Towards a better understanding of the systematics and diversity of Cortinarius, with an emphasis on species growing in boreal and temperate zones of Europe and North America

Towards a better understanding of the systematics and diversity of Cortinarius, with an emphasis on species growing in boreal and temperate zones of Europe and North America Cortinarius is the largest genus of Agaricales with a global distribution and thousands of species. Cortinarius species are important ectomycorrhizal fungi associated with different trees. At the moment, any estimation of the true diversity of Cortinarius is impossible to determine. Species level taxonomy is the foundation of biological studies, without knowing the species and their boundaries it is very difficult to do ecological, applied or other research. Our aims as taxonomists are to discover and describe all species, define relationships, and provide tools for unambiguous identification. It is clear that with morphology we are not able to achieve that goal. The development and use of molecular methods in taxonomic studies of fungi has revolutionized our field of science. Molecular data is more objective and has a better repeatability than morphological data. This thesis project has been the biggest effort to stabilize the nomenclature of Cortinarius so far and also is an example of using only reliable names in taxonomic studies by excluding names without sequences from type material. Also, our view on the amount of diversity has changed dramatically, e.g. estimation of diversity of Cortinarius in Finland is now about same than it was previously estimated to be in Europe. Our studies have revealed much needed information about species composition and have allowed for comparisons between North-America and Europe. New information about potential cryptic species in Cortinarius and limitations of ITS for species level taxonomy has been gained. The sequences of type specimens published create thus far the largest, reliable ITS identification database for Cortinarius containing over 200 species. Our results also confirm the finding of earlier molecular studies that the traditional infra-generic groupings are at least partly artificial and should be reevaluated.Kohti luotettavampaa seitikkien tunnistamista ja nimeämistä DNA-sekvenssien avulla. Sienten luokittelua on tutkittu morfologiaan eli silmillä nähtäviin ominaisuuksiin perustuen jo yli kaksisataa vuotta. Viimeisen vuosikymmenen aikana sienten tutkimuksessa on pystytty käyttämään myös geneettistä tietoa, joka on mullistanut käsityksiämme. Silti tietämyksemme sienten todellisesta monimuotoisuudesta ja nimistön oikeellisuudesta on vielä hyvin puutteellista. Tietoa tarvitaan, jotta sekä sienten perustutkimus että myös tiedon soveltaminen esimerkiksi luonnonsuojelullisissa päätöksissä olisi mielekästä. Seitikit (Cortinarius) on helttasienten lajirikkain suku, jonka edustajia löytyy kaikkialta. Seitikit elävät symbioosissa monien puulajien kanssa. Esimerkiksi Suomessa kaikki taloudellisesti tärkeät puulajimme tarvitsevat seitikkejä kasvaakseen. Maailmanlaajuisesti seitikkejä on kuvattu jo yli 4000 lajia. Silti seitikit ovat vielä hyvin puutteellisesti tunnettuja ja monia yleisiäkään lajeja ei ole vielä tieteelle kuvattu. Suomen tunnettujen seitikkilajien lukumäärä on kaksinkertaistunut viimeisen kymmenen vuoden aikana ja todellinen lajimäärä on vähintään 500 tietämillä. Tällaisen suuren lajimäärän nopea ja luotettava tunnistaminen onnistuu vain DNA-sekvenssien avulla. DNA-tunnistukseen tarvitaan kuitenkin DNA-tietokanta, joka sisältää luotettavasti nimetyt ja luokitellut DNA-sekvenssit mahdollisimman monesta sienilajista. Tämän tutkimuksen myötä saatiin toistaiseksi suurin helttasienten luotettavaan tunnistukseen tarkoitettu DNA-tietokanta luoduksi. Tietokanta sisältää yli 200 seitikkilajia Euroopasta ja Pohjois-Amerikasta

Telamonioid Cortinarius species of the C-puellaris group from calcareous Tilia forests

Four species belonging to the Cortinarius puellaris group are presented, including the two new species C. biriensis, C. subpuellaris and the recently described C. puellaris. Based on type studies, it is shown that the fourth species in the group should be named C. intempestivus (=C. cristatosporus). The species co-occur and are all studied mainly from SE Norwegian calcareous Tilia forests, but at least some of them also occur in Quercus(-Carpinus) forests in temperate-mediterranean areas of C-S Europe, and are apparently widespread and much overlooked. These are all tiny, small, ochre-redbrown telamonioid cortinarii with strongly to spiny ornamented spores. Phylogenetically, the taxa are well-distinguished by 8 to 22 ITS differences. Together with two taxa only known from ectomycorrhizal ITS sequences, they constitute an apparently well-supported Glade with uncertain affinity.Peer reviewe

Taming the beast: a revised classification of Cortinariaceae based on genomic data

