared to Danish estimates. We detected 319 species not known for Denmark. Our results indicate that this method can assess the flying insect fauna to a wide extent, but may be, like other methods, biased towards certain insect orders. Furthermore, car net sampling and DNA metabarcoding can update species records for Denmark while preserving specimens for natural history collections.

Recent studies report declines in biomass, abundance and diversity of terrestrial insect groups. While anthropogenic land use is one likely contributor to this decline, studies assessing land cover as a driver of insect dynamics are rare and mostly restricted in spatial scale and types of land cover. In this study, we used rooftop-mounted car nets in a citizen science project (‘InsectMobile’) to allow for large-scale geographic sampling of flying insects across Denmark and parts of Germany. Citizen scientists sampled insects along 278 10 km routes in urban, farmland and semi-natural (grassland, wetland and forest) landscapes in the summer of 2018. We assessed the importance of local to landscape-scale effects and land use intensity by relating insect biomass to land cover in buffers of 50, 250, 500 and 1000 m along the routes. We found a negative association of urban cover and a positive association of farmland on insect biomass at a landscape-scale (1000 m buffer) in both countries. In Denmark, we also found positive effects of all semi-natural land covers, i.e. grassland (largest at the landscape-scale, 1000 m), forests (largest at intermediate scales, 250 m), and wetlands (largest at the local-scale, 50 m). The negative association of insect biomass with urban land cover and positive association with farmland were not clearly modified by any variable associated with land use intensity. Our results show that land cover has an impact on flying insect biomass with the magnitude of this effect varying across spatial scales. Since we consistently found negative effects of urban land cover, our findings highlight the need for the conservation of semi-natural areas, such as wetlands, grasslands and forests, in Europe.

Aim We investigated the consistency between richness and trait-based diversity metrics in capturing the effects of management-related habitat factors on biodiversity. The choice of biodiversity metrics can substantially affect the evaluation of conservation tools. However, the relative sensitivity of different metrics is not well investigated, especially in a multi-taxon framework. Location European beech forests in Denmark. Methods We studied 20 beech stands comprising four management types (from intensively managed to long unmanaged stands). We analyzed how management-related environmental variables were reflected in the measure of: (i) species richness, (ii) number of conservation-relevant species (red-listed species and old-growth forest indicators) and (iii) functional diversity targeting five organism groups with different habitat requirements, i.e. vascular plants, epiphytic lichens and bryophytes, saproxylic fungi and breeding birds. Results Plain species richness at stand level was generally misleading, as it did not capture changes in the number of conservation relevant species with changes in management-related environmental variables. The interpretation of functional responses was most informative for the better known vascular plants, while responses were more fragmented for the other organism groups. Overall, however, functional responses were consistent with a loss of specialization and progressive simplification of species assemblages from long-unmanaged to intensively managed stands. Conclusions Our findings suggest that the occurrence of conservation-relevant species is a sound and relevant metric for planning and evaluating conservation actions, especially for less studied organism groups (e.g., saproxylic fungi and epiphytes). The functional approach is promising, but presupposes the availability of databases of relevant traits.

Aim: Macroecological scales of species compositional trends are well documented for a variety of plant and animal groups, but remain sparse for fungi, despite their ecological importance in carbon and nutrient cycling. It is, thus, essential to understand the composition of fungal assemblages across broad geographical scales and the underlying drivers. Our overall aim was to describe these patterns for fungi across two nutritional modes (saprotrophic and ectomycorrhizal). Furthermore, we aimed to elucidate the temporal component of fruiting patterns and to relate these to soil carbon and nitrogen deposition. Location: Central and Northern Europe. Methods: A total of 4.9 million fungal fruit body observations throughout Europe, collected between 1970 and 2010, were analysed to determine the two main environmental and geographical gradients structuring fungal assemblages for two main nutritional modes, saprotrophic and ectomycorrhizal fungi. Results: Two main gradients explaining the geography of compositional patterns were identified, for each nutritional mode. Mean annual temperature (and related collinear, seasonal measures) correlated most strongly with the first gradient for both nutritional modes. Soil organic carbon was the highest correlate of the second compositional gradient for ectomycorrhizal fungi, suspected as an indicator of vegetation- and pH-related covariates. In contrast, nitrogen deposition constituted a second gradient for saprotrophic fungi, likely a proxy for anthropogenic pollution. Compositional gradients and environmental conditions correlated similarly when the data were divided into two time intervals of 1970–1990 and 1991–2010. Evidence of compositional temporal change was highest with increasing elevation and latitude. Main conclusions: Fungal assemblage patterns demonstrate clear biogeographical patterns that relate the nutritional modes to their main environmental correlates of temperature, soil organic carbon and nitrogen deposition. With respect to global change impacts, the highest rates of compositional change by time suggest targeting higher latitudes and elevations for a better understanding of fungal dynamics. We, finally, suggest further examination of the ranges and dispersal abilities of fungi to better assess responses to global change.

Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to amend the International Code of Nomenclature for algae, fungi, and plants (ICN), allowing DNA sequences alone (of any region and extent) to serve as types of taxon names for voucherless fungi (mainly putative taxa from environmental DNA sequences), have been submitted to be voted on at the 11th International Mycological Congress (Puerto Rico, July 2018). We consider various genetic processes affecting the distribution of alleles among taxa and find that alleles may not consistently and uniquely represent the species within which they are contained. Should the proposals be accepted, the meaning of nomenclatural types would change in a fundamental way from physical objects as sources of data to the data themselves. Such changes are conducive to irreproducible science, the potential typification on artefactual data, and massive creation of names with low information content, ultimately causing nomenclatural instability and unnecessary work for future researchers that would stall future explorations of fungal diversity. We conclude that the acceptance of DNA sequences alone as types of names of taxa, under the terms used in the current proposals, is unnecessary and would not solve the problem of naming putative taxa known only from DNA sequences in a scientifically defensible way. As an alternative, we highlight the use of formulas for naming putative taxa (candidate taxa) that do not require any modification of the ICN. Publisher’s Note A first version of this text was prepared by the first eight authors and the last one, given here. The other listed co-authors in the article PDF support the content, and their actual contributions varied from only support to additions that substantially improved the content. The full details of all co-authors, with their affiliations, are included in Supplementary Table 1 after p.175 of the article for reasons of clarity and space. Slavomír Adamčík Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Teuvo Ahti Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland M. Catherine Aime Purdue University, 915 W. State St., West Lafayette, Indiana 47907, U.S.A. A. Martyn Ainsworth Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom László Albert Hungarian Mycological Society, 1087 Könyves Kálmán krt. 40, Budapest, Hungary Edgardo Albertó Instituto de Investigaciones Biotecnológicas-Instituto Tecnológico de Chascomús, Universidad Nacional de San Martin-Consejo Nacional de Investigaciones Científicas y Técnicas, Buenos Aires, Argentina Alberto Altés García Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Dmitry Ageev SIGNATEC Ltd., 630090, Novosibirsk, Akademgorodok (Novosibirsk Scientific Center), Inzhenernaya str., 22, Russia Reinhard Agerer Ludwig-Maximilians-Universität München, Menzinger Str. 67, 80638 München, Germany Begona Aguirre-Hudson Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom Joe Ammirati University of Washington, Seattle, Washington 98195-1800, U.S.A. Harry Andersson Eichhahnweg 29a, 38108 Braunschweig, Germany Claudio Angelini Jardín Botánico Nacional Dr. Rafael Ma. Moscoso, Apartado 21-9, Santo Domingo, Dominican Republic Vladimír Antonín Moravian Museum, Zeny trh 6, 659 37 Brno, Czech Republic Takayuki Aoki Genetic Resources Center, National Agriculture and Food Research Organization, 2-1-2 Kannondai, Tsukuba, Ibaraki 305-8602, Japan André Aptroot ABL Herbarium, G.v.d.Veenstraat 107, 3762 XK Soest, The Netherlands Didier Argaud 40 rue du Justemont, 57290 Fameck, France Blanca Imelda Arguello Sosa Instituto Tecnológico de Ciudad Victoria, Tecnológico Nacional de México, Ciudad Victoria, Tamaulipas, Mexico Arne Aronsen Torødveien 54, 3135 Torød, Norway Ulf Arup Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Bita Asgari Iranian Research Institute of Plant Protection, Agricultural Research, Education and Extension Organization, Tehran, Iran Boris Assyov Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Str., 1113 Sofia, Bulgaria Violeta Atienza Facultad de Ciencias Biológicas, Universitat de València, C/Dr Moliner 50, 46100, Burjasot, Valencia, Spain Ditte Bandini Panoramastr 47, 69257 Wiesenbach, Germany João Luís Baptista-Ferreira Instituto de Biossistemas e Ciências Integrativas, Faculdade de Ciências da Universidade de Lisboa, 1749-016 Lisboa, Portugal Hans-Otto Baral Blaihofstr. 42, 72074 Tübingen, Germany Tim Baroni The State University of New York, 340 Bowers Hall, P.O. Box 2000, Cortland, New York 13045, U.S.A. Robert Weingart Barreto Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Henry Beker (1) Royal Holloway, University of London, United Kingdom; (2) Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Ann Bell 45 Gurney Road, Lower Hutt, New Zealand Jean-Michel Bellanger CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Francesco Bellù Naturmusem of Bolzano, CP 104, 39100, Bolzano, Italy Martin Bemmann Kleingemünderstraße 111, 69118 Heidelberg, Germany Mika Bendiksby NTNU, University Museum, Norwegian University of Science and Technology, 7491 Trondheim, Norway Egil Bendiksen Norwegian Institute for Nature Research, Gaustadalleen 21, 0349 Oslo, Norway Katriina Bendiksen Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Lajos Benedek Szent Istvan University, Hungary Anna Bérešová-Guttová Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Franz Berger University of Salzburg, Salzburg, Austria Reinhard Berndt Herbaria Z+ZT, ETH Zürich, CHN D37, Universitätstr. 16, 8092 Zürich, Switzerland Annarosa Bernicchia Via A. Guidotti 39, 40134 Bologna, Italy Alona Yu. Biketova Institute of Biochemistry, BRC-HAS, 6726 Szeged, Temesvari krt. 62, 6726 Szeged, Hungary Enrico Bizio Società Veneziana di Micologia, Società Veneziana di Scienze Naturali, Fontego dei Turchi, Santa Croce 1730, 30135 Venice, Italy Curtis Bjork UBC Herbarium, Beaty Biodiversity Museum, University of British Columbia, Canada Teun Boekhout (1) Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; (2) Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Amsterdam, The Netherlands David Boertmann Aarhus University, Frederiksborgvej 399, 4000 Roskilde, Denmark Tanja Böhning AG Geobotanik Schleswig-Holstein & Hamburg, c/o University of Kiel, Olshausenstraße 75, 24098 Kiel, Germany Florent Boittin Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Carlos G. Boluda Conservatoire et Jardin botaniques de la Ville de Genève, 1292 Genève, Switzerland Menno W. Boomsluiter T.v.Lohuizenstraat 34, 8172xl, Vaassen, The Netherlands Jan Borovička Institute of Geology, Czech Academy of Sciences, Rozvojova 269, 165 00 Prague 6, Czech Republic Tor Erik Brandrud Norwegian Institute for Nature Research, Gaustadalleen 21, 0349 Oslo, Norway Uwe Braun Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik, und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle, Germany Irwin Brodo Canadian Museum of Nature, 240 McLeod Street, Ottawa, Ontario, Canada Tatiana Bulyonkova A.P. Ershov Institute of Informatics Systems, Russian Academy of Sciences, Siberian Branch, 6 Acad. Lavrentjev pr., Novosibirsk 630090, Russia Harold H. Burdsall Jr. Fungal & Decay Diagnostics, LLC, 9350 Union Valley Road, Black Earth, Wisconsin 53515, U.S.A. Bart Buyck Muséum National d’Histoire Naturelle, CP 39, ISYEB, UMR 7205 CNRS MNHN UPMC EPHE, 12 rue Buffon, 75005 Paris, France Ana Rosa Burgaz Facultad de Ciencias Biológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain Vicent Calatayud Fundación CEAM, c/ Charles R. Darwin, 14, Parque Tecnológico, 46980 Paterna, Valencia, Spain Philippe Callac INRA, MycSA, CS 20032, 33882 Villenave d’Ornon, France Emanuele Campo Associazione Micologica Bresadola, Via Alessandro Volta 46, 38123 Trento, Italy Massimo Candusso Via Ottone Primo 90, 17021, Alassio, Savona, Italy Brigitte Capoen Queffioec, rue de Saint Gonval, 22710 Penvenan, France Joaquim Carbó Roser, 60, 17257 Torroella de Montgrí, Girona, Spain Matteo Carbone Via Don Luigi Sturzo 173 16148 Genova, Italy Rafael F. Castañeda-Ruiz Instituto de Investigaciones Fundamentales en Agricultura, Tropical ‘Alejandro de Humboldt’, OSDE, Grupo Agrícola, Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana 17200, Cuba Michael A. Castellano USDA, Forest Service, Northern Research Station, Corvallis, Oregon 97330, U.S.A. Jie Chen Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Philippe Clerc Conservatoire et Jardin botaniques de la Ville de Genève, 1292 Genève, Switzerland Giovanni Consiglio Via C. Ronzani 61, 40033 Casalecchio Bologna, Italy Gilles Corriol National Botanical Conservatory for Pyrenees and Midi-Pyrénées Region of France and BBF Herbarium, Vallon de Salut. B.P. 315. 65203 Bagnères-de-Bigorre, France Régis Courtecuisse Université Lille, Fac. Pharma. Lille, EA4483 IMPECS, 59000 Lille, France Ana Crespo Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Cathy Cripps Plant Sciences & Plant Pathology, 119 Plant Biosciences Building, Montana State University, Bozeman, Montana 59717, U.S.A. Pedro W. Crous Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Gladstone Alves da Silva Universidade Federal de Pernambuco, Centro de Biociências, Avenida da Engenharia, S/N, Cidade Universitária, Recife, Pernambuco, Brazil Meiriele da Silva Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Marjo Dam Hooischelf 13, 6581 SL Malden, The Netherlands Nico Dam Hooischelf 13, 6581 SL Malden, The Netherlands Frank Dämmrich The Bavarian Natural History Collections (SNSB Munich), Menzinger Strasse 71, 80638, München, Germany Kanad Das Botanical Survey of India, Cryptogamic Unit, P.O. Botanic Garden, Howrah 711103, W.B., India Linda Davies Centre for Environmental Policy, Imperial College London, SW7 2AZ, United Kingdom Eske De Crop Ghent University K.L. Ledeganckstraat 35, 9000 Ghent, Belgium Andre De Kesel Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Ruben De Lange Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Bárbara De Madrignac Bonzi Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Thomas Edison E. dela Cruz University of Santo Tomas, Espana 1008 Manila, Philippines Lynn Delgat Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Vincent Demoulin Institut de Botanique, B.22, Université de Liège, 4000 Liège I, Belgium Dennis E. Desjardin HD Thiers Herbarium (SFSU), San Francisco State University, 1600 Holloway Ave, San Francisco, California 94132, U.S.A. Paul Diederich Musée national d’histoire naturelle, 25 rue Münster, 2160 Luxembourg, Luxembourg Bálint Dima (1) Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; (2) Viikki Plant Science Centre, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland Maria Martha Dios Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Catamarca, Av Belgrano 300, 4700 San Fernando del Valle de Catamarca, Argentina Pradeep Kumar Divakar Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Clovis Douanla-Meli Julius Kühn-Institut, Federal Research Centre for Cultivated Plants, Institute for National and International Plant Health, Messeweg 11-12, 38104 Braunschweig, Germany Brian Douglas Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom Elisandro Ricardo Drechsler-Santos Universidade Federal de Santa Catarina, Campus Universitário Reitor João David Ferreira Lima, Trindade, Florianópolis, Santa Catarina CEP 88040-900, Brazil Paul S. Dyer School of Life Sciences, University of Nottingham, University Park, Nottingham NG7 2RD, United Kingdom Ursula Eberhardt Abt. Botanik, Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany Damien Ertz Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Fernando Esteve-Raventós Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Javier Angel Etayo Salazar Navarro Villoslada 16, 3º dcha., 31003 Pamplona, Navarra, Spain Vera Evenson Sam Mitchel Herbarium of Fungi, Denver Botanic Gardens, 1007 York Street, Denver, Colorado 80206, U.S.A. Guillaume Eyssartier Muséum national d’histoire naturelle, Jardin des plantes, 57 rue Cuvier, 75005 Paris, France Edit Farkas Institute of Ecology and Botany, MTA Centre for Ecological Research, 2163 Vácrátót, Hungary Alain Favre Fédération Mycologique et Botanique Dauphiné Savoie, Le Prieuré, 144 Place de l’Eglise, 74320 Sevrier, France Anna G. Fedosova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Mario Filippa Regione Monsarinero 36, 14041 Agliano Terme, Italy Péter Finy 8000 Székesfehérvár, Zsombolyai u. 56, Hungary Adam Flakus W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Simón Fos Facultad de Ciencias Biológicas, Universitat de València, C/Dr Moliner 50, 46100, Burjasot, Valencia, Spain Jacques Fournier Las Muros, F. 09420 Rimont, France André Fraiture Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Paolo Franchi Associazione Micologica Bresadola, Via Alessandro Volta 46, 38123 Trento, Italy Ana Esperanza Franco Molano Escuela de Microbiología, Universidad de Antioquia, AA1226, Fundación Biodiversa Colombia, Medellín, Colombia Gernot Friebes Centre of Natural History, Botany & Mycology, Universalmuseum Joanneum, Weinzöttlstraße 16, 8045 Graz, Austria Andreas Frisch NTNU, University Museum, Norwegian University of Science and Technology, 7491 Trondheim, Norway Alan Fryday Michigan State University, East Lansing, Michigan 48824, U.S.A. Giuliana Furci The Fungi Foundation, Paseo Bulnes 79 of. 112A, Santiago, Chile Ricardo Galán Márquez Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Matteo Garbelotto University of California, 130 Mulford Hall #3114 Berkeley, California 94720, U.S.A. Joaquina Maria Garcia-Martin Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Mónica A. García Otálora Herbaria Z+ZT, ETH Zürich, CHN D37, Universitätstr. 16, 8092 Zürich, Switzerland Dania García Sánchez Universitat Rovira i Virgili, C/ Sant Llorenç 21, 43201 Reus, Tarragona, Spain Alain Gardiennet 14 rue Roulette, 21260 Véronnes, France Sigisfredo Garnica Instituto de Bioquímica y Microbiología, Universidad Austral de Chile, Isla Teja Campus, Casilla 567, Valdivia, Chile Isaac Garrido Benavent Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Genevieve Gates Tasmanian Institute of Agriculture, Private Bag 54, Hobart, Tasmania 7001, Australia Alice da Cruz Lima Gerlach Conservatoire et Jardin Botaniques de la ville de Genève, Genève, Switzerland Masoomeh Ghobad-Nejhad Iranian Research Organization for Science and Technology, P.O. Box 15815-3538, Tehran 15819, Iran Tatiana B. Gibertoni Universidade Federal de Pernambuco, Centro de Biociências, Avenida da Engenharia, S/N, Cidade Universitária, Recife, Pernambuco, Brazil Tine Grebenc Slovenian Forestry Institute, Vecna pot 2, 100 Ljubljana, Slovenia Irmgard Greilhuber University of Vienna, Rennweg 14, 1030 Vienna, Austria Bella Grishkan Institute of Evolution, University of Haifa, Aba Khoushi Ave. 199, Mt. Carmel, Haifa 3498838, Israel Johannes Z. Groenewald Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Martin Grube Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Gérald Gruhn Office National des Forêts, 2 Avenue de Saint-Mandé, 75570 Paris Cedex 12, France Cécile Gueidan CSIRO — Australian National Herbarium, Clunies Ross Street, Canberra ACT 2601, Australia Gro Gulden Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Luis FP Gusmão Universidade Estadual de Feira de Santana, Av. Transnordestina, s/n, Bairro Novo Horizonte, CEP:44036-900, Feira de Santana, Bahia, Brazil Josef Hafellner Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Michel Hairaud 2 Impasse des Marronniers, 79360 Poivendre de Marigny, France Marek Halama Museum of Natural History, Wrocław University, ul. H. Sienkiewicza 5, 50-335 Wrocław, Poland Nils Hallenberg University of Gothenburg, Box 461, 40530 Göteborg, Sweden Roy E. Halling Institute of Systematic Botany, New York Botanical Garden, 2900 Southern Blvd, Bronx, New York 10458-5126, U.S.A. Karen Hansen Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Christoffer Bugge Harder Texas Tech University, Box 42122, Lubbock, Texas 79409, U.S.A. Jacob Heilmann-Clausen Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Stip Helleman Sweelinck 78, 5831KT Boxmeer, The Netherlands Alain Henriot Mycological Society of France, 20 rue Rottembourg, 12th arrondissement, Paris, France Margarita Hernandez-Restrepo Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Raphaël Herve 24 rue des Fougères, 86550 Mignaloux-Beauvoir, France Caroline Hobart 84 Stafford Road, Sheffield, South Yorkshire S2 2SF, United Kingdom Mascha Hoffmeister Julius Kühn-Institut, Institute for Epidemiology and Pathogen Diagnostics, Braunschweig, Germany Klaus Høiland University of Oslo, P.O.Box 1066, Blindern, 0316 Oslo, Norway Jan Holec National Museum, Herbarium PRM, Cirkusová 1740, 193 00 Praha 9, Czech Republic Håkon Holien Faculty of Bioscience and Aquaculture, NORD University, P.O. Box 2501, 7729 Steinkjer, Norway Karen Hughes University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Vit Hubka Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Seppo Huhtinen Herbarium TUR, Biodiversity Unit, University of Turku, 20014 Turku, Finland Boris Ivančević Natural History Museum, Njegoševa 51, P.O. Box 401, 11000 Belgrade, Serbia Marian Jagers Reelaan 13, 7522 LR Enschede, The Netherlands Walter Jaklitsch Institute of Forest Entomology, Forest Pathology and Forest Protection, University of Natural Resources and Life Sciences Vienna, Vienna, Austria AnnaElise Jansen Stationsstraat 10, 6701 AM Wageningen, the Netherlands Ruvishika S. Jayawardena Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Thomas Stjernegaard Jeppesen Global Biodiversity Information Facility, Universitetsparken 15, 2100 København Ø, Denmark Mikael Jeppson Lilla Håjumsgatan 4, 46135 Trollhättan, Sweden Peter Johnston Manaaki Whenua Landcare Research, Private Bag 92170, Auckland 1072, New Zealand Per Magnus Jørgensen University of Bergen, Allégaten 41, P.O. Box 7800, 5020 Bergen, Norway Ingvar Kärnefelt Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Liudmila B. Kalinina Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Gintaras Kantvilas Tasmanian Herbarium (HO), Tasmanian Museum and Art Gallery, P.O. Box 5058, UTAS LP.O., Sandy Bay, Tasmania 7005, Australia Mitko Karadelev Institute of Biology, Faculty of Natural Science and Mathematics, Ss. Cyril and Methodius University, Arhimedova 5, 1000 Skopje, Republic of Macedonia Taiga Kasuya Faculty of Risk and Crisis Management, Chiba Institute of Science, 3 Shiomi-cho, Choshi, Chiba 288-0025, Japan Ivona Kautmanová Natural History Museum, Slovak National Museum, Bratislava, Slovakia Richard W. Kerrigan RWK Research, Kittanning, Pennsylvania 16201, U.S.A. Martin Kirchmair Institut für Mikrobiologie, Universität Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria Anna Kiyashko Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Dániel G. Knapp Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary Henning Knudsen Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Kerry Knudsen Faculty of Environmental Sciences, University of Life Sciences at Prague, Prague, Czech Republic Tommy Knutsson Nedra Västerstad 111, 380 62 Mörbylånga, Sweden Miroslav Kolařík Institute of Microbiology ASCR, Videnska 1083, 142 20 Prague 4, Czech Republic Urmas Kõljalg Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Alica Košuthová Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Attila Koszka Faculty of Agricultural and Environmental Sciences, Kaposvar University, 7400 Kaposvar, Hungary Heikki Kotiranta Finnish Environment Institute, P.O. Box 140, 00251 Helsinki, Finland Vera Kotkova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Ondřej Koukol Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Jiří Kout University of West Bohemia, Faculty of Education, Klatovska 51, 306 19 Pilsen, Czech Republic Gábor M. Kovács Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary Martin Kříž Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Åsa Kruys Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Viktor Kučera Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Linas Kudzma 37 Maple Ave. Annandale, New Jersey 08801, U.S.A. Francisco Kuhar Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina Martin Kukwa Faculty of Biology, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland T. K. Arun Kumar The Zamorin’s Guruvayurappan College, Kozhikode, Kerala 673014, India Vladimír Kunca Technical University in Zvolen, Ul. T. G. Masaryka 24, 960 53 Zvolen,Slovakia Ivana Kušan Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Thomas W. Kuyper Wageningen University & Research, Droevendaalsesteeg 4, 6708 PB Wageningen, The Netherlands Carlos Lado Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Thomas Læssøe Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Patrice Lainé 123 rue Saint Antoine, 75004, Paris, France Ewald Langer University of Kassel, Heinrich-Plett-Strasse 40, 34132 Kassel, Germany Ellen Larsson University of Gothenburg, Box 461, 40530 Göteborg, Sweden Karl-Henrik Larsson Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Gary Laursen Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, Alaska 99775-7000, U.S.A. Christian Lechat Ascofrance, 64 route de Chizé, 79360 Villiers en Bois, France Serena Lee Herbarium Singapore Botanic Gardens, National Parks Board, Singapore James C. Lendemer (1) Institute of Systematic Botany, New York Botanical Garden, 2900 Southern Blvd, Bronx, New York 10458-5126, U.S.A.; (2) Graduate Center, City University of New York, 365 5th Ave, New York, New York 10016, U.S.A. Laura Levin University of Buenos Aires, Junin 956, 1113, Buenos Aires, Argentina Uwe Lindemann Landesmuseum für Naturkunde, Münster, Germany Håkan Lindström Östansjö 150, 840 64 Kälarne, Sweden Xingzhong Liu Institute of Microbiology, Chinese Academy of Sciences, No 3 Park 1, Beichen West Road, Chaoyang District, Beijing 100101, China Regulo Carlos Llarena Hernandez Facultad de Ciencias Biológicas y Agropecuarias, Peñuela, Universidad Veracruzana, Amatlán de los Reyes, Ver., Mexico Esteve Llop Facultat de Biologia, Universitat de Barcelona, Av. Diagonal, 643, 08028 Barcelona, Spain Csaba Locsmándi Hungarian Natural History Museum, 1087 Budapest, Hungary Deborah Jean Lodge USDA Forest Service, NRS, P.O. Box 1377, Luquillo, Puerto Rico 00773-1377, U.S.A. Michael Loizides P.O. Box 58499, 3734 Limassol, Cyprus László Lőkös Hungarian Natural History Museum, 1087 Budapest, Hungary Jennifer Luangsa-ard National Center for Genetic Engineering and Biotechnology (BIOTEC), NSTDA, 113 Thailand Science Park, Phaholyothin Rd., Klong Nueng, Klong Luang, Pathum Thani 12120, Thailand Matthias Lüderitz AG Geobotanik Schleswig-Holstein & Hamburg, c/o University of Kiel, Olshausenstraße 75, 24098 Kiel, Germany Thorsten Lumbsch Field Museum, 1400 S. Lake Shore Dr., Chicago, Illinois 60605, U.S.A. Matthias Lutz Institute of Evolution and Ecology, University of Tübingen, Auf der Morgenstelle 5, 72076 Tübingen, Germany Dan Mahoney Callaghan Innovation, 69 Gracefield Road, Lower Hutt 5010, New Zealand Ekaterina Malysheva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Vera Malysheva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Patinjareveettil Manimohan University of Calicut, Kerala, 673 635, India Yasmina Marin-Felix Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Guilhermina Marques Centre for the Research and Technology of Agro-Environmental and Biological Sciences, University of Tras-os-Montes and Alto Douro, Laboratory of Mycology and Soil Microbiology, 5000-801 Vila Real, Portugal Rubén Martínez-Gil Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Guy Marson Musée national d’histoire naturelle, 25 rue Münster, 2160 Luxembourg, Luxembourg Gerardo Mata Instituto de Ecología, A.C., Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, Mexico P. Brandon Matheny University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Geir Harald Mathiassen Tromsø University Museum, University of Tromsø — The Arctic University of Norway, 9037 Tromsø, Norway Neven Matočec Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Helmut Mayrhofer Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Mehdi Mehrabi Iranian Research Institute of Plant Protection, Agricultural Research, Education and Extension Organization, Tehran, Iran Ireneia Melo Botanical Garden, National Museum of Natural History and Science, University of Lisbon, Portugal Armin Mešić Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Andrew S. Methven Savannah State University, Savannah, Georgia 31404, U.S.A Otto Miettinen Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Ana M. Millanes Romero Universidad Rey Juan Carlos, 28933 Móstoles, Madrid, Spain Andrew N. Miller Illinois Natural History Survey, University of Illinois Urbana-Champaign, 1816 South Oak Street, Champaign, Illinois 61820-6970, U.S.A. James K. Mitchell Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Roland Moberg Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Pierre-Arthur Moreau Université Lille, Fac. Pharma. Lille, EA4483 IMPECS, 59000 Lille, France Gabriel Moreno Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Olga Morozova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Asunción Morte Facultad de Biología, Universidad de Murcia, Campus de Espinardo, 30100 Murcia, Spain Lucia Muggia University of Trieste, via L. Giorgieri 10, 34127 Trieste, Italy Guillermo Muñoz González Avda Valvanera N.32, 5D, 26500 Calahorra, La Rioja, Spain Leena Myllys Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland István Nagy Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary László G. Nagy Institute of Biochemistry, BRC-HAS, 6726 Szeged, Temesvari krt. 62, 6726 Szeged, Hungary Maria Alice Neves Universidade Federal de Santa Catarina, Campus Universitário Reitor João David Ferreira Lima, Trindade, Florianópolis, Santa Catarina CEP 88040-900, Brazil Tuomo Niemelä Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Pier Luigi Nimis University of Trieste, via L. Giorgieri 10, 34127 Trieste, Italy Nicolas Niveiro Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Machiel E. Noordeloos Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA, Leiden, The Netherlands Anders Nordin Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Sara Raouia Noumeur Faculty of Natural and Life Sciences, University of Batna 2, 05000 Batna, Algeria Yuri Novozhilov Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Jorinde Nuytinck Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands Esteri Ohenoja Botanical Museum, University of Oulu, Finland Patricia Oliveira Fiuza Universidade Estadual de Feira de Santana, Av. Transnordestina, s/n, Bairro Novo Horizonte, CEP:44036-900, Feira de Santana, Bahia, Brazil Alan Orange National Museum of Wales, Cardiff CF10 3NP, United Kingdom Alexander Ordynets University of Kassel, Heinrich-Plett-Strasse 40, 34132 Kassel, Germany Beatriz Ortiz-Santana USDA Forest Service, Northern Research Station, One Gifford Pinchot Dr, Madison, Wisconsin 53726, U.S.A. Leticia Pacheco Universidad Autónoma Metropolitana-Iztapalapa, Av. San Rafael Atlixco 186, Col. Vicentina, 09340 México, D. F., Mexico Ferenc Pál-Fám Faculty of Agricultural and Environmental Sciences, Kaposvar University, 7400 Kaposvar, Hungary Melissa Palacio Universidade Federal do Rio Grande do Sul, Brazil Zdeněk Palice Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Viktor Papp Szent Istvan University, 1118 Budapest, Menesi st. 44, Hungary Kadri Pärtelv Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Julia Pawlowska Faculty of Biology, Biological and Chemical Research Centre, University of Warsaw, Żwirki i Wigury 101, 02-089 Warsaw, Poland Aurelia Paz Urb. La Llosa, 219, 39509 Villanueva de la Peña, Mazcuerras, Cantabria, Spain Ursula Peintner Institut für Mikrobiologie, Universität Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria Shaun Pennycook Manaaki Whenua Landcare Research, Private Bag 92170, Auckland 1072, New Zealand Olinto Liparini Pereira Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Pablo Pérez Daniëls University of Córdoba, 14071, Córdoba, Spain Miquel À. Pérez-De-Gregorio Capella C/ Pau Casals, 6, 1º, 1ª, 17001, Girona, Spain Carlos Manuel Pérez del Amo C/ Luis de Ulloa, 1, 7º I, 26004 Logroño, Navarra, Spain Sergio Pérez Gorjón Universidad de Salamanca, Avda. Licenciado Mendez Nieto s/n, 37007 Salamanca, Spain Sergio Pérez-Ortega Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Israel Pérez-Vargas Facultad de Farmacia, Universidad de La Laguna, c/ Astrofísico Sánchez s/n 38071 La Laguna, Tenerife, Canary Islands, Spain Brian A. Perry California State University East Bay, Hayward, California 94542, U.S.A. Jens H. Petersen Nøruplundvej 2, 8400 Ebeltoft, Denmark Ronald H. Petersen University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Donald H. Pfister Harvard University, 22 Divinity Avenue, Cambridge MA 02138, U.S.A. Chayanard Phukhamsakda Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Marcin Piątek W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Meike Piepenbring Faculty of Biosciences, Goethe University Frankfurt am Main, Germany Raquel Pino-Bodas Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Juan Pablo Pinzón Esquivel Universidad Autónoma de Yucatán, Carretera Mérida-Xmatkuil Km. 15.5, Apdo. Postal: 4-116 Itzimná,C.P: 97100, Merida, Yucatan, Mexico Paul Pirot Rue des Peupliers 10, 6840 Neufchâteau, Belgium Eugene S. Popov Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Orlando Popoff Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina María Prieto Álvaro Universidad Rey Juan Carlos, 28933 Móstoles, Madrid, Spain Christian Printzen Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Abteilung Botanik und Molekulare Evolutionsforschung, Herbarium Senckenbergianum (FR), Senckenberganlage 25, 60325 Frankfurt am Main, Germany Nadezhda Psurtseva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Witoon Purahong Helmholtz Centre for Environmental Research GmbH — UFZ, Theodor-Lieser-Straße 4, 06120 Halle, Germany Luis Quijada Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Gerhard Rambold University of Bayreuth, Universitätsstrasse 30, 95447 Bayreuth, Germany Natalia A. Ramírez Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Huzefa Raja University of North Carolina at Greensboro, 435 Sullivan Science Building, PO Box 26170, Greensboro North Carolina 27402-6170, U.S.A. Olivier Raspé Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Tania Raymundo Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Prolongación de Carpio y Plan de Ayala s/n, Miguel Hidalgo, Santo Tomás, 11340 Ciudad de México, Mexico Martina Réblová Institute of Botany, Czech Academy of Sciences, Pruhonice, Czech Republic Yury A. Rebriev Southern Scientific Center, Russian Academy of Sciences, 41 Chehova str., Rostov-on-Don, 344006, Russia Juan de Dios Reyes García Paseo Virgen de Linarejos 6 2 D, Linares, Jaen, Spain Miguel Ángel Ribes Ripoll Avda. Pablo Neruda 120 F, 2°D, 28018 Madrid, Spain Franck Richard CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Mike J. Richardson Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, United Kingdom Víctor J. Rico Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Gerardo Lucio Robledo Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina Flavia Rodrigues Barbosa Universidade Federal de Mato Grosso, Av. Alexandre Ferronato, 1200, Setor Industrial, Sinop, Mato Grosso, Brazil Cristina Rodriguez-Caycedo UT Southwestern Medical Center, Dallas, Texas 75390, U.S.A. Pamela Rodriguez-Flakus W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Anna Ronikier W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Luis Rubio Casas Ul. Andrieja Sacharowa 1/1, 30-806 Kraków, Poland Katerina Rusevska Institute of Biology, Faculty of Natural Science and Mathematics, Ss. Cyril and Methodius University, Arhimedova 5, 1000 Skopje, Republic of Macedonia Günter Saar Dammenmühle 7, 77933 Lahr-Sulz, Germany Irja Saar Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Isabel Salcedo University of the Basque Country (UPV/EHU), Apdo 644, 48080 Bilbao, Spain Sergio M. Salcedo Martínez Facultad de Ciencias Biológicas, UANL. Ave. Pedro de Alba s/n esq. Manuel Barragán Cd. Universitaria, San Nicolás de los Garza Nuevo León, CP. 66451, Mexico Carlos A. Salvador Montoya Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Santiago Sánchez-Ramírez University of Toronto, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada J. Vladimir Sandoval-Sierra Health Science Centre, University of Tennessee, U.S.A. Sergi Santamaria Facultat de Biociències, Edifici C, Despatx C1/331, Campus de la UAB, 08193 Bellaterra, (Cerdanyola del Vallès), Barcelona, Spain Josiane Santana Monteiro Botany Coordination, Museu Paraense Emílio Goeldi, 66077-830, Belém, Pará, Brazil Hans Josef Schroers Agricultural Institute of Slovenia, Hacquetova ulica 17, 1000 Ljubljana, Slovenia Barbara Schulz Institute of Microbiology, Technische Universität Braunschweig, Germany Geert Schmidt-Stohn Burgstr. 25, 29553 Bienenbüttel, Germany Trond Schumacher University of Oslo, P.O.Box 1066, Blindern, 0316 Oslo, Norway Beatrice Senn-Irlet RU Biodiversity and Conservation Biology, Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Zürcherstr. 111, 8903 Birmensdorf, Switzerland Hana Ševčíková Moravian Museum, Zeny trh 6, 659 37 Brno, Czech Republic Oleg Shchepin Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Takashi Shirouzu Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan Anton Shiryaev Institute of Plant & Animal Ecology Ural Branch of the Russian Academy of Sciences, 620144 Ekaterinburg, 8 March str., 202/3, Russia Klaus Siepe Geeste 133, 46342 Velen, Germany Esteban B. Sir Fundación Miguel Lillo, Consejo Nacional de Investigaciones Científicas y Técnicas, Miguel Lillo 251, San Miguel de Tucumán 4000, Tucumán, Argentina Mohammad Sohrabi Iranian Research Organization for Science and Technology, P.O. Box 33535111, Tehran, Iran Karl Soop Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Viacheslav Spirin Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Toby Spribille University of Alberta, Edmonton, Alberta T6G 2R3, Canada Marc Stadler Helmholtz Centre for Infection Research, Inhoffenstrasse 7, 38124 Braunschweig, Germany Joost Stalpers Torenlaan 43, 3742CR Baarn, The Netherlands Soili Stenroos Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Ave Suija Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Stellan Sunhede Hökaskog Sandbacken 1, 533 92 Lundsbrunn, Sweden Sten Svantesson University of Gothenburg, Box 461, 40530 Göteborg, Sweden Sigvard Svensson Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Tatyana Yu. Svetasheva (1) Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia; (2) Department of Technologies of Living Systems, Tula State Lev Tolstoy Pedagogical University, Lenin ave. 125, Tula, 300026, Russia Krzysztof Świerkosz Museum of Natural History, Wrocław University, ul. H. Sienkiewicza 5, 50-335 Wrocław, Poland Heidi Tamm Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Hatira Taskin Faculty of Agriculture, University of Çukurova, 01330 Adana, Turkey Adrien Taudière CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Jan-Olof Tedebrand Sundsvall Mycological Society, Medelpad, Sweden Raúl Tena Lahoz C/Arreñales del Portillo B 21 1°D, 44003 Teruel, Spain Marina Temina Institute of Evolution, University of Haifa, Aba Khoushi Ave. 199, Mt. Carmel, Haifa 3498838, Israel Arne Thell Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Marco Thines Faculty of Biosciences, Goethe University Frankfurt am Main, Germany Göran Thor Swedish University of Agricultural Sciences, P. O. Box 7044, 750 07 Uppsala, Sweden Holger Thüs State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany Leif Tibell Evolutionary Biology Centre, Norbyvägen 18D, 75236 Uppsala, Sweden Sanja Tibell Evolutionary Biology Centre, Norbyvägen 18D, 75236 Uppsala, Sweden Einar Timdal Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Zdenko Tkalčec Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Tor Tønsberg University of Bergen, Allégaten 41, P.O. Box 7800, 5020 Bergen, Norway Gérard Trichies 5, impasse des Écoles S7700 Neufchef, France Dagmar Triebel Botanische Staatssammlung München, Menzinger Straße 67, 80638 München, Germany Andrei Tsurykau (1) F. Skorina Gomel State University, Sovetskaja Str. 104, 246019 Gomel, Belarus; (2) Institute of Natural Sciences, Samara National Research University, Moskovskoye shosse 34, 443086 Samara, Russia Rodham E. Tulloss Herbarium Amanitarum Rooseveltensis, P. O. Box 57, Roosevelt, New Jersey 08555-0057, U.S.A. Veera Tuovinen University of Alberta, Edmonton, Alberta T6G 2R3, Canada Miguel Ulloa Sosa Universidad Nacional Autónoma de México, Tercer Circuito/Sin Número, Ciudad Universitaria, Ciudad de México, C. P. 04510, Mexico Carlos Urcelay Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina François Valade 11 rue haras, le boqueteau, 91240 Saint-Michel-sur-Orge, France Ricardo Valenzuela Garza Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Prolongación de Carpio y Plan de Ayala s/n, Miguel Hidalgo, Santo Tomás, 11340 Ciudad de México, Mexico Pieter van den Boom Arafura 16, 5691 JA Son, The Netherlands Nicolas Van Vooren Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Aida M. Vasco-Palacios Escuela de Microbiología, Universidad de Antioquia, AA1226, Fundación Biodiversa Colombia, Medellín, Colombia Jukka Vauras Herbarium TUR, Biodiversity Unit, University of Turku, 20014 Turku, Finland Juan Manuel Velasco Santos 3 C/Pontevedra, 18, 1º C, 37003 Salamanca, Spain Else Vellinga 861 Keeler Avenue, Berkeley, California 94708, U.S.A. Annemieke Verbeken Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Per Vetlesen Norges Sopp- og Nyttevekstforbund, Schweigaards gate 34F, 0191 Oslo, Norway Alfredo Vizzini University of Torino, Viale P.A. Mattioli 25, 10125, Torino, Italy Hermann Voglmayr University of Vienna, Rennweg 14, 1030 Vienna, Austria Sergey Volobuev Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Wolfgang von Brackel Kirchenweg 2, 91341 Röttenbach, Germany Elena Voronina Lomonosov Moscow State University, Biology Faculty, Moscow, Russia Grit Walther Arvid-Harnack-Str. 4, 07743 Jena, Germany Roy Watling Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, United Kingdom Evi Weber Blaihofstr. 42, 72074 Tübingen, Germany Mats Wedin Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Øyvind Weholt Nord University, Nesna, 8700 Nesna, Norway Martin Westberg Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Eugene Yurchenko Polessky State University, Dnyaprouskai flatylii str. 23, 225710, Pinsk, Belarus Petr Zehnálek Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Huang Zhang Kunming University of Science & Technology, Kunming, China Mikhail P. Zhurbenko Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia The following institutions are also supporting the present text (institutional support means that committees from the mentioned) Asociación Micológica Carlos Spegazzini (Argentina) Austrian Mycological Society (Austria) Croatian Mycological Society (Croatia) Committee of the Czech Scientific Society for Mycology (Czech Republic) The Netherlands Mycological Society (The Netherlands) Sociedad Española de Liquenología (Portugal/Spain) Iberian Mycological Society (which is under constitution, Portugal/Spain) Federación de Asociaciones Micológicas Andaluzas (FAMA) (Spain) Asociación Botánica y Micológica de Jaén (Spain) Asociación Micológica Hispalense Muscaria (Spain) Societat Micològica Valenciana (Spain)

Here we assess the impact of geographically dependent (latitude, longitude, and altitude) changes in bioclimatic (temperature, precipitation, and primary productivity) variability on fungal fruiting phenology across Europe. Two main nutritional guilds of fungi, saprotrophic and ectomycorrhizal, were further separated into spring and autumn fruiters. We used a path analysis to investigate how biogeographic patterns in fungal fruiting phenology coincided with seasonal changes in climate and primary production. Across central to northern Europe, mean fruiting varied by approximately 25 d, primarily with latitude. Altitude affected fruiting by up to 30 d, with spring delays and autumnal accelerations. Fruiting was as much explained by the effects of bioclimatic variability as by their large-scale spatial patterns. Temperature drove fruiting of autumnal ectomycorrhizal and saprotrophic groups as well as spring saprotrophic groups, while primary production and precipitation were major drivers for spring-fruiting ectomycorrhizal fungi. Species-specific phenology predictors were not stable, instead deviating from the overall mean. There is significant likelihood that further climatic change, especially in temperature, will impact fungal phenology patterns at large spatial scales. The ecological implications are diverse, potentially affecting food webs (asynchrony), nutrient cycling and the timing of nutrient availability in ecosystems.

We assessed how environmental drivers influence the occurrences of wood-inhabiting macrofungi in European beech forests, using an extensive dataset of fruit body records collected in 53 reserves across twelve European countries. We found that the 105 species included in this study varied greatly in their responses to covariates related to resource quality, climate and forest connectivity, both in the strength and direction of the observed effects. Climate was the most important driver for some species, while others responded more to connectivity, or simply to the presence of high quality substrates within the reserves. Species occurrences varied also across geographical regions, especially between the UK and the rest of Europe. Our results show that wood-inhabiting fungi in European beech forests respond individualistically to habitat filters and differ in their biogeographical distribution patterns, and they thus provide a detailed perspective of how wood-inhabiting fungal communities are structured across Europe.

Ectomycorrhizal and saprotrophic agaricoid basidiomycetes show diverse morphological reproductive traits, a phenomenon which has been attributed to their different lifestyles. From previous studies, we know that such differences are also reflected in assembly formation. Regardless of these differences, and assuming that dispersal fitness, predominantly by air movement, is one of the prevalent factors in fungal lifecycles, spores of both guilds should become on average more elongate and smaller with resource depletion. In our study we defined resource depletion as the decrease of living and dead organic biomass due to climate constraints along an elevational gradient in the Bavarian Forest (Germany). We found that spores of both guilds indeed become more elongate along the resource depletion gradient. Unexpectedly, the ectomycorrhizal assemblages showed larger spores under resource constraints, which could be a survivability trade-off. The spore trait syndrome responses to environmental constraints suggest ecological relevance, i.e. being advantageous in environments with patchily distributed resources. To deepen our mechanistic understanding of the underlying patterns, we particularly recommend experiments (artificial resource gradient free of confounding effects such as climate), and application of genomics and transcriptomics for elucidating the evolution of spore morphology.

The Danish Fungal Atlas was a citizen science project aiming to map Danish macrofungi over five years (2009–13). The atlas contributed >235,000 records of fruit-body forming Basidiomycota, adding to about 195,000 fungal records from earlier periods. The new records increased the average number of species known per 10 km × 10 km grid cell by 75% from 125 to 218 species. We recorded 197 species as new to Denmark, extending the number of known basidiomycote species by 7%. At least 15 species appeared to be new to science. Among the new Danish records, species with northern distribution ranges were significantly overrepresented, in marked contrast to climate change predictions. Species with inconspicuous or subterranean fruit bodies were overrepresented among both the new Danish species and those only recorded before the project period, indicating low recording probability to be an important driver for the turnover in species recorded. Hence, the main drivers of novel fungal discoveries were 1) intensive sampling effort by citizen scientists guided by professional mycologists and 2) improved taxonomic knowledge. Summarizing over the last 100 years, an exponential increase in known macrofungal diversity in Denmark is evident, suggesting that we are still far from having a complete overview. This is striking, considering that Denmark is among the best-studied land areas on the globe. We conclude that citizen science projects, if appropriately designed, have a huge potential to boost primary knowledge on fungal biodiversity.

