Abstract

INTRODUCTION

Studies using DNA sequence data have shown that the diversity of fungi far exceeds earlier expectations with many common species of macrofungi still to be described and named (Schmit & Mueller 2007). In addition, it is not always easy to determine which species have been described and which are new to science, unless the type specimens of existing names can be carefully studied, including DNA sequencing.

For many species of macrofungi the names have been difficult or even impossible to interpret. The most difficult ones are the older names with brief descriptions and usually without type material. But even if type material exists it can be very difficult to be certain of the identification based only on morphology, especially in challenging genera like Cortinarius where there is considerable convergence in morphology, colouration, and microscopic features. Furthermore, the literature is often difficult to obtain, making it hard to get information on available names and their application by earlier taxonomists. Consequently, many names have not been used consistently and in some cases the same species has been described two or more times under separate names. In instances where there is no type material available, a neotype (or a lectotype if collections of the author are available) is required to stabilize the use of the name. Finally, old type collections that are considered historical materials may not be available for study or DNA sequencing requiring the selection of an epitype.

The development of molecular techniques has provided a more unambiguous tool to identify species. Currently the most commonly used locus in species level taxonomy is the nuclear ribosomal internal transcribed spacer (ITS), which has been proposed as a universal barcode marker for fungi (Schoch et al. 2012). The region is present in several chromosomes and is arranged in tandem repeats that are thousands of copies long (Burnett 2003). Due to the high copy number the region usually is easy to amplify and sequence, even from very old specimens (Larsson & Jacobsson 2004). In Cortinarius the ITS was proposed as a species-identifier sequence already in 2007 by Frøslev et al. and in 2008 by Ortega et al. It has also been shown that in the majority of the cases ITS is suitable for species delimitation in Cortinarius. The results of the multi-gene phylogenetic study based on ITS, rpb1, and rpb2 regions by Frøslev et al. (2005) showed that inference from ITS alone is indicative of the species level phylogenetic delimitations of multi-gene analyses. Furthermore, the delimitations inferred from ITS usually correlate with the morphospecies (Frøslev et al. 2007, Ortega et al. 2008, Niskanen et al. 2012b).

Cortinarius is the largest genus of Agaricales with a cosmopolitan distribution and over 2 000 described species (Kirk et al. 2008). Cortinarius species are important ectomycorrhizal fungi associated with different trees and shrubs, belonging to the order Fagales, families Caesalpiniaceae, Cistaceae, Dipterocarpaceae, Myrtaceae, Pinaceae, Rhamnaceae, Rosaceae and Salicaceae as well as a few herbaceous plants in the Cyperaceae. Owing to their often narrow ecological preferences and sensitivity to environmental change, many Cortinarius species have been used as indicator species for valuable natural environments, e.g. in Sweden and Denmark (Vesterholt 1991, Hallingbäck & Aronsson 1998, Frøslev & Jeppesen 2011). Lately it also was suggested that they have a key role in the carbon cycling of boreal forests (Bödeker et al. 2011).

Many of the major studies in Cortinarius have dealt with North American and especially European species, while the species of southern hemisphere are somewhat less studied (Moser & Horak 1975, Garnica et al. 2002, Cleland 1976 (1935), Gasparini & Soop 2008). In Europe the most extensive studies have been done by Fries (e.g., 1821, 1836–1838, 1851) from Sweden, Henry (e.g., 1951, 1958, 1981, 1986) and Bidaud et al. (e.g., 1992, 2010) from France, Moser (e.g. 1960, 1969–1970, 1983) from Central Europe, Høiland (1984), Brandrud (e.g., 1996, 1998), Brandrud et al. (e.g., 1990, 2012) and Niskanen et al. (e.g., 2009, 2011a, 2013a) mainly from northern Europe, Frøslev et al. (e.g., 2006, 2007) from northern and Central Europe, Orton (1955, 1958) from Great Britain, and Ortega et al. (2008) and Suárez-Santiago et al. (2009) from mediterranean area. Selected papers of some of the major contributors to Cortinarius systematics in North America include Peck (1873; also see Gilbertson 1962), Kauffman (1918, 1923, 1932), Smith (1939, 1942, 1944), Ammirati (1972), Ammirati et al. (2013), Garnica et al. (2009), Liu et al. (1997), Moser & Ammirati (1996, 1999), Moser et al. (1995), Bojantchev (2011a, b) and Niskanen et al. (2013b, c).

Very little is known about the distribution of Cortinarius species on a larger scale or the differences in the species composition between the continents, although species in the southern hemisphere are distinct from those in the northern hemisphere (Peintner et al. 2004, Garnica et al. 2005). However, recent molecular studies on Cortinarius have shed some light on these questions for North America and Europe (Moser & Peintner 2002, Matheny & Ammirati 2006, Garnica et al. 2009, 2011, Harrower et al. 2011, Ammirati et al. 2013, Niskanen et al. 2011b, 2012c, 2013b, c). These studies show several patterns of species distributions. There are species common to North America and Europe, especially those species from more northern and montane conifer forests, i.e. Cortinarius aureofulvus M.M. Moser s.auct., C. napus Fr. and C. pinophilus Soop, but also endemic species occur both in western North America, eastern North America and Europe, i.e. C. elegantio-occidentalisGarnica & Ammirati in western North America, C. hesleri Ammirati, Niskanen, Liimat. & Matheny in eastern North America and C. puniceus P.D. Orton in Europe. Cortinarius species composition is somewhat similar between eastern North America and Europe, but there appears to be less similarity between Europe and western North America.

Different infrageneric classification systems have been proposed for Cortinarius, i.e. Brandrud et al. (1990) divided the genus in four subgenera: Cortinarius, Myxacium, Phlegmacium and Telamonia. Recent molecular analyses have shown that Cortinarius is monophyletic, but also that many of the infrageneric groups such as subg. Phlegmacium are not monophyletic (Peintner et al. 2004, Garnica et al. 2005). Nonetheless, several authors have treated subg. Phlegmacium in similar ways (Moser 1983, Bidaud et al. 1994, Brandrud et al. