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Mushroom-forming fungi (Agaricomycetes) have the greatest morphological diversity and complexity of any group of fungi. They have radiated into most niches and fulfil diverse roles in the ecosystem, including wood decomposers, pathogens or mycorrhizal mutualists. Despite the importance of mushroom-forming fungi, large-scale patterns of their evolutionary history are poorly known, in part due to the lack of a comprehensive and dated molecular phylogeny. Here, using multigene and genome-based data, we assemble a 5,284-species phylogenetic tree and infer ages and broad patterns of speciation/extinction and morphological innovation in mushroom-forming fungi. Agaricomycetes started a rapid class-wide radiation in the Jurassic, coinciding with the spread of (sub)tropical coniferous forests and a warming climate. A possible mass extinction, several clade-specific adaptive radiations and morphological diversification of fruiting bodies followed during the Cretaceous and the Paleogene, convergently giving rise to the classic toadstool morphology, with a cap, stalk and gills (pileate-stipitate morphology). This morphology is associated with increased rates of lineage diversification, suggesting it represents a key innovation in the evolution of mushroom-forming fungi. The increase in mushroom diversity started during the Mesozoic-Cenozoic radiation event, an era of humid climate when terrestrial communities dominated by gymnosperms and reptiles were also expanding. A phylogenetic tree of 5,284 fungal species is used to infer patterns of extinction, diversification and morphological innovation in mushroom-forming fungi.

