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The genus Morchella has gone through turbulent taxonomic treatments. Although significant progress in Morchella systematics has been achieved in the past decade, several problems remain unresolved and taxonomy in the genus is still in flux. In late 2019, a paper published in the open-access journal Scientific Reports raised serious concerns about the taxonomic stability of the genus, but also about the future of academic publishing. The paper, entitled “High diversity of Morchella and a novel lineage of the esculenta clade from the north Qinling Mountains revealed by GCPSR-based study” by Phanpadith and colleagues, suffered from gross methodological errors, included false results and artifactual phylogenies, had misapplied citations throughout, and proposed a new species name invalidly. Although the paper was eventually retracted by Scientific Reports in 2021, the fact that such an overtly flawed and scientifically unsound paper was published in a high-ranked Q1 journal raises alarming questions about quality controls and safekeeping procedures in scholarly publishing. Using this paper as a case study, we provide a critical review on the pitfalls of Morchella systematics followed by a series of recommendations for the delimitation of species, description of taxa, and ultimately for a sustainable taxonomy in Morchella. Problems and loopholes in the academic publishing system are also identified and discussed, and additional quality controls in the pre- and post-publication stages are proposed.

There were errors in the name of author Laszlo G. Nagy and in affiliation no. 31 in the original publication. The original article has been corrected.

The study deals with the last unexplored morphological group of the genus Hodophilus defined by absence of distinct odours, absence of yellow colours and absence of darker dots on the stipe. The phylogenetic reconstruction of the whole genus based on nrITS, nrLSU and RPB2 sequences placed all European members having these morphological characters in a monophyletic group defined here as a new section H. sect. Phaeophylli. The remaining European members of the genus are placed in two additional groups classified as section H. sect. Hodophilus and the new section H. sect. Micacei. Five species are recognised within section Phaeophylli which is typified by H. phaeophyllus that is lecto- and epitypified. Three new species belonging to this section are described: H. carpathicus, H. decurrentior and H. stramineus. Hodophilus decurrentior is the only species showing distinct morphological differences under the microscope. The identification of other species of the section depends mainly on the colour of basidiomata. An updated key to all European members of the genus is provided.

Hebeloma parvisporum is described as new and placed within H. sect. Porphyrospora. This mushroom is sold as an edible in markets of Laos under the local name “wai khom.” Hebeloma sect. Porphyrospora is discussed and expanded to include the species formerly included in the genus Anamika and recently transferred to Hebeloma. Hebeloma sect. Porphyrospora currently comprises 16 species, 14 of which are known only from the western Pacific and Indian subcontinent. All species in this section share the character of having red-brown spores when fresh, atypical for other sections of Hebeloma, which causes the lamellae to be red-brown. However, this red-brown color fades when the material is dried. The close links, morphologically and molecularly, between H. parvisporum and other members of H. sect. Porphyrospora, particularly H. victoriense, are shown.

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.

Novel species of fungi described in this study include those from various countries as follows: Australia, Chaetomella pseudocircinoseta and Coniella pseudodiospyri on Eucalyptus microcorys leaves, Cladophialophora eucalypti, Teratosphaeria dunnii and Vermiculariopsiella dunnii on Eucalyptus dunnii leaves, Cylindrium grande and Hypsotheca eucalyptorum on Eucalyptus grandis leaves, Elsinoe salignae on Eucalyptus saligna leaves, Marasmius lebeliae on litter of regenerating subtropical rainforest, Phialoseptomonium eucalypti (incl. Phialoseptomonium gen. nov.) on Eucalyptus grandis × camaldulensis leaves, Phlogicylindrium pawpawense on Eucalyptus tereticornis leaves, Phyllosticta longicauda as an endophyte from healthy Eustrephus latifolius leaves, Pseudosydowia eucalyptorum on Eucalyptus sp. leaves, Saitozyma wallum on Banksia aemula leaves, Teratosphaeria henryi on Corymbia henryi leaves. Brazil, Aspergillus bezerrae, Backusella azygospora, Mariannaea terricola and Talaromyces pernambucoensis from soil, Calonectria matogrossensis on Eucalyptus urophylla leaves, Calvatia brasiliensis on soil, Carcinomyces nordestinensis on Bromelia antiacantha leaves, Dendryphiella stromaticola on small branches of an unidentified plant, Nigrospora brasiliensis on Nopalea cochenillifera leaves, Penicillium alagoense as a leaf endophyte on a Miconia sp., Podosordaria nigrobrunnea on dung, Spegazzinia bromeliacearum as a leaf endophyte on Tilandsia catimbauensis, Xylobolus brasiliensis on decaying wood. Bulgaria, Kazachstania molopis from the gut of the beetle Molops piceus. Croatia, Mollisia endocrystallina from a fallen decorticated Picea abies tree trunk. Ecuador, Hygrocybe rodomaculata on soil. Hungary, Alfoldia vorosii (incl.Alfoldia gen. nov.) from Juniperus communis roots, Kiskunsagia ubrizsyi (incl. Kiskunsagia gen. nov.) from Fumana procumbens roots. India, Aureobasidium tremulum as laboratory contaminant, Leucosporidium himalayensis and Naganishia indica from windblown dust on glaciers. Italy, Neodevriesia cycadicola on Cycas sp. leaves, Pseudocercospora pseudomyrticola on Myrtus communis leaves, Ramularia pistaciae on Pistacia lentiscus leaves, Neognomoniopsis quercina (incl. Neognomoniopsis gen. nov.) on Quercus ilex leaves. Japan, Diaporthe fructicola on Passiflora edulis × P. edulis f. flavicarpa fruit, Entoloma nipponicum on leaf litter in a mixed Cryptomeria japonica and Acer spp. forest. Macedonia, Astraeus macedonicus on soil. Malaysia, Fusicladium eucalyptigenum on Eucalyptus sp. twigs, Neoacrodontiella eucalypti (incl. Neoacrodontiella gen. nov.) on Eucalyptus urophylla leaves. Mozambique, Meliola gorongosensis on dead Philenoptera violacea leaflets. Nepal, Coniochaeta dendrobiicola from Dendriobium lognicornu roots. New Zealand, Neodevriesia sexualis and Thozetella neonivea on Archontophoenix cunninghamiana leaves. Norway, Calophoma sandfjordenica from a piece of board on a rocky shoreline, Clavaria parvispora on soil, Didymella finnmarkica from a piece of Pinus sylvestris driftwood. Poland, Sugiyamaella trypani from soil. Portugal, Colletotrichum feijoicola from Acca sellowiana. Russia, Crepidotus tobolensis on Populus tremula debris, Entoloma ekaterinae, Entoloma erhardii and Suillus gastroflavus on soil, Nakazawaea ambrosiae from the galleries of Ips typographus under the bark of Picea abies. Slovenia, Pluteus ludwigii on twigs of broadleaved trees. South Africa, Anungitiomyces stellenboschiensis (incl. Anungitiomyces gen. nov.) and Niesslia stellenboschiana on Eucalyptus sp. leaves, Beltraniella pseudoportoricensis on Podocarpus falcatus leaf litter, Corynespora encephalarti on Encephalartos sp. leaves, Cytospora pavettae on Pavetta revoluta leaves, Helminthosporium erythrinicola on Erythrina humeana leaves, Helminthosporium syzygii on a Syzygium sp. barkcanker, Libertasomyces aloeticus on Aloe sp. leaves, Penicillium lunae from Musa sp. fruit, Phyllosticta lauridiae on Lauridia tetragona leaves, Pseudotruncatella bolusanthi (incl. Pseudotruncatellaceae fam. nov.) and Dactylella bolusanthi on Bolusanthus speciosus leaves. Spain, Apenidiella foetida on submerged plant debris, Inocybe grammatoides on Quercus ilex subsp. ilex forest humus, Ossicaulis salomii on soil, Phialemonium guarroi from soil. Thailand, Pantospora chromolaenae on Chromolaena odorata leaves. Ukraine, Cadophora helianthi from Helianthus annuus stems. USA, Boletus pseudopinophilus on soil under slash pine, Botryotrichum foricae, Penicillium americanum and Penicillium minnesotense from air. Vietnam, Lycoperdon vietnamense on soil. Morphological and culture characteristics are supported by DNA barcodes.

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 cryptic lifestyle of most fungi necessitates molecular identification of the guild in environmental studies. Over the past decades, rapid development and affordability of molecular tools have tremendously improved insights of the fungal diversity in all ecosystems and habitats. Yet, in spite of the progress of molecular methods, knowledge about functional properties of the fungal taxa is vague and interpretation of environmental studies in an ecologically meaningful manner remains challenging. In order to facilitate functional assignments and ecological interpretation of environmental studies we introduce a user friendly traits and character database FungalTraits operating at genus and species hypothesis levels. Combining the information from previous efforts such as FUNGuild and FunFun together with involvement of expert knowledge, we reannotated 10,210 and 151 fungal and Stramenopila genera, respectively. This resulted in a stand-alone spreadsheet dataset covering 17 lifestyle related traits of fungal and Stramenopila genera, designed for rapid functional assignments of environmental studies. In order to assign the trait states to fungal species hypotheses, the scientific community of experts manually categorised and assigned available trait information to 697,413 fungal ITS sequences. On the basis of those sequences we were able to summarise trait and host information into 92,623 fungal species hypotheses at 1% dissimilarity threshold. Fungal traits · Trophic modes · Function · Guild · Bioinformatics · High-throughput sequencing · Community ecology

Novel species of fungi described in this study include those from various countries as follows: Algeria, Phaeoacremonium adelophialidum from Vitis vinifera. Antarctica, Comoclathris antarctica from soil. Australia, Coniochaeta salicifolia as endophyte from healthy leaves of Geijera salicifolia, Eremothecium peggii in fruit of Citrus australis, Microdochium ratticaudae from stem of Sporobolus natalensis, Neocelosporium corymbiae on stems of Corymbia variegata, Phytophthora kelmanii from rhizosphere soil of Ptilotus pyramidatus, Pseudosydowia backhousiae on living leaves of Backhousia citriodora, Pseudosydowia indoor oopillyensis, Pseudosydowia louisecottisiae and Pseudosydowia queenslandica on living leaves of Eucalyptus sp. Brazil, Absidia montepascoalis from soil. Chile, Ilyonectria zarorii from soil under Maytenus boaria. Costa Rica, Colletotrichum filicis from an unidentified fern. Croatia, Mollisia endogranulata on deteriorated hardwood. Czech Republic, Arcopilus navicularis from tea bag with fruit tea, Neosetophoma buxi as endophyte from Buxus sempervirens, Xerochrysium bohemicum on surface of biscuits with chocolate glaze and filled with jam. France, Entoloma cyaneobasale on basic to calcareous soil, Fusarium aconidiale from Triticum aestivum, Fusarium juglandicola from buds of Juglans regia. Germany, Tetraploa endophytica as endophyte from Microthlaspi perfoliatum roots. India, Castanediella ambae on leaves of Mangifera indica, Lactifluus kanadii on soil under Castanopsis sp., Penicillium uttarakhandense from soil. Italy, Penicillium ferraniaense from compost. Namibia, Bezerromyces gobabebensis on leaves of unidentified succulent, Cladosporium stipagrostidicola on leaves of Stipagrostis sp., Cymostachys euphorbiae on leaves of Euphorbia sp., Deniquelata hypolithi from hypolith under a rock, Hysterobrevium walvisbayicola on leaves of unidentified tree, Knufia hypolithi and Knufia walvisbayicola from hypolith under a rock, Lapidomyces stipagrostidicola on leaves of Stipagrostis sp., Nothophaeotheca mirabibensis (incl. Nothophaeotheca gen. nov.) on persistent inflorescence remains of Blepharis obmitrata, Paramyrothecium salvadorae on twigs of Salvadora persica, Preussia procaviicola on dung of Procavia sp., Sordaria equicola on zebra dung, Volutella salvadorae on stems of Salvadora persica. Netherlands, Entoloma ammophilum on sandy soil, Entoloma pseudocruentatum on nutrient poor(acid)soil, Entoloma pudens on plant debris, amongst grasses. [...], Leslie W.S. de Freitas and colleagues express their gratitude to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for scholarships provided to Leslie Freitas and for the research grant provided to André Luiz Santiago; their contribution was financed by the projects ‘Diversity of Mucoromycotina in the different ecosystems of the Atlantic Rainforest of Pernambuco’ (FACEPE–First Projects Program PPP/ FACEPE/CNPq–APQ–0842-2.12/14) and ‘Biology of conservation of fungi s.l. in areas of Atlantic Forest of Northeast Brazil’ (CNPq/ICMBio 421241/ 2017-9) H.B. Lee was supported by the Graduate Program for the Undiscovered Taxa of Korea (NIBR202130202). The study of O.V. Morozova, E.F. Malysheva, V.F. Malysheva, I.V. Zmitrovich, and L.B. Kalinina was carried out within the framework of a research project of the Komarov Botanical Institute RAS (АААА-А19-119020890079-6) using equipment of its Core Facility Centre ‘Cell and Molecular Technologies in Plant Science’. The work of O. V. Morozova, L.B. Kalinina, T. Yu. Svetasheva, and E.A. Zvyagina was financially supported by Russian Foundation for Basic Research project no. 20-04-00349. E.A. Zvyagina and T.Yu. Svetasheva are grateful to A.V. Alexandrova, A.E. Kovalenko, A.S. Baykalova for the loan of specimens, T.Y. James, E.F. Malysheva and V.F. Malysheva for sequencing. J.D. Reyes acknowledges B. Dima for comparing the holotype sequence of Cortinarius bonachei with the sequences in his database. A. Mateos and J.D. Reyes acknowledge L. Quijada for reviewing the phylogeny and S. de la Peña- Lastra and P. Alvarado for their support and help. Vladimir I. Kapitonov and colleagues are grateful to Brigitta Kiss for help with their molecular studies. This study was conducted under research projects of the Tobolsk Complex Scientific Station of the Ural Branch of the Russian Academy of Sciences (N АААА-А19-119011190112-5). E. Larsson acknowledges the Swedish Taxonomy Initiative, SLU Artdatabanken, Uppsala (dha.2019.4.3-13). The study of D.B. Raudabaugh and colleagues was supported by the Schmidt Science Fellows, in partnership with the Rhodes Trust. Gregorio Delgado is grateful to Michael Manning and Kamash Pillai (Eurofins EMLab P&K) for provision of laboratory facilities. Jose G. Maciá-Vicente acknowledges support from the German Research Foundation under grant MA7171/1-1, and from the Landes-Offensive zur Entwicklung Wissenschaftlich-ökonomischer Exzellenz (LOEWE) of the state of Hesse within the framework of the Cluster for Integrative Fungal Research (IPF). Thanks are also due to the authorities of the Cabañeros National Park and Los Alcornocales Natural Park for granting the collection permit and for support during field work. The study of Alina V. Alexandrova was carried out as part of the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University No. 121032300081-7. Michał Gorczak was financially supported by the Ministry of Science and Higher Education through the Faculty of Biology, University of Warsaw intramural grant DSM 0117600- 13. M. Gorczak acknowledges M. Klemens for sharing a photo of the Białowieża Forest logging site and M. Senderowicz for help with preparing the illustration. Ivona Kautmanová and D. Szabóová were funded by the Operational Program of Research and Development and co-financed with the European Fund for Regional Development (EFRD). ITMS 26230120004: ‘Building of research and development infrastructure for investigation of genetic biodiversity of organisms and joining IBOL initiative’. Ishika Bera, Aniket Ghosh, Jorinde Nuytinck and Annemieke Verbeken are grateful to the Director, Botanical Survey of India (Kolkata), Head of the Department of Botany & Microbiology & USIC Dept. HNB Garhwal University, Srinagar, Garhwal for providing research facilities. Ishika Bera and Aniket Ghosh acknowledge the staff of the forest department of Arunachal Pradesh for facilitating the macrofungal surveys to the restricted areas. Sergey Volobuev was supported by the Russian Science Foundation (RSF project N 19-77- 00085). Aleksey V. Kachalkin and colleagues were supported by the Russian Science Foundation (grant No. 19-74-10002). The study of Anna M. Glushakova was carried out as part of the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University No. 121040800174-6. Tracey V. Steinrucken and colleagues were supported by AgriFutures Australia (Rural Industries Research and Development Corporation), through funding from the Australian Government Department of Agriculture, Water and the Environment, as part of its Rural Research and Development for Profit program (PRJ-010527). Neven Matočec and colleagues thank the Croatian Science Foundation for their financial support under the project grant HRZZ-IP-2018-01-1736 (ForFungiDNA). Ana Pošta thanks the Croatian Science Foundation for their support under the grant HRZZ-2018-09-7081. The research of Milan Spetik and co-authors was supported by Internal Grant of Mendel University in Brno No. IGAZF/ 2021-SI1003. K.C. Rajeshkumar thanks SERB, the Department of Science and Technology, Government of India for providing financial support under the project CRG/2020/000668 and the Director, Agharkar Research Institute for providing research facilities. Nikhil Ashtekar thanks CSIR-HRDG, INDIA, for financial support under the SRF fellowship (09/670(0090)/2020-EMRI), and acknowledges the support of the DIC Microscopy Facility, established by Dr Karthick Balasubramanian, B&P (Plants) Group, ARI, Pune. The research of Alla Eddine Mahamedi and co-authors was supported by project No. CZ.02.1.01/0.0/0.0/16_017/0002334, Czech Republic. Tereza Tejklová is thanked for providing useful literature. A. Polhorský and colleagues were supported by the Operational Program of Research and Development and co-financed with the European fund for Regional Development (EFRD), ITMS 26230120004: Building of research and development infrastructure for investigation of genetic biodiversity of organisms and joining IBOL initiative. Yu Pei Tan and colleagues thank R. Chen for her technical support. Ernest Lacey thanks the Cooperative Research Centres Projects scheme (CRCPFIVE000119) for its support. Suchada Mongkolsamrit and colleagues were financially supported by the Platform Technology Management Section, National Center for Genetic Engineering and Biotechnology (BIOTEC), Project Grant No. P19-50231. Dilnora Gouliamova and colleagues were supported by a grant from the Bulgarian Science Fund (KP-06-H31/19). The research of Timofey A. Pankratov was supported by the Russian Foundation for Basic Research (grant No. 19-04-00297a). Gabriel Moreno and colleagues wish to express their gratitude to L. Monje and A. Pueblas of the Department of Drawing and Scientific Photography at the University of Alcalá for their help in the digital preparation of the photographs, and to J. Rejos, curator of the AH herbarium, for his assistance with the specimens examined in the present study. Vit Hubka was supported by the Charles University Research Centre program No. 204069. Alena Kubátová was supported by The National Programme on Conservation and Utilization of Microbial Genetic Resources Important for Agriculture (Ministry of Agriculture of the Czech Republic). The Kits van Waveren Foundation (Rijksherbariumfonds Dr E. Kits van Waveren, Leiden, Netherlands) contributed substantially to the costs of sequencing and travelling expenses for M. Noordeloos. The work of B. Dima was supported by the ÚNKP-20-4 New National Excellence Program of the Ministry for Innovation and Technology from the source of the National Research, Development and Innovation Fund, and by the ELTE Thematic Excellence Programme 2020 supported by the National Research, Development and Innovation Office of Hungary (TKP2020-IKA-05). The Norwegian Entoloma studies received funding from the Norwegian Biodiversity Information Centre (NBIC), and the material was partly sequenced through NorBOL. Gunnhild Marthinsen and Katriina Bendiksen (Natural History Museum, University of Oslo, Norway) are acknowledged for performing the main parts of the Entoloma barcoding work. Asunción Morte is grateful to AEI/FEDER, UE (CGL2016-78946-R) and Fundación Séneca - Agencia de Ciencia y Tecnología de la Región de Murcia (20866/PI/18) for financial support. Vladimír Ostrý was supported by the Ministry of Health, Czech Republic - conceptual development of research organization (National Institute of Public Health – NIPH, IN 75010330). Konstanze Bensch (Westerdijk Fungal Biodiversity Institute, Utrecht) is thanked for correcting the spelling of various Latin epithets.

Tree species is one of the most important determinants of wood-inhabiting fungal community composition, yet its relationship with fungal reproductive and dispersal traits remains poorly understood. We studied fungal communities (total of 657 species) inhabiting broadleaved and coniferous dead wood (total of 192 logs) in 12 semi-natural boreal forests. We utilized a trait-based hierarchical joint species distribution model to examine how the relationship between dead wood quality and species occurrence correlates with reproductive and dispersal morphological traits. Broadleaved trees had higher species richness than conifers, due to discomycetoids and pyrenomycetoids specializing in them. Resupinate and pileate species were generally specialized in coniferous dead wood. Fungi inhabiting broadleaved trees had larger and more elongated spores than fungi in conifers. Spore size was larger and spore shape more spherical in species occupying large dead wood units. These results indicate the selective effect of dead wood quality, visible not only in species diversity, but also in reproductive and dispersal traits. (C) 2019 Elsevier Ltd and British Mycological Society. All rights reserved. peerReviewed

The North European species of Elaphomyces section Elaphomyces (Eurotiales, Pezizomycotina) are studied. Three new species, E. citrinopapillatus, E. pusillus, and E. roseoviolaceus are introduced and verified by morphology and sequence data from ITS, nuclear LSU, mitochondrial SSU, and β-tubulin. A lectotype for Elaphomyces granulatus is selected. Elaphomyces granulatus and E. muricatus are epitypified with sequenced material from the Femsjö region in South Sweden. Elaphomyces striatosporus is epitypified with sequenced material from the vicinity of the type locality in Norway. A key to all species of Elaphomyces occurring in Denmark, Norway, and Sweden is provided.

Typhuloid fungi are a very poorly known group of tiny clavarioid homobasidiomycetes. The phylogenetic position and family classification of the genera targeted here, Ceratellopsis, Macrotyphula, Pterula sensu lato and Typhula, are controversial and based on unresolved phylogenies. Our six-gene phylogeny with an expanded taxon sampling shows that typhuloid fungi evolved at least twice in the Agaricales (Pleurotineae, Clavariineae) and once in the Hymenochaetales. Macrotyphula, Pterulicium and Typhula are nested within the Pleurotineae. The type of Typhula (1818) and Sclerotium (1790), T. phacorrhiza and S. complanatum (synonym T. phacorrhiza), are encompassed in the Macrotyphula clade that is distantly related to a monophyletic group formed by species usually assigned to Typhula. Thus, the correct name for Macrotyphula (1972) and Typhula is Sclerotium and all Typhula species but those in the T. phacorrhiza group need to be transferred to Pistillaria (1821). To avoid undesirable nomenclatural changes, we suggest to conserve Typhula with T. incarnata as type. Clavariaceae is supported as a separate, early diverging lineage within Agaricales, with Hygrophoraceae as a successive sister taxon to the rest of the Agaricales. Ceratellopsis s. auct. is polyphyletic because C. acuminata nests in Clavariaceae and C. sagittiformis in the Hymenochaetales. Ceratellopsis is found to be an earlier name for Pterulicium, because the type, C. queletii, represents Pterulicium gracile (synonym Pterula gracilis), deeply nested in the Pterulicium clade. To avoid re-combining a large number of names in Ceratellopsis we suggest to conserve it with C. acuminata as type. The new genus Bryopistillaria is created to include C. sagittiformis. The families Sarcomyxaceae and Phyllotopsidaceae, and the suborder Clavariineae, are described as new. Six new combinations are proposed and 15 names typified.

Morphological and molecular phylogenetic studies of true morels (Morchella) in North America, the Dominican Republic, Venezuela, Ecuador, and Peru led to the discovery of four undescribed species of Morchella. Two new species in the Elata clade, one from the Dominican Republic, initially distinguished by the informal designation Mel-18, and a newly discovered sister species from northern Arizona, are now recognized. Mel-18 is described as a novel phylogenetically distinct species, M. hispaniolensis. Its sister species from Arizona is described as M. kaibabensis, also recovered as an endophyte of Rocky Mountain juniper. Two additional species in the Esculenta clade, M. peruviana discovered in Peru and M. gracilis (previously reported as Mes-14) from the Dominican Republic, Venezuela, and Ecuador, are described as new. We also demonstrate that scanning electron microscopy (SEM) imaging of ascospores using rehydration/dehydration/critical point drying preparation techniques provides for enhanced resolution of spore wall surfaces, thereby increasing the number of morphological traits available to assess differences among otherwise closely related species.

Phylogenetic reconstruction of Hodophilus species with a yellow colour on the stipe based on nrITS, nrLSU and rpb2 sequences revealed six European species. All these species correspond to the widely accepted European concept of a single species Hodophilus micaceus. Four of these species are described and illustrated. H. micaceus and H. phaeoxanthus are recognised as two separate species and H. albofloccipes as a synonym of the latter. Two species, H. anatinus and H. cambriensis, are described as new. Possible endemism of H. micaceus and H. cambriensis to the British Isles is discussed. All analysed North American samples represent different species to those found in Europe. The North American species Hygrophorus rugulosus is combined in the genus Hodophilus. The preliminary key uses position and development of the yellow colour during maturation as the most important distinguishing character. The presence of the yellow colour is discussed as a possible synapomorphic character.

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. 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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. 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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. 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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)

Hodophilus atropunctus is traditionally defined as the only species of this genus with dark brown or black dots on the stipe. Multi-locus phylogenetic reconstruction recognised two distinct clades morphologically corresponding to this species concept. The limited morphological description in the protologue of H. atropunctus and absence of a type specimen were limitations in an assignment of this name to one of the recognised phylogenetic species. The emended species concept and the selection of a neotype are based on careful analyses of the colour of the basidiomata and how this changes during maturation and drying. The name H. atropunctus is assigned to the paler of the two species which also shows colour change across the pileus and along the length of the stipe when dry. The second darker species is described here as new, H. variabilipes, but only seven out of 14 collections examined belonging to this taxon had distinct dark coloured dots on the stipe surface.

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.