© The Author(s) 2022. This article is licensed under a Creative Commons Attribution 4.0 International License, http://creativecommons.org/licenses/by/4.0/.Abstract: Family Cortinariaceae currently includes only one genus, Cortinarius, which is the largest Agaricales genus, with thousands of species worldwide. The species are important ectomycorrhizal fungi and form associations with many vascular plant genera from tropicals to arctic regions. Genus Cortinarius contains a lot of morphological variation, and its complexity has led many taxonomists to specialize in particular on infrageneric groups. The previous attempts to divide Cortinarius have been shown to be unnatural and the phylogenetic studies done to date have not been able to resolve the higher-level classification of the group above section level. Genomic approaches have revolutionized our view on fungal relationships and provide a way to tackle difficult groups. We used both targeted capture sequencing and shallow whole genome sequencing to produce data and to perform phylogenomic analyses of 75 single-copy genes from 19 species. In addition, a wider 5-locus analysis of 245 species, from the Northern and Southern Hemispheres, was also done. Based on our results, a classification of the family Cortinariaceae into ten genera—Cortinarius, Phlegmacium, Thaxterogaster, Calonarius, Aureonarius, Cystinarius, Volvanarius, Hygronarius, Mystinarius, and Austrocortinarius—is proposed. Seven genera, 10 subgenera, and four sections are described as new to science and five subgenera are introduced as new combinations in a new rank. In addition, 41 section names and 514 species names are combined in new genera and four lecto- and epitypes designated. The position of Stephanopus in suborder Agaricineae remains to be studied. Targeted capture sequencing is used for the first time in fungal taxonomy in Basidiomycetes. It provides a cost-efficient way to produce -omics data in species-rich groups. The -omics data was produced from fungarium specimens up to 21 years old, demonstrating the value of museum specimens in the study of the fungal tree of life. This study is the first family revision in Agaricales based on genomics data and hopefully many others will soon follow.Peer reviewedFinal Published versio

New national and regional biological records for Finland 1. Contributions to agaricoid and ascomycetoid taxa of fungi

Here we present records on 59 fungal taxa, based on recent field work and on herbarium studies. Of those 22 are reported here as new to Finland. Information concerning 37 taxa published elsewhere, is brought here together. Coordinates are given in Uniform Coordinate System (UCS)

Type studies and fourteen new North American species of Cortinarius section Anomali reveal high continental species diversity

Section is a species-rich group in North America belonging to , the most diverse genus in the Agaricales. This study is based on extensive morphological investigations and molecular methods using 191 nrDNA ITS sequence data and recovered 43 phylogenetic species from which 14 are described here as new to science. We sequenced ten type materials which belonged to eight species. The synonymy of with and with is proposed here. The North American occurrence of four species ( , , , and ), so far known only from Europe, was confirmed. Thirteen species were not formally described here due to lack of relevant information. An identification key to the known species in North America is provided

Pushing the Frontiers of Biodiversity Research: Unveiling the Global Diversity, Distribution, and Conservation of Fungi

Fungi comprise approximately 20% of all eukaryotic species and are connected to virtually all life forms on Earth. Yet, their diversity remains contentious, their distribution elusive, and their conservation neglected. We aim to flip this situation by synthesizing current knowledge. We present a revised estimate of 2–3 million fungal species with a “best estimate” at 2.5 million. To name the unknown >90% of these by the end of this century, we propose recognition of species known only from DNA data and call for large-scale sampling campaigns. We present an updated global map of fungal richness, highlighting tropical and temperate ecoregions of high diversity. We call for further Red List assessments and enhanced management guidelines to aid fungal conservation. Given that fungi play an inseparable role in our lives and in all ecosystems, and considering the fascinating questions remaining to be answered, we argue that fungi constitute the next frontier of biodiversity research

New national and regional biological records for Finland 4. Contributions to agaricoid and ascomycetoid taxa of fungi 3

Agaricoid fungi (Basidiomycota): Cortinarius chromataphilus, Cortinarius croceocaeruleus, Cortinarius disjungendulus, Cortinarius nolaneiformis, Cortinarius olididisjungendus, Cortinarius piceidisjungendus, Crepidotus stenocystis, Entoloma callirhodon, Entoloma rhombisporum, Hydropus marginellus, Macrocystidia cucumis var. latifolia, Russula clementinae, Russula fellea, Russula mairei, Russula roseicolor and three ascomycetoid fungi (Ascomycota): Eutypella extensa, Octospora coccinea and Sporormiella megalospora are reported as new to Finland. Information of species recently published elsewhere: Cortinarius balteatialutaceus, Cortinarius balteatibulbosus, Cortinarius balteaticlavatus, Cortinarius boreicyanites, Cortinarius boreidionysae, Cortinarius brunneiaurantius, Cortinarius caesiocolor, Cortinarius caesiolamellatus, Cortinarius caesiophylloides, Cortinarius cremeiamarescens, Cortinarius flavipallens, Cortinarius infractiflavus, Cortinarius kytoevuorii, Cortinarius luteiaureus, Cortinarius myrtilliphilus, Cortinarius ochribubalinus, Cortinarius pallidirimosus, Cortinarius subrubrovelatus, Cortinarius talimultiformis, Inocybe fuscescentipes and Inocybe subpaleacea is brought here together. New records of little collected and rare taxa Camarops tubulina, Catathelasma imperiale, Entoloma longistriatum, Galerina calyptrospora, Hyaloscypha epiporia, Hymenogaster tener, Porpoloma metapodium and Sowerbyella imperialis are also listed. Corrections of previous information are given on: Cortinarius balteatoalbus, Cortinarius crassifolius (under C. coracis), Cortinarius cyanites, Cortinarius dionysae and Inocybe lindrothii

New national and regional biological records for Finland 3. Contributions to agaricoid and ascomycetoid taxa of fungi 2

Agaricoid fungi (Basidiomycota): Amanita flavescens, Cortinarius brunneotinctus, Cortinarius claroplaniusculus, Cortinarius orasericeus, Cortinarius pardinipes, Cortinarius rufoallutus, Entoloma querquedula, Galerina calyptrospora, Galerina lacustris and Tricholomopsis flammula are reported as new to Finland. Three genera of ascomycetoid fungi (Ascomycota): Ascocodinea, Pseudoproboscispora, Scabropezia and 16 species are reported as new to Finland: Arachnocrea stipata, Ascocodinea polyporicola, Capronia parasitica, Diaporthe dulcamarae, Endoxylina astroidea, Gnomoniopsis tormentillae, Gorgoniceps hypothallosa, Helvella latispora, Helvella terrestris, Peziza celtica, Hypocrea estonica, Mycosphaerella topographica, Ophiobolus galii, Pseudoproboscispora caudae-suis, Roesleria subterranean, Scabropezia scabrosa. Information of species recently published elsewhere: Cortinarius alticaudus, Cortinarius bovinatus, Cortinarius cobaltinus, Cortinarius olympianus, Cortinarius sannio, Cortinarius sobrius, Cortinarius subfoetens is brought here together. New records of little collected and rare taxa: Cortinarius barbaricus, Desmazierella piceicola and Tuber anniae s.lat. are also listed. Corrections of previous information are given on: Cortinarius luteobrunnescens, Cortinarius metarius, Cortinarius patibilis and Cortinarius pseudonaevosus

New national and regional biological records for Finland 5. Contributions to agaricoid and ascomycetoid taxa of fungi 4

One genera of agaricoid fungi (Basidiomycota): Romagnesiella and 12 species are reported as new to Finland: Agaricus macrocarpus, Arrhenia obatra, Arrhenia obscurata, Arrhenia rigidipes, Coprinellus brevisetulosus, Coprinus candidatus, Entoloma plebejum, Hydnum vesterholtii, Inocybe phaeocystidiosa, Mycena clavata, Omphalina arctica and Romagnesiella clavus. Two genera of ascomycetoid fungi (Ascomycota): Strossmayeria, Phaeomollisia and 8 species are reported as new to Finland: Arachnopeziza delicatula, Hyaloscypha diabolica, Hyalopeziza cf. tianschanica, Phaeomollisia piceae, Phialina pseudopuberula, Sphaeropezia hepaticarum, Strossmayeria basitricha and Trichopeziza subsulphurea. Information of species recently published elsewhere: Cortinarius angustisporus, C. cacaodiscus, C. caesioarmeniacus, C. centrirufus, C. crassisporus, C. cruentiphyllus, C. davemallochii, C. ferrugineovelatus, C. furvus, C. fuscescens, C. murinascens, C. privignipallens, C. pseudofervidus, C. roseivelatus, C. roseocastaneus, C. subbrunneoideus, C. subexitiosus, C. squamivenetus, C. uraceisporus, Hebeloma eburneum, H. salicicola, Hygrophorus exiguus and Psathyrella fennoscandica is brought here together. New records of little collected and rare taxa Coprinopsis patouillardii, Cuphophyllus cinerellus, Galerina perplexa, Galerina pruinatipes, Gorgoniceps hypothallosa, Inocybe boreocarelica, Marasmius setosus, Psathyrella potteri, Psathyrella tenuicula, Russula adulterina, Russula pyriodora, Scutellinia trechispora, Sowerbyella imperialis and Volvariella murinella are also listed. Corrections of previous information are given on: Cortinarius angulosus (under C. duristipes), Coprinopsis patouillardii and Psathyrella potteri