This paper is a compilation of notes on 142 fungal taxa, including five new families, 20 new genera, and 100 new species, representing a wide taxonomic and geographic range. The new families, Ascocylindricaceae, Caryosporaceae and Wicklowiaceae (Ascomycota) are introduced based on their distinct lineages and unique morphology. The new Dothideomycete genera Pseudomassariosphaeria (Amniculicolaceae), Heracleicola, Neodidymella and Pseudomicrosphaeriopsis (Didymellaceae), Pseudopithomyces (Didymosphaeriaceae), Brunneoclavispora, Neolophiostoma and Sulcosporium (Halotthiaceae), Lophiohelichrysum (Lophiostomataceae), Galliicola, Populocrescentia and Vagicola (Phaeosphaeriaceae), Ascocylindrica (Ascocylindricaceae), Elongatopedicellata (Roussoellaceae), Pseudoasteromassaria (Latoruaceae) and Pseudomonodictys (Macrodiplodiopsidaceae) are introduced. The newly described species of Dothideomycetes (Ascomycota) are Pseudomassariosphaeria bromicola (Amniculicolaceae), Flammeascoma lignicola (Anteagloniaceae), Ascocylindrica marina (Ascocylindricaceae), Lembosia xyliae (Asterinaceae), Diplodia crataegicola and Diplodia galiicola (Botryosphaeriaceae), Caryospora aquatica (Caryosporaceae), Heracleicola premilcurensis and Neodidymella thailandicum (Didymellaceae), Pseudopithomyces palmicola (Didymosphaeriaceae), Floricola viticola (Floricolaceae), Brunneoclavispora bambusae, Neolophiostoma pigmentatum and Sulcosporium thailandica (Halotthiaceae), Pseudoasteromassaria fagi (Latoruaceae), Keissleriella dactylidicola (Lentitheciaceae), Lophiohelichrysum helichrysi (Lophiostomataceae), Aquasubmersa japonica (Lophiotremataceae), Pseudomonodictys tectonae (Macrodiplodiopsidaceae), Microthyrium buxicola and Tumidispora shoreae (Microthyriaceae), Alloleptosphaeria clematidis, Allophaeosphaeria cytisi, Allophaeosphaeria subcylindrospora, Dematiopleospora luzulae, Entodesmium artemisiae, Galiicola pseudophaeosphaeria, Loratospora luzulae, Nodulosphaeria senecionis, Ophiosphaerella aquaticus, Populocrescentia forlicesenensis and Vagicola vagans (Phaeosphaeriaceae), Elongatopedicellata lignicola, Roussoella magnatum and Roussoella angustior (Roussoellaceae) and Shrungabeeja longiappendiculata (Tetraploasphaeriaceae). The new combinations Pseudomassariosphaeria grandispora, Austropleospora archidendri, Pseudopithomyces chartarum, Pseudopithomyces maydicus, Pseudopithomyces sacchari, Vagicola vagans, Punctulariopsis cremeoalbida and Punctulariopsis efibulata Dothideomycetes. The new genera Dictyosporella (Annulatascaceae), and Tinhaudeus (Halosphaeriaceae) are introduced in Sordariomycetes (Ascomycota) while Dictyosporella aquatica (Annulatascaceae), Chaetosphaeria rivularia (Chaetosphaeriaceae), Beauveria gryllotalpidicola and Beauveria loeiensis (Cordycipitaceae), Seimatosporium sorbi and Seimatosporium pseudorosarum (Discosiaceae), Colletotrichum aciculare, Colletotrichum fusiforme and Colletotrichum hymenocallidicola (Glomerellaceae), Tinhaudeus formosanus (Halosphaeriaceae), Pestalotiopsis subshorea and Pestalotiopsis dracaenea (Pestalotiopsiceae), Phaeoacremonium tectonae (Togniniaceae), Cytospora parasitica and Cytospora tanaitica (Valsaceae), Annulohypoxylon palmicola, Biscogniauxia effusae and Nemania fusoideis (Xylariaceae) are introduced as novel species to order Sordariomycetes. The newly described species of Eurotiomycetes are Mycocalicium hyaloparvicellulum (Mycocaliciaceae). Acarospora septentrionalis and Acarospora castaneocarpa (Acarosporaceae), Chapsa multicarpa and Fissurina carassensis (Graphidaceae), Sticta fuscotomentosa and Sticta subfilicinella (Lobariaceae) are newly introduced in class Lecanoromycetes. In class Pezizomycetes, Helvella pseudolacunosa and Helvella rugosa (Helvellaceae) are introduced as new species. The new families, Dendrominiaceae and Neoantrodiellaceae (Basidiomycota) are introduced together with a new genus Neoantrodiella (Neoantrodiellaceae), here based on both morphology coupled with molecular data. In the class Agaricomycetes, Agaricus pseudolangei, Agaricus haematinus, Agaricus atrodiscus and Agaricus exilissimus (Agaricaceae), Amanita melleialba, Amanita pseudosychnopyramis and Amanita subparvipantherina (Amanitaceae), Entoloma calabrum, Cora barbulata, Dictyonema gomezianum and Inocybe granulosa (Inocybaceae), Xerocomellus sarnarii (Boletaceae), Cantharellus eucalyptorum, Cantharellus nigrescens, Cantharellus tricolor and Cantharellus variabilicolor (Cantharellaceae), Cortinarius alboamarescens, Cortinarius brunneoalbus, Cortinarius ochroamarus, Cortinarius putorius and Cortinarius seidlii (Cortinariaceae), Hymenochaete micropora and Hymenochaete subporioides (Hymenochaetaceae), Xylodon ramicida (Schizoporaceae), Colospora andalasii (Polyporaceae), Russula guangxiensis and Russula hakkae (Russulaceae), Tremella dirinariae, Tremella graphidis and Tremella pyrenulae (Tremellaceae) are introduced. Four new combinations Neoantrodiella gypsea, Neoantrodiella thujae (Neoantrodiellaceae), Punctulariopsis cremeoalbida, Punctulariopsis efibulata (Punctulariaceae) are also introduced here for the division Basidiomycota. Furthermore Absidia caatinguensis, Absidia koreana and Gongronella koreana (Cunninghamellaceae), Mortierella pisiformis and Mortierella formosana (Mortierellaceae) are newly introduced in the Zygomycota, while Neocallimastix cameroonii and Piromyces irregularis (Neocallimastigaceae) are introduced in the Neocallimastigomycota. Reference specimens or changes in classification and notes are provided for Alternaria ethzedia, Cucurbitaria ephedricola, Austropleospora, Austropleospora archidendri, Byssosphaeria rhodomphala, Lophiostoma caulium, Pseudopithomyces maydicus, Massariosphaeria, Neomassariosphaeria and Pestalotiopsis montellica.