In light of the biodiversity crisis and our limited ability to explain variation in biodiversity, tools to quantify spatial and temporal variation in biodiversity and its underlying drivers are critically needed. Inspired by the recently published ecospace framework, we developed and tested a sampling design for environmental and biotic mapping. We selected 130 study sites (40 × 40 m) across Denmark using stratified random sampling along the major environmental gradients underlying biotic variation. Using standardized methods, we collected site species data on vascular plants, bryophytes, macrofungi, lichens, gastropods and arthropods. To evaluate sampling efficiency, we calculated regional coverage (relative to the known species number per taxonomic group), and site scale coverage (i.e., sample completeness per taxonomic group at each site). To extend taxonomic coverage to organisms that are difficult to sample by classical inventories (e.g., nematodes and non-fruiting fungi), we collected soil for metabarcoding. Finally, to assess site conditions, we mapped abiotic conditions, biotic resources and habitat continuity. BACKGROUND: In light of the biodiversity crisis and our limited ability to explain variation in biodiversity, tools to quantify spatial and temporal variation in biodiversity and its underlying drivers are critically needed. Inspired by the recently published ecospace framework, we developed and tested a sampling design for environmental and biotic mapping. We selected 130 study sites (40 × 40 m) across Denmark using stratified random sampling along the major environmental gradients underlying biotic variation. Using standardized methods, we collected site species data on vascular plants, bryophytes, macrofungi, lichens, gastropods and arthropods. To evaluate sampling efficiency, we calculated regional coverage (relative to the known species number per taxonomic group), and site scale coverage (i.e., sample completeness per taxonomic group at each site). To extend taxonomic coverage to organisms that are difficult to sample by classical inventories (e.g., nematodes and non-fruiting fungi), we collected soil for metabarcoding. Finally, to assess site conditions, we mapped abiotic conditions, biotic resources and habitat continuity.RESULTS: Despite the 130 study sites only covering a minute fraction (0.0005%) of the total Danish terrestrial area, we found 1774 species of macrofungi (54% of the Danish fungal species pool), 663 vascular plant species (42%), 254 bryophyte species (41%) and 200 lichen species (19%). For arthropods, we observed 330 spider species (58%), 123 carabid beetle species (37%) and 99 hoverfly species (33%). Overall, sample coverage was remarkably high across taxonomic groups and sufficient to capture substantial spatial variation in biodiversity across Denmark. This inventory is nationally unprecedented in detail and resulted in the discovery of 143 species with no previous record for Denmark. Comparison between plant OTUs detected in soil DNA and observed plant species confirmed the usefulness of carefully curated environmental DNA-data. Correlations among species richness for taxonomic groups were predominantly positive, but did not correlate well among all taxa suggesting differential and complex biotic responses to environmental variation.CONCLUSIONS: We successfully and adequately sampled a wide range of diverse taxa along key environmental gradients across Denmark using an approach that includes multi-taxon biodiversity assessment and ecospace mapping. Our approach is applicable to assessments of biodiversity in other regions and biomes where species are structured along environmental gradient.

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.

Plants regulate soils and microclimate, provide substrate for heterotrophic taxa, are easy to observe and identify and have a stable taxonomy, which strongly justifies the use of plants as bioindicators in monitoring and conservation. However, insects and fungi make up the vast majority of species. Surprisingly, it remains untested whether plants are strong predictors of total multi-taxon species richness. To answer this question, we collected an extensive data set on species richness of vascular plants, bryophytes, macrofungi, lichens, plant-galling arthropods, gastropods, spiders, carabid beetles, hoverflies and OTU richness from environmental DNA metabarcoding. Plant species richness per se was a moderate predictor of richness of other taxa. Taking an ecospace approach to modelling, the addition of plant-derived bioindicators revealed 1) a consistently positive effect of plant richness on other taxa, 2) prediction of 12-55% of variation in other taxa and 48 % of variation in the total species richness.

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

1. Plants regulate soils and microclimate, provide substrate for heterotrophic taxa, are easy to observe and identify and have a stable taxonomy, which strongly justifies their use as indicators in monitoring and conservation. However, there is no consensus as to whether plants are strong predictors of total multi-taxon species richness. In this study we investigate if general terrestrial species richness can be predicted by vascular plant richness and bioindication. 2. To answer this question, we collected an extensive data set on species richness of vascular plants, bryophytes, macrofungi, lichens, plant-galling arthropods, gastropods, spiders, carabid beetles, hoverflies and genetic richness (Operational Taxonomic Units = OTUs) from environmental DNA metabarcoding. We also constructed a Conservation Index based on threatened red list species. Besides using richness of vascular plants for prediction of other taxonomic groups, we also used plant-derived calibration of the abiotic environment (moisture, soil fertility and light conditions) as well as the degree of anthropogenic impact. 3. Bivariate relationships between plant species richness and other species groups showed no consistent pattern. After taking environmental calibration by bioindication into account, we found a consistent, and for most groups significant, positive effect of plant richness. Plant species richness was also important for richness of fungal OTUs, Malaise OTUs and for the Conservation Index. Our multiple regression analyses revealed 1) a consistently positive effect of plant richness on other taxa, 2) prediction of 12-55% of variation in other taxa and 48% of variation in the total species richness when bioindication and plant richness were used as predictors. 4. Our results justify that vascular plants are strong indicators of total biodiversity across environmental gradients and broad taxonomic realms and therefore a natural first choice for biodiversity monitoring and conservation planning. Plants regulate soils and microclimate, provide substrate for heterotrophic taxa, are easy to observe and identify and have a stable taxonomy, which strongly justifies their use as indicators in monitoring and conservation. However, there is no consensus as to whether plants are strong predictors of total multi-taxon species richness. In this study, we investigate if general terrestrial species richness can be predicted by vascular plant richness and bioindication. To answer this question, we collected an extensive dataset on species richness of vascular plants, bryophytes, macrofungi, lichens, plant-galling arthropods, gastropods, spiders, carabid beetles, hoverflies, and genetic richness (operational taxonomic units = OTUs) from environmental DNA metabarcoding. We also constructed a Conservation Index based on threatened red list species. Besides using richness of vascular plants for prediction of other taxonomic groups, we also used plant-derived calibration of the abiotic environment (moisture, soil fertility and light conditions) as well as the degree of anthropogenic impact. Bivariate relationships between plant species richness and other species groups showed no consistent pattern. After taking environmental calibration by bioindication into account, we found a consistent, and for most groups significant, positive effect of plant richness. Plant species richness was also important for richness of fungal OTUs, Malaise OTUs and for the Conservation Index. Our multiple regression analyses revealed (a) a consistently positive effect of plant richness on other taxa, (b) prediction of 12%–55% of variation in other taxa and 48% of variation in the total species richness when bioindication and plant richness were used as predictors. Our results justify that vascular plants are strong indicators of total biodiversity across environmental gradients and broad taxonomic realms and therefore a natural first choice for biodiversity monitoring and conservation planning.

