Ilkka Kytövuori

Cortinarius sect. Brunnei (Basidiomycota, Agaricales) in North Europe.

The section Brunnei was extensively studied based on material from North Europe. To stabilise the nomenclature we studied the relevant types of taxa included in this section. Phylogenetic relationships and species limits were investigated using rDNA ITS sequences and the results were compared with the morphological data. We recognised 11 species: Cortinarius brunneus, C. clarobrunneus comb. nov., C. coleoptera, C. ectypus, C. gentilis, C. glandicolor (neotypified), C. pseudorubricosus, and four species described as new C. caesiobrunneus, C. albogaudis, C. carabus, and C. cicindela. They are described here and their taxonomy, ecology, distribution, and relationships are discussed. In addition, a key to species of the section Brunnei is provided. A total of 77 new sequences of 11 species are published including nine type sequences. Also the taxonomic assignments of sequences in the public databases belonging to the section Brunnei are revised.

The largest type study of Agaricales species to date: bringing identification and nomenclature of Phlegmacium (Cortinarius) into the DNA era

Cortinarius is a species-rich and morphologically challenging genus with a cosmopolitan distribution. Many names have not been used consistently and in some instances the same species has been described two or more times under separate names. This study focuses on subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project were to: i) study type material to determine which species already have been described; ii) stabilize the use of Friesian and other older names by choosing a neo- or epitype; iii) describe new species that were discovered during the process of studying specimens; and iv) establish an accurate ITS barcoding database for Phlegmacium species. A total of 236 types representing 154 species were studied. Of these 114 species are described only once whereas 40 species had one ore more synonyms. Of the names studied only 61 were currently represented in GenBank. Neotypes are proposed for 21 species, and epitypes are designated for three species. In addition, 20 new species are described and six new combinations made. As a consequence ITS barcodes for 175 Cortinarius species are released.

Cortinarius sect. Armillati in northern Europe

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.

Cortinarius sanguineus and equally red species in Europe with an emphasis on northern European material

The red species of Cortinarius subgenus Dermocybe in Europe were studied based on morphological and molecular data. Three completely red species were recognized: C. sanguineus (syn. C. sanguineus var. aurantiovaginatus), C. puniceus (syn. C. cruentus, C. rubrosanguineus) and C. vitiosus comb. nov. Cortinarius sanguineus has dusky red to red pileus, reddish yellow mycelium and lacking or with only slightly encrusted hyphae in pileipellis. It occurs in mesic to damp forests with Picea, often on rich soil in the boreal and montane areas of Europe, presumably also in eastern Canada. Cortinarius puniceus differs from C. sanguineus by its stronger purplish red, narrower spores and spot-like encrusted hyphae in pileipellis. It grows with deciduous trees in the temperate zone of Europe. Cortinarius vitiosus is known only from Fennoscandia and occurs in dry to mesic coniferous forests. It has fairly thin, often zonate, dark red to dark reddish brown pileus, pale red mycelium, small spores and encrusted lamellar trama and pileipellis hyphae. In addition to these three species C. fervidus and C. phoeniceus occasionally have red basidiomes. The relationships of the species were inferred by analysis of ITS sequences. Our study suggests that the section Sanguinei, as earlier defined, is polyphyletic. Here the section is limited to include C. sanguineus, C. puniceus and North American D. sierraensis. The relationships with other red species were not determined. Section Dermocybe, including C. cinnamomeus, C. croceus and C. uliginosus, formed a monophyletic group, and the section Malicoriae had some support. A total of 34 new sequences are published including nine from type specimens.

Two new species in Cortinarius subgenus Telamonia, Cortinarius brunneifolius and C. leiocastaneus, from Fennoscandia (Basidiomycota, Agaricales)

Two new Cortinarius species, C. brunneifolius and C. leiocastaneus, are described based on molecular and morphological data. Cortinarius brunneifolius looks like a trivial brown Telamonia. It is characterised by the fairly dark brown cap and gills contrasting with the white stem. The major distinctive features are the refracting, colourless granules in the epicutis hyphae, broadly ellipsoid, equally verrucose spores, and fruitbody without bluish tints. Cortinarius leiocastaneus can be fairly easily recognised by the appearance of miniature brown Telamonia species, narrowly amygdaloid spores, and a habitat with Betula. Detailed descriptions of the species are provided as well as comparisons with macro- and microscopically similar existing telamonioid taxa. Phylogenetic relationships within the genus were studied by use of ITS rDNA sequence data. Ten new ITS sequences are published. Cortinarius brunneifolius and C. leiocastaneus are fairly closely related and belong to the subgenus Telamonia sensu stricto. The infrasubgeneric relationships were not well resolved, but the new species seem to be unrelated to the macroscopically similar species and do not clearly belong to any of the existing sections.

Two new species of Cortinarius, subgenus Telamonia, sections Colymbadini and Uracei, from Europe

Two new Cortinarius species are described from European Quercus forests and one new combination is made based on molecular and morphological data. Cortinarius uraceomajalis is a vernal species currently only known from Hungary, Bulgaria and Italy, but likely is common throughout central and south-eastern Europe. Cortinarius uraceonemoralis is a widely distributed species in Europe. The new combination Cortinarius nolaneiformis is based on Hydrocybe nolaneiformis Velen. and is a widespread vernal species in Europe. Cortinarius uraceomajalis and C. nolaneiformis are preliminarily placed in sect. Colymbadini, characterized by having a positive (yellow) UV reaction, while C. uraceonemoralis with a UV negative reaction is placed in sect. Uracei. A neotype is selected for C. colymbadinus and C. uraceus to stabilize the nomenclature. Taxonomic novelties: Cortinarius uraceomajalis Dima, Liimat., Niskanen & Bojantchev, Cortinarius uraceonemoralis Niskanen, Liimat., Dima, Kytöv., Bojantchev & H. Lindstr., Cortinarius nolaneiformis (Velen.) Dima, Niskanen & Liimat.

The species of Cortinarius, section Bovini, associated with conifers in northern Europe

Cortinarius bovinus and morphologically similar conifer-associated species were studied using material mainly from northern Europe. To stabilize the nomenclature, relevant types were examined. Phylogenetic relationships and species limits were investigated with rDNA ITS and nuclear rpb2 sequences as well as morphological data. We recognize seven species: C. bovinus (neotypified) and six new species, C. anisochrous, C. bovinaster, C. bovinatus, C. fuscobovinus, C. fuscobovinaster and C. oulankaënsis. Their taxonomy, ecology, distribution and relationships are discussed, and a key to species is provided. Based on our phylogeny and morphological data the species were placed in section Bovini.

Cortinarius, subgenus Telamonia, section Disjungendi, cryptic species in North America and Europe

Cortinarius disjungendus and morphologically similar species were studied using ITS sequences of nine type collections and 18 specimens in the study group, as well as 13 sequences from public sequence repositories, including sequences of three type specimens. In addition, previously published rpb2 sequences were used to provide the phylogenetic position of species in the subgenus Telamonia. Based on these results, the new section Disjungendi, subgenus Telamonia, is proposed. Six species belong to this section: C. claroplaniusculus, C. disjungendus, C. disjungendulus, C. olididisjungendus, C. orasericeus, and C. piceidisjungendus. Of these, C. disjungendus, C. disjungendulus, and C. claroplaniusculus are to date only known from Europe, whereas the others occur both in North America and Europe. Based on ITS sequences, three of the six species had taxonomic synonyms: C. claroplaniusculus (synonym C. hymenoluctus and C. planodepressus), C. disjungendus (synonyms C. remauxii, C. conicosordescens, and C. solivagus), and C. orasericeus (synonym C. brunneofulvus f. myrtillophilus). This study demonstrates how difficult it can be to identify species only on the basis of morphology, and emphasizes the importance of molecular studies of type and of other specimens, particularly in groups with cryptic taxa.

Cortinarius badiolaevis, a new conifer-associated, darkening species in the subgenus Telamonia (Basidiomycota, Agaricales)

A new species, Cortinarius badiolaevis Niskanen, Liimat., Mahiques, Ballarà, Kytöv. in subgenus Telamonia, is described based on morphological and molecular data. Typical for the species is a dark brown cap, pale flesh, indistinct universal veil, small amygdaloid to ellipsoid spores [7.0–8.5 × 4.5–5.0(−5.5) µm], and dark exsiccata. It occurs in mesic to dryish coniferous forests, and grows in calcareous soil presumably with Pinus and Picea. So far, it is only known from Spain and Sweden, and considered rare. Based on the phylogenetic analyses of the ITS regions, it does not belong to any of the known sections, nor does it have any very closely related sister species.

Cortinarius section Bicolores and section Saturnini (Basidiomycota, Agaricales), a morphogenetic overview of European and North American species

Cortinarius is the largest genus of ectomycorrhizal fungi worldwide. Recent molecular studies have shown high levels of morphological homoplasy within the genus. Importantly, DNA phylogenies can reveal characteristics that have been either over- or underemphasized in taxonomic studies. Here we sequenced and phylogenetically analysed a large set of pan-European and North American collections taxonomically studied and placed in Cortinarius sect. Bicolores and sect. Saturnini, according to traditional morpho-anatomical criteria. Our goal was to circumscribe the evolutionary boundaries of the two sections, to stabilize both the limits and nomenclature of relevant species, and to identify described taxa which, according to our current understanding, belong to other lineages. Our analysis resolves two clades: /Bicolores, including 12 species, one of which is new to science, and /Saturnini, including 6 species. Fifteen binomials, traditionally treated in these two sections based on morphology, do not belong to the above two phylogenetic clades. Instead, six of these latter are clearly placed in other clades that represent sect. Bovini, sect. Sciophylli, sect. Duracini and sect. Brunneotincti. The presence or absence of blue pigments and the detection of specific odours emerge as clearly misleading taxonomic features, but more surprisingly, spore size and ecology can be misleading as well. A total of 63 type specimens were sequenced, 4 neotypes and 2 epitypes are proposed here, and 1 new combination is made.