Research and publication of the planet's remaining plant and fungal species as yet unknown to science is essential if we are to address the United Nations Sustainable Development Goal (SDG) 15 “Life on Land” which includes the protection of terrestrial ecosystems and halting of biodiversity loss. If species are not known to science, they cannot be assessed on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species and so the possibility to protect them from extinction is reduced. Furthermore, until species are known to science they cannot be fully scientifically evaluated for their potential as new foods, medicines, and products which would help address SDGs 1,2,3, and 8. Societal Impact Statement Research and publication of the planet's remaining plant and fungal species as yet unknown to science is essential if we are to address the United Nations Sustainable Development Goal (SDG) 15 “Life on Land” which includes the protection of terrestrial ecosystems and halting of biodiversity loss. If species are not known to science, they cannot be assessed on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species and so the possibility to protect them from extinction is reduced. Furthermore, until species are known to science they cannot be fully scientifically evaluated for their potential as new foods, medicines, and products which would help address SDGs 1,2,3, and 8. Summary Scientific discovery, including naming new taxa, is important because without a scientific name, a species is invisible to science and the possibilities of researching its ecology, applications and threats, and conserving it, are greatly reduced. We review new scientific discoveries in the plant and fungal kingdoms, based largely on new names of taxa published in 2019 and indexed in the International Plant Names Index and Index Fungorum. Numbers of new species in both kingdoms were similar with 1942 new species of plant published and 1882 species of fungi. However, while >50% of plant species have likely been discovered, >90% of fungi remain unknown. This gulf likely explains the greater number of higher order taxa for fungi published in 2019: three classes, 18 orders, 48 families and 214 genera versus one new family and 87 new genera for plants. We compare the kingdoms in terms of rates of scientific discovery, globally and in different taxonomic groups and geographic areas, and with regard to the use of DNA in discovery. We review species new to science, especially those of interest to humanity as new products, and also by life‐form. We consider where future such discoveries can be expected. We recommend an urgent increase in investment in scientific discovery of plant and fungal species, while they still survive. Priorities include more investment in training taxonomists, in building and equipping collections‐based research centers for them, especially in species‐rich, income‐poor countries where the bulk of species as yet unknown to science are thought to occur.

Summary Plant pathogenic fungi are a large and diverse assemblage of eukaryotes with substantial impacts on natural ecosystems and human endeavours. These taxa often have complex and poorly understood life cycles, lack observable, discriminatory morphological characters, and may not be amenable to in vitro culturing. As a result, species identification is frequently difficult. Molecular (DNA sequence) data have emerged as crucial information for the taxonomic identification of plant pathogenic fungi, with the nuclear ribosomal internal transcribed spacer (ITS) region being the most popular marker. However, international nucleotide sequence databases are accumulating numerous sequences of compromised or low-resolution taxonomic annotations and substandard technical quality, making their use in the molecular identification of plant pathogenic fungi problematic. Here we report on a concerted effort to identify high-quality reference sequences for various plant pathogenic fungi and to re-annotate incorrectly or insufficiently annotated public ITS sequences from these fungal lineages. A third objective was to enrich the sequences with geographical and ecological metadata. The results – a total of 31,954 changes – are incorporated in and made available through the UNITE database for molecular identification of fungi ( http://unite.ut.ee ), including standalone FASTA files of sequence data for local BLAST searches, use in the next-generation sequencing analysis platforms QIIME and mothur, and related applications. The present initiative is just a beginning to cover the wide spectrum of plant pathogenic fungi, and we invite all researchers with pertinent expertise to join the annotation effort.