International audience; Documenting succession in forest canopy gaps provides insights into the ecological processes governing the temporal dynamics of species within communities. We analyzed the fruiting patterns of a rare but widely distributed saproxylic macro- mycete, Xylobolus subpileatus, during the ageing of natural canopy gaps in oak forests. In one of the last remaining Quercus ilex L. old-growth forests (on the island of Corsica, western Mediterranean basin), we systematically recorded and conducted molecular analyses of X. subpileatus basidiomes in 80 dated natural canopy gaps representing a 45-year long sequence of residence time of tree logs on the forest floor. Xylobolus subpileatus fruited exclusively on Q. ilex logs. The probability of fruiting of X. subpileatus significantly increases during the process of wood decomposition to reach its maximum in the oldest gaps, approximately 40 years after treefall. In contrast, the abundance and the richness of saprobic and ectomycorrhizal fruitbodies decrease as canopy gaps age. Our results emphasize the high ecological specialization of X. subpileatus. They also highlight the imperative need to conserve the last patches of old-growth Mediterranean forests to secure the persistence of this endangered and functionally unique macromycete whose presence is highly dependent on old wood in advanced stages of decomposition.; L’étude des successions écologiques ayant lieu dans les ouvertures de canopée forestière offre l’opportunité de mieux comprendre les processus écologiques gouvernant la dynamique temporelle des espèces a` l’intérieur des communautés. Nous avons analysé la distribution des fructifications d’une espèce, rare mais largement distribuée, de macromycète saproxilique, Xylobolus subpileatus, au cours du vieillissement de trouées naturelles dans les forêts de chênes. Dans l’une des dernières forêts anciennes de chêne vert (Quercus ilex L.) (île de Corse, ouest du bassin méditerranéen), nous avons échantillonné systématique- ment et analysé moléculairement les basidiomes de X. subpileatus dans 80 trouées datées qui représentent une chronoséquence de 45 ans de temps de résidence au sol des troncs d’arbres. Xylobolus subpileatus ne fructifie que sur les troncs de chêne vert. Sa probabilité de fructification est positivement corrélée a` l’âge des trouées et atteint un maximum dans les plus vieilles trouées (environ 40 ans après la chute de l’arbre). A contrario, l’abondance et la richesse des fructifications de champignons décompo- seurs et ectomycorhiziens décroissent avec le vieillissement de la trouée. Nos résultats soulignent la spécialisation écologique poussée de X. subpileatus et mettent en avant le besoin impératif de préserver les dernières parcelles de forêts anciennes méditerranéennes afin de protéger cette espèce de macromycète fonctionnellement originale dont la présence dépend forte- ment de la complétion du processus naturel de décomposition.

BackgroundIn light of the biodiversity crisis and our limited ability to explain variation in biodiversity, tools to quantify spatial and temporal variation in biodiversity and its underlying drivers are critically needed. Inspired by the recently published ecospace framework, we developed and tested a sampling design for environmental and biotic mapping. We selected 130 study sites (40 × 40 m) across Denmark using stratified random sampling along the major environmental gradients underlying biotic variation. Using standardized methods, we collected site species data on vascular plants, bryophytes, macrofungi, lichens, gastropods and arthropods. To evaluate sampling efficiency, we calculated regional coverage (relative to the known species number per taxonomic group), and site scale coverage (i.e., sample completeness per taxonomic group at each site). To extend taxonomic coverage to organisms that are difficult to sample by classical inventories (e.g., nematodes and non-fruiting fungi), we collected soil for metabarcoding. Finally, to assess site conditions, we mapped abiotic conditions, biotic resources and habitat continuity.ResultsDespite the 130 study sites only covering a minute fraction (0.0005 %) of the total Danish terrestrial area, we found 1774 species of macrofungi (54 % of the Danish fungal species pool), 663 vascular plant species (42 %), 254 bryophyte species (41 %) and 200 lichen species (19 %). For arthropods, we observed 330 spider species (58 %), 123 carabid beetle species (37 %) and 99 hoverfly species (33 %). Correlations among species richness for taxonomic groups were predominantly positive. Overall, sample coverage was remarkably high across taxonomic groups and sufficient to capture substantial spatial variation in biodiversity across Denmark. This inventory is nationally unprecedented in detail and resulted in the discovery of 143 species with no previous record for Denmark. Comparison between plant OTUs detected in soil DNA and observed plant species confirmed the usefulness of carefully curated environmental DNA-data. Species richness did not correlate well among taxa suggesting differential and complex biotic responses to environmental variation.ConclusionsWe successfully and adequately sampled a wide range of diverse taxa along key environmental gradients across Denmark using an approach that includes multi-taxon biodiversity assessment and ecospace mapping. Our approach is applicable to assessments of biodiversity in other regions and biomes where species are structured along environmental gradient.

Elaphomyces ('deer truffles') is one of the most important ectomycorrhizal fungal genera in temperate and subarctic forest ecosystems, but also one of the least documented in public databases. The current systematics are mainly based on macromorphology, and is not significantly different from that proposed by Vittadini (1831). Within the 49 species recognised worldwide, 23 were originally described from Europe and 17 of these were described before the 20th century. Moreover, very recent phylogenetic treatments of the genus are mainly based on a few extra-European species and most common European species are still poorly documented. Based on an extensive taxonomic sampling mainly made in the biogeographically rich Cantabrian area (Spain), complemented with collections from France, Greece, Italy, Norway, Portugal and Sweden, all currently recognized species in Europe have been sequenced at the ITS and 28S of the rDNA. Combined phylogenetic analyses yielded molecular support to sections Elaphomyces and Ceratogaster (here emended), while a third, basal lineage encompasses the sections Malacodermei and Ascoscleroderma as well as the tropical genus Pseudotulostoma. Species limits are discussed and some taxa formerly proposed as genuine species based on morphology and biogeography are re-evaluated as varieties or forms. Spore size and ornamentation, features of the peridial surface, structure of the peridium, and the presence of mycelium patches attached to the peridial surface emerge as the most significant systematic characters. Four new species: E. barrioi, E. quercicola, E. roseolus and E. violaceoniger, one new variety: E. papillatus var. sulphureopallidus, and two new forms: E. granulatus forma pallidosporus and E. anthracinus forma talosporus are introduced, as well as four new combinations in the genus: E. muricatus var. reticulatus, E. muricatus var. variegatus, E. papillatus var. striatosporus and E. morettii var. cantabricus. Lectotypes and epitypes are designated for most recognised species. For systematic purposes, new infrageneric taxa are introduced: E. sect. Ascoscleroderma stat. nov., E. subsect. Sclerodermei stat. nov., E. subsect. Maculati subsect. nov., E. subsect. Muricati subsect. nov., and E. subsect. Papillati subsect. nov. Lastly, E. laevigatus, E. sapidus, E. sulphureopallidus and E. trappei are excluded from the genus and referred to Rhizopogon roseolus, Astraeus sapidus comb. nov., Astraeus hygrometricus and Terfezia trappei comb. nov. (syn.: Terfezia cistophila), respectively.

Biodiversity is declining on a global scale and the limited resources available for conservation efforts and research need to be used efficiently. Fungi are a megadiverse taxonomic group where lack of knowledge impedes sufficient conservation focus. Inventories of fungi require a high level of expertise and are difficult and expensive due to seasonal variation and fluctuation in sporocarp formation. Lidar offers objective, fine-resolution and potentially cheap and broad scale data for representing vegetation structure and has proven useful for prediction of the diversity of plant and animal species. In this study, we used lidar-derived variables as explanatory variables in models of fungal species richness and controlling gradients of species composition, represented by three ordination axes derived from multidimensional scaling. The fungal data comprised a data set of 1527 species recorded in three inventories of 121 sites covering all major terrestrial habitat types across Denmark. We compared the performance of lidar-derived explanatory variables with two field inventory-based sets of variables: 1) lists of vascular plants and 2) recordings of soil, microclimate and vegetation structure. Lidar-derived variables performed best in predicting fungal richness and the first ordination axis, interpreted as a gradient from low and open herb-dominated communities over closed swards and scrubs to closed-canopy forests (cross validated r 2 of 0.50 and 0.81, respectively). The number of red-listed species and the second ordination axis were poorly predicted by lidar, and the third ordination axis was equally well detected by the three explanatory sets of variables (cross validated r 2 of 0.38). From a fungal conservation perspective, it is promising that lidar-based variables hold information suitable for detecting major gradients in fungal richness and composition.

The new species Chlorostroma vestlandicum is described from coarse dead wood of Ulmus glabra in western Norway. It was invariably found in close association with Hypoxylon vogesiacum and appears to be mycoparasitic on this species. With a strikingly orange entostroma, tiny perithecia and specialized habitat association it is a highly distinctive species. C. vestlandicum differs from the type species by the color of the entostroma (bright yellow orange as opposed to ochraceous), iodine reaction of the apical apparatus, ascospores (more or less ellipsoid as opposed to more or less cuboid). The surface is not green or bluegreen as in the previously described species, albeit dark greenish blackish in section. Its distribution seems to cover mainly the hemiboreal regions of western Norway, an area not yet affected but threatened by Dutch elm disease. It is probably a rare species restricted to the most dead wood rich sites with big populations of H. vogesiacum.

For a monograph based on a polythetic concept, several thousands of herbarium specimens, and several hundreds of freshly collected and cultured specimens of Daldinia and allied Xylariaceae, originating from around the world, were studied for morphological traits, including by SEM, and chemically by HPLC profiles using UV-visible and mass spectrometric detection. Emphasis was given to tropical material, and importantly, ancient specimens, including as many types as possible, were tracked and studied to review earlier taxonomic concepts. An epitype of D. eschscholtzii was selected as representative of the morphochemotype that is most widely distributed in the tropics. Six new species of Daldinia from the tropics and the southern Hemisphere are described. Daldinia asphalatum is resurrected, and D. cudonia is regarded as its synonym. In addition, the following binomials are epi-, iso-, neo- and/or lectotypified: Daldinia asphalatum, D. caldariorum, D. clavata, D. cuprea, D. durissima, D. eschscholtzii, D. grandis, D. loculata, and D. vernicosa. Annellosporium and Versiomyces are regarded as synonyms of Daldinia. Many new synonymies in Daldinia are proposed, and some previously published names are rejected. In total, 47 taxa in Daldinia are recognised and a key is provided. Their biogeography, chorology, and ecology, as well as the importance of their secondary metabolites, are also discussed. The previous definition of the genus is emended. The species concept is based mainly on morphological and other phenotype-derived characters because, despite diligent search, no molecular data or cultures of several of the accepted species could be obtained. Daldinia is segregated into five major groups, based on phenotypic characteristics. Some unnamed but aberrant specimens were not found in good condition and are therefore not formally described as new species. However, they are illustrated in detail in a hope that this will facilitate the discovery of fresh material in future. A preliminary molecular phylogeny based on 5.8S/ITS nrDNA including numerous representatives of all hitherto described taxa for which cultures are extant, was found basically in agreement with the above mentioned segregation of the genus, based on morphological and chemotaxonomic evidence. In the rDNA based phylogenetic tree, Daldinia appears clearly distinct from members of the genera Annulohypoxylon and Hypoxylon; nevertheless, representatives of small genera of predominantly tropical origin (Entonaema, Phylacia, Ruwenzoria, Rhopalostroma, Thamnomyces) appear to have evolved from daldinioid ancestors and are nested inside the Daldinia clade. Interestingly, these findings correlate with chemotaxonomic characters to a great extent, especially regarding the distribution of marker metabolites in their mycelial cultures. Hence, the current study revealed for the first time that fungal secondary metabolite profiles can have taxonomic value beyond the species rank and even coincide with phylogenetic data.Taxonomic novelties: Daldinia andina sp. nov., D. australis sp. nov., D. hausknechtii sp. nov., D. rehmii sp. nov., D. starbaeckii sp. nov., D. theissenii sp. nov., D. cahuchosa comb. nov., D. nemorosa comb. nov.

Phylogenetic analyses of Mycena sect. Calodontes using ITS previously suggested ten cryptic monophyletic ITS lineages within the Mycena pura morphospecies. Here, we compare ITS data (645 bp incl. gaps) from 46 different fruit bodies that represent the previously described ITS diversity with partial tEF-1-α (423 bp) and RNA polymerase II (RPB1) (492 bp) sequence data to test the genealogical concordance. While neither of the markers were in complete topological agreement, the branches differing between the tEF and RPB1 trees had a low bootstrap (

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

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

In Mycena sectio Calodontes with otherwise amyloid spores, the inamyloid spores of Mycena pearsoniana Dennis ex Singer were a distinguishing feature for this species and its subsection Violacella. Although the original concept of this species was European, Singer chose to typify it with material collected in Mexico. The name has since been applied to all European collections with inamyloid spores and decurrent lamellae. Our phylogenetic analysis of 91 ITS sequences from European, North and South American Calodontes collections shows that European collections identified as M. pearsoniana fall into two well-supported sibling clades together with both inamyloid and weakly amyloid North American collections. Since the holotype of M. pearsoniana is in an advanced state of decay, we have selected an epitype from a North American locality with a climate comparable to the Mexican type locality. Our results show weakly and inamyloid spore reactions to be homoplastic in Calodontes, and furthermore that spores of M. pearsoniana can show either amyloid or inamyloid reactions interchangeably. This raises doubt about the taxonomic value of this trait in Mycena systematics.

The tropical genus Thamnomyces is characterized by having wiry, black, brittle stromata and early deliquescent asci, lacking an amyloid apical apparatus. Thamnomyces is regarded as a member of the Xylariaceae because the morphology of its ascospores and the anamorphic structures are typical for this family. However, its relationship to other xylariaceous genera remained to be clarified. Cultures of three Thamnomyces species were obtained and studied for morphological characters, and their secondary metabolite profiles as inferred from high performance liquid chromatography coupled with mass spectrometric and diode array detection (HPLC–MS/DAD) were also compared. Cultures of Thamnomyces closely resembled those of the genera Daldinia and Phylacia and even produced several secondary metabolite families that are known to be chemotaxonomic markers for the aforementioned genera. These findings were corroborated by a comparison of their 5.8S/ITS nrDNA sequences. We conclude that Thamnomyces, Daldinia, and Phylacia are derived from the same evolutionary lineage, despite these genera differing drastically in their stromatal morphology and anatomy. Along with Entonaema and Rhopalostoma, these fungi comprise an evolutionarily derived lineage of the hypoxyloid Xylariaceae. A new species of Thamnomyces is erected, and preliminary descriptions of three further, potentially new taxa are also provided.

The members of Mycena sect. Calodontes (Tricholomataceae s.l., Basidiomycota) are characterised by a collybioid aspect and more or less purplish to reddish colours and a distinct raphanoid odour. In Europe, nine species have been recognised though some of these based on somewhat dubious morphological differences. Historically, most were assigned to Mycena pura. However, since Mycena pura displays one of the most striking colour variabilities within European agarics, many attempts have been made to subdivide it into independent entities, and several forms, varieties and species have been split from Mycena pura s.l. based largely on differences in colouration, gross macromorphology or other phenetic traits. We developed a large sample of ITS sequences of all species of sect. Calodontes known from Europe for which vouchers exist. Furthermore, partial LSU data were developed and additional sequences downloaded from GENBANK to assess the relationship of Calodontes with other Mycena spp. We show that most Calodontes form a monophyletic group including a few North and South American collections, but that this cannot be conclusively shown when an additional North American sequence is added. For all other species than M. pura and M. diosma, we found morphological species recognition to be in agreement with the ITS data. Several significantly different clades can be recognised within the M. pura morphospecies, none of which can be linked to the observed (and described by proxy) colour varieties/forms. Indications of a possible environmental basis of the colour differentiation in the M. pura morphospecies are discussed.

During a foray to the mountain rainforests of the Democratic Republic of the Congo, a peculiar species of Xylariaceae was found, which could not be accommodated in any of the existing genera. It is recognised as representative of a new genus, named Ruwenzoria, owing to the presence of a new combination of teleomorphic and anamorphic characters that are regarded as significant for generic segregation within the Xylariaceae. Studies on its secondary metabolites in stromata and cultures by high performance liquid chromatography, coupled with diode array and mass spectrometric detection (HPLC-DAD/MS), and on its phylogenetic affinities based on 5.8S/ITS rDNA sequence data, respectively, revealed a close relationship of the new taxon to the genera Daldinia and Entonaema, from which it differs by having neither a hollow, gelatinous nor a conspicuously zonate stromatal interior, and an anamorph featuring enteroblastic rather than holoblastic conidiogenesis. A specimen from the same geographic region, previously identified as Daldinia bakeri by R.W.G. Dennis was found to constitute a mixture of stromata of Ruwenzoria and an additional, undescribed Daldinia species. The latter fungus is not formally described due to the scantiness of the material, but its morphological characteristics are illustrated.

Two new species of Camillea are described from localities on an east–west transect through Ecuador between the latitudes 1°00’N and 1°00’S. Camillea ovalispora was collected in San Vicente de Huaticocha, an area characterized as humid lowland rainforest and premontane cloud forest; C. unistoma was collected at Cuyabeno, a protected area with black water inundated forest and primary rainforest on terra firme. Further, C. heterostoma var. macrospora is elevated to full species status as C. macrospora. New country records of six Camillea species are annotated from the same east-west transect. These are C. flosculosa, C. fossulata, C. hainesii, C. macrospora, C. scriblita, and C. tinctor. Also, previously recorded species have been annotated with new specimens added. Additionally, six distinct taxa from the transect that appear to be undescribed were included in the analysis; one is partly described but not the others due to insufficient material. The data showed an obvious altitude influence on the distribution and frequency of Camillea species. The diversity of species is lower on the western part of the transect compared to the eastern side, probably due to multiple factors such as differences in search intensity between the two slopes, difference in annual rainfall, and possibly the availability of host plants. Camillea taxa were only found at altitudes below 2,000 m, with the highest diversity below 600 m. Palmae is reported as a new host for Camillea species.

Fungus-growing termites live in obligate mutualistic symbiosis with species of the basidiomycete genus Termitomyces, which are cultivated on a substrate of dead plant material. When the termite colony dies, or when nest material is incubated without termites in the laboratory, fruiting bodies of the ascomycete genus Xylaria appear and rapidly cover the fungus garden. This raises the question whether certain Xylaria species are specialised in occupying termite nests or whether they are just occasional visitors. We tested Xylaria specificity at four levels: (1) fungus-growing termites, (2) termite genera, (3) termite species, and (4) colonies. In South Africa, 108 colonies of eight termite species from three termite genera were sampled for Xylaria. Xylaria was isolated from 69% of the sampled nests and from 57% of the incubated fungus comb samples, confirming high prevalence. Phylogenetic analysis of the ITS region revealed 16 operational taxonomic units of Xylaria, indicating high levels of Xylaria species richness. Not much of this variation was explained by termite genus, species, or colony; thus, at level 2-4 the specificity is low. Analysis of the large subunit rDNA region, showed that all termite-associated Xylaria belong to a single clade, together with only three of the 26 non-termite-associated strains. Termite-associated Xylaria thus show specificity for fungus-growing termites (level 1). We did not find evidence for geographic or temporal structuring in these Xylaria phylogenies. Based on our results, we conclude that termite-associated Xylaria are specific for fungus-growing termites, without having specificity for lower taxonomic levels.

A study of the diversity, taxonomy, and ecology of endophytic Xylariaceae (Ascomycota) was carried out. In this study, we obtained isolates of Xylariaceae from healthy, attached leaves and teleomorphic stromata on decayed plant materials in a permanent plot at Khao Yai National Park (Thailand). In addition, strains deposited beforehand were selected in which both endophytic strains isolated from living plant tissues and saprobic strains from fruit bodies were included. Consequently, 405 strains of Xylariaceae (273 endophytic and 132 saprobic strains, including identified strains) were studied to reveal the diversity and taxonomy of endophytes and the relationships between those endophytes and saprobic Xylariaceae in Thailand that have been recorded according to fruit-body formation on decayed plant materials. Analysis of 28S rDNA D1/D2 sequences revealed 21 xylariaceous species inhabiting tropical foliage at the site, and several species that are already known as saprobes appear to be among those isolated from living leaves. Furthermore, several clades that consisted of only endophytic strains were found, and some of these have no known matches in public DNA sequence banks.