The majority of Earth's biodiversity is unknown. This is particularly true for the vast part of soil biodiversity, which rarely can be observed directly. Metabarcoding of DNA extracted from the environment (eDNA) has become state-of-the-art in assessing soil biodiversity. For assessing fungal communities, eDNA metabarcoding is seen as an attractive alternative to classical surveying based on fruitbodies. However, it is unknown whether eDNA metabarcoding provides a representative sample of fungal diversity and census of threatened species. Therefore conservation planning and assessment are still based on fruitbody inventories. Based on a dataset of large ecological width, representing both soil eDNA metabarcoding and expert inventories of fungal fruitbodies in Denmark, we document for the first time the validity of eDNA sampling and metabarcoding as a practical inventory method and a measure of conservation value for fungi. Fruitbody data identified fewer species in total and per site, and had larger variance in site richness. Focusing on macrofungi – the class Agaricomycetes, and in turn the order Agaricales – metrics of total richness and compositional similarity converged between the methods. eDNA was suboptimal for recording the non-soil dwelling fungi such as lichens and polypores. β-Diversity was similar between methods, but more variation in community composition could be explained by environmental predictors in the eDNA data. The fruitbody survey was slightly better at finding red listed species. We find a better correspondence between biodiversity indices derived from fungal fruitbodies and DNA-based approaches than indicated in earlier studies. We argue that (historical) fungal community data based on fruitbody forays – with careful selection of taxonomic groups – may be interpreted together with modern DNA-based approaches to obtain a richer picture of the full mycobiota of the site, and for addressing historical changes. We estimated the costs of the two inventory approaches to be approximately similar for practical applications.

Based on molecular and morphological data we investigated the taxonomy and phylogeny of the ectomycorrhizal genus Tricholoma in northern Europe. Our phylogenetic tree confirmed the presence of at least 72 well circumscribed species within the region. Of these, three species, viz. T. boreosulphurescens, T. bryogenum and T. ilkkae are described as new to science, based on morphological, distributional, ecological and molecular data. Several other terminal branches represent putative cryptic taxa nested within classical species or species groups. Molecular type studies and/or designation of sequenced neotypes are needed in these groups, before the taxonomy can be settled. In general our phylogenetic analysis supported previous suprageneric classification systems, but with some substantial changes. Most notably, T. virgatum and allies were found to belong to sect. Tricholoma rather than sect. Atrosquamosa, while T. focale was found to be clearly nested in sect. Genuina rather than in sect. Caligata. In total, ten sections are accepted, with five species remaining unassigned. The combination of morphological and molecular data showed pileus colour, pileipellis structure, presence of clamp connections and spore size to be rather conservative characters within accepted sections, while the presence of a distinct ring, and especially host selection were highly variable within these.

The biodiversity value of production forests is substantially lower than that of natural forests. This is related to differences in hydrology, stand age and amounts of old trees and deadwood. Using a predictive model framework we show that restoring hydrology and old-growth characteristics in a forest formerly managed for timber extraction results in changes to forest composition and structure, ultimately increasing its biodiversity value. We inventoried biodiversity and stand variables in 102 sample plots in a temperate mixed broadleaved forest, which is in focus of a LIFE+ programme aiming to restore hydrology and old-growth structure. We collected presence/absence data for four organism groups (vascular plants, epiphytic bryophytes and lichens, wood-inhabiting fungi) and measured environmental variables associated with species occurrence and influenced by restoration (dead or living tree characteristics, stand age, water level). We investigated biodiversity consequences of restoration towards pristine environmental characteristics by using a space-for-time substitution model. We evaluated how and through what mechanisms species richness is likely to react when pre-forestry hydrological conditions and old-growth structures are restored. The model results show that reversing the effects of a long history of management for timber extraction increased availability of suitable habitat, and hence the local species richness for three of four of the organism groups, compared to the pre-restoration conditions. Furthermore, the increase in soil moisture shifted the forest plots towards an alder carr, while the stand ageing process sustained the shade-tolerant beech despite its low tolerance for high soil humidity. Our prediction shows an increase in species richness for plants directly driven by the restoration of natural water level, and for fungi as an indirect effect of a change in suitable substrate availability. Lichens responded positively to both processes. Plants stabilized their richness levels earlier than tree-dwelling organisms, as water level recovered faster than old-growth structures. The projection of stable bryophytes richness values under restoration is potentially biased by their lower diversity and more limited affiliation to forest structural variation than other groups. We suggest applying our space-for-time approach as a tool to assess forest and biodiversity responses in similar restoration projects involving management actions of open-ended habitat creation, promoting development of natural processes in the long-term. This modelling tool turns to be especially relevant in dynamic habitats where the outcomes for biodiversity are uncertain.

Reintroduction of locally extinct species is increasingly applied as a conservation tool for re-establishing species within their historical ranges. Thus far, this option has however not been investigated for fungi other than lichens. A large fraction of wood-inhabiting fungal species have declined because of forest loss and fragmentation, in addition to a decrease in dead wood. Here, we show the results from an experiment carried out in southern Finland, which demonstrates that inoculation is an effective method for reintroducing threatened wood-inhabiting fungi. All selected red-listed fungal species successfully established in the inoculated logs as mycelia, and three out of the seven produced fruit-bodies. Success rate was greater when the strains were inoculated in early-decay logs, including species that usually fruit in late decay stages. Inoculation can provide an effective tool for reintroducing fungal species, as the source populations remain intact and it is possible to produce massive amounts of inoculation-units with relatively low cost. Reintroductions of fungi should however be preceded by a risk assessment of the species to be reintroduced, by using source populations from nearby localities, and they should be considered complementary to the primary target of increasing the amount of their habitat. Our results suggest that the reintroductions of threatened fungi via inoculation in combination with other conservation measures can have important bearings for forest conservation and restoration.

The conservation status of European temperate forests is overall unfavorable, and many associated species are listed in national or European red-lists. A better understanding of factors increasing survival probability of red-listed species is needed for a more efficient conservation effort. Here, we investigated the importance of current forest cover, historical forest cover and a number of soil and climate variables on the incidence and richness of red-listed forest species in Denmark. We considered eight major taxa separately (mammals, saproxylic beetles, butterflies, vascular plants and four groups of fungi), using mainly citizen science data from several national mapping projects. Taxa were selected to represent important forest habitats or properties (soil, dead wood, forest glades and landscape context) and differ in dispersal potential and trophic strategy. For all groups, presence and richness of red-listed species was positively related with current forest cover, but-for most taxa-forest cover 200 years ago was an even better predictor. The intersection of past and current deciduous forest was used to identify the area of continuous, lost and new forest. Continuous and lost deciduous forest cover were strong predictors of red-listed species occurrence in most groups, but surprisingly species richness of butterflies and hydnoid fungi, and presence of mammals, was significantly, positively affected by coniferous forest area. The positive effect of lost deciduous forests on red-listed species, suggest an extinction debt of at least 200 years, with some areas hosting more red-listed species than the current area of old forest can sustain in the long run. Our results suggest that current priorities for forest conservation in Denmark are not efficient in protecting red-listed forest species, and that more focus should be put on conserving deciduous forest with long continuity. Furthermore, a multi-taxa approach including a wide array of organism groups with contrasting habitat affiliations, results in a more comprehensive understanding of the requirements of red-listed forest species and necessitate a more focused approach to conservation planning. (C) 2016 Elsevier B.V. All rights reserved.

Aim To understand how the integration of contextual spatial data on land cover and human infrastructure can help reduce spatial bias in sampling effort, and improve the utilization of citizen science-based species recording schemes. By comparing four different citizen science projects, we explore how the sampling design's complexity affects the role of these spatial biases. Location Denmark, Europe. Methods We used a point process model to estimate the effect of land cover and human infrastructure on the intensity of observations from four different citizen science species recording schemes. We then use these results to predict areas of under- and oversampling as well as relative biodiversity ‘hotspots’ and ‘deserts’, accounting for common spatial biases introduced in unstructured sampling designs. Results We demonstrate that the explanatory power of spatial biases such as infrastructure and human population density increased as the complexity of the sampling schemes decreased. Despite a low absolute sampling effort in agricultural landscapes, these areas still appeared oversampled compared to the observed species richness. Conversely, forests and grassland appeared undersampled despite higher absolute sampling efforts. We also present a novel and effective analytical approach to address spatial biases in unstructured sampling schemes and a new way to address such biases, when more structured sampling is not an option. Main conclusions We show that citizen science datasets, which rely on untrained amateurs, are more heavily prone to spatial biases from infrastructure and human population density. Objectives and protocols of mass-participating projects should thus be designed with this in mind. Our results suggest that, where contextual data is available, modelling the intensity of individual observation can help understand and quantify how spatial biases affect the observed biological patterns.

The richness of saproxylic species is commonly found to be compromised in managed forests. However, it is often less clear if this reflects local scarcity of high quality deadwood objects or changes to the overall forest structure. Further, anthropogenic and natural drivers working at the landscape scale may have a strong impact. Hence, it is difficult to give detailed advice on how to conserve biodiversity in a cost-effective manner. Here we explored macrofungal richness on deadwood across a gradient of forest management intensity in Danish Fagus sylvatica forests. We aimed to disentangle the importance of habitat quality per se from the effects of variables recorded at the forest stand and landscape scales. Data was collected from 40 beech stands, each representing one of four broad management types: conventionally managed, near-to-nature managed, recently unmanaged and long unmanaged. Stands were aggregated within six larger forest clusters, to account for variation at the landscape scale. Fungal species were recorded as fruitbodies on deadwood at five decay stages. We found that the four management types showed strong differences in fungal richness, deadwood volume and forest structure, with long unmanaged stands having notably higher structural complexity and species richness. Fungal species richness and presence of red-listed fungi appeared to be mainly driven by deadwood volume, tree species and decay stage, but some stand-level variables such as canopy gaps, basal area and stand age were also of importance. Lying coarse deadwood had the highest species richness while standing coarse woody debris and fine deadwood had lower species density. Saproxylic fungal richness on individual deadwood objects was largely affected by resource quality and size, which is often limited in managed forests. Our results indicate that the best measure to increase fungal species richness, especially in managed stands, is to increase the amount of large diameter deadwood.

International audience; Aim: The tinder fungus Fomes fomentarius is a pivotal wood decomposer in European beech Fagus sylvatica forests. The fungus, however, has regionally declined due to centuries of logging. To unravel biogeographical drivers of arthropod communities associated with this fungus, we investigated how space, climate and habitat amount structure alpha and beta diversity of arthropod communities in fruitbodies of F. fomentarius. Location: Temperate zone of Europe. Taxon: Arthropods. Methods: We reared arthropods from fruitbodies sampled from 61 sites throughout the range of European beech and identified 13 orders taxonomically or by metabarcoding. We estimated the total number of species occurring in fruitbodies of F. fomentarius in European beech forests using the Chao2 estimator and determined the relative importance of space, climate and habitat amount by hierarchical partitioning for alpha diversity and generalized dissimilarity models for beta diversity. A subset of fungi samples was sequenced for identification of the fungus' genetic structure. Results: The total number of arthropod species occurring in fruitbodies of F. fomentarius across European beech forests was estimated to be 600. Alpha diversity increased with increasing fruitbody biomass; it decreased with increasing longitude, temperature and latitude. Beta diversity was mainly composed by turnover. Patterns of beta diversity were only weakly linked to space and the overall explanatory power was low. We could distinguish two genotypes of F. fomentarius, which showed no spatial structuring. Main conclusion: Fomes fomentarius hosts a large number of arthropods in European beech forests. The low biogeographical and climatic structure of the communities suggests that fruitbodies represent a habitat that offers similar conditions across large gradients of climate and space, but are characterized by high local variability in community composition and colonized by species with high dispersal ability. For European beech forests, retention of trees with F. fomentarius and promoting its recolonization where it had declined seems a promising conservation strategy.

Premise of the Study Fungal diversity (richness) trends at large scales are in urgent need of investigation, especially through novel situations that combine long‐term observational with environmental and remotely sensed open‐source data. Methods We modeled fungal richness, with collections‐based records of saprotrophic (decaying) and ectomycorrhizal (plant mutualistic) fungi, using an array of environmental variables across geographical gradients from northern to central Europe. Temporal differences in covariables granted insight into the impacts of the shorter‐ versus longer‐term environment on fungal richness. Results Fungal richness varied significantly across different land‐use types, with highest richness in forests and lowest in urban areas. Latitudinal trends supported a unimodal pattern in diversity across Europe. Temperature, both annual mean and range, was positively correlated with richness, indicating the importance of seasonality in increasing richness amounts. Precipitation seasonality notably affected saprotrophic fungal diversity (a unimodal relationship), as did daily precipitation of the collection day (negatively correlated). Ectomycorrhizal fungal richness differed from that of saprotrophs by being positively associated with tree species richness. Discussion Our results demonstrate that fungal richness is strongly correlated with land use and climate conditions, especially concerning seasonality, and that ongoing global change processes will affect fungal richness patterns at large scales. publishedVersion