Cortinarius sect. Armillati (subgenus Telamonia) was studied extensively based on morphology and molecular data. A total of about 1000 specimens, mostly from Fennoscandia, were revised. The nomenclature of the species was confirmed by sequencing the type material. Phylogenetic relationships were inferred by analyses of ITS, and the results were compared with the morphological and ecological data. Based on macro- and micromorphological characters, as well as molecular data, section Armillati contains only the medium to large species with slightly hygrophanous pileus and ± reddish or in some species yellowish brown to rose brown universal veils. The other red-brown-veiled species, previously included in Armillati, seem to belong to at least seven different sections or clades: sect. Anthracini, sect. Boulderenses, sect. Brunneotincti p.p., sect. Cinnabarini, sect. Fulvescentes, /Fuscoperonatus, and /Praestigiosus. Our study recognized six Armillati species from northern Europe: C. armillatus, C. luteo-ornatus, C. paragaudis, and three species described as new, C. pinigaudis, C. roseoarmillatus, and C. suboenochelis. The former three also occur in North America. Two additional species, C. subarmillatus (Japan) and C. quercoarmillatus (Costa Rica), are known outside the area. Based on the phylogenetic analysis, the species associated with deciduous trees, C. armillatus, C. quercoarmillatus, and C. roseoarmillatus, all with dextrinoid, thick-walled spores, formed a separate group from the mainly conifer-associated species, C. luteo-ornatus, C. paragaudis, C. pinigaudis and C. suboenochelis, all with fairly thin to moderately thick-walled, indextrinoid to moderately dextrinoid spores. Descriptions of the northern European species are provided, the distribution is mapped and their taxonomy, ecology, distribution, and relationships are discussed. A total of 64 new sequences of 12 species are reported including 17 sequences from type material. Our study also suggests that ITS sequences are not always sufficiently variable for species-rank recognition (barcoding) in Cortinarius.

This paper is a contribution to the current knowledge of taxonomy, ecology and distribution of South American Cortinarius (Pers.) Gray. Cortinarius is among the most widely distributed and species-rich basidiomycete genera occurring with South American Nothofagaceae and species are found in many distinct habitats, including shrublands and forests. Due to their ectomycorrhizal role, Cortinarius species are critical for nutrient cycling in forests, especially at higher latitudes. Some species have also been reported as edible fungi with high nutritional quality. Our aim is to unravel the taxonomy of selected Cortinarius belonging to phlegmacioid and myxotelamonioid species based on morphological and molecular data. After widely sampling Cortinarius specimens in Patagonian Nothofagaceae forests and comparing them to reference collections (including holotypes), we propose five new species of Cortinarius in this work. Phylogenetic analyses of concatenated rDNA ITS-LSU and RPB1 sequences failed to place these new species into known Cortinarius sections or lineages. These findings highlight our knowledge gaps regarding the fungal diversity of South American Nothofagaceae forests. Due to the high diversity of endemic Patagonian taxa, it is clear that the South American Cortinarius diversity needs to be discovered and described in order to understand the evolutionary history of Cortinarius on a global scale.

The North American species of Cortinarius section Sanguinei were studied using morphological characters and ITS and RPB2 sequence data. Several type collections also were examined. Four species were identified: C. harrisonii sp. nov, C. neosanguineus sp. nov., C. sanguineus and C. sierraensis comb. nov. Of these, C. sanguineus also occurs in Europe together with C. puniceus, a fifth member of the section. Typical features of these species include ± red, fairly small basidiomata, stipe basal mycelium often with yellow to reddish yellow tints, amygdaloid to ellipsoid spores, and aniline-red lamellar trama and pileipellis hyphae when mounted in KOH. Two other species with red lamellae C. marylandensis comb. nov. and C. smithii stat. nov. & nom. nov. also are discussed.

In this study, eight species of Chroogomphus are recognized from Europe: C. britannicus , C. aff. filiformis 1, C. fulmineus , C. cf. helveticus , C. mediterraneus , C . cf. purpurascens , C. rutilus , and C. subfulmineus . Different candidates for the application of the name C. rutilus are evaluated and the best fit to the description is selected; lecto- and epitypes are chosen to fix the name. Chroogomphus fulmineus and C. mediterraneus are also epitypified and a new species, C. subfulmineus , is described. The infrageneric classification is revised and a new subgenus Siccigomphus and three new sections, Confusi, Filiformes , and Fulminei are introduced. The former sections Chroogomphus and Floccigomphus are elevated to subgeneric level. Comparison of the ITS regions (nuc rDNA ITS1-5.8S-ITS2) of all species studied shows that there is a minimum interspecific difference of 1.5 %, with the exception of the two species belonging to sect. Fulminei which differ by a minimum of 0.9 %. Ecological specimen data indicate that species of Chroogomphus form basidiomes under members of Pinaceae , with a general preference for species of Pinus . Five European species have been recorded under Picea , while Abies and Larix have also been recorded as tree associates, although the detailed nutritional relationships of the genus, involving other suilloid fungi in particular, have yet to be fully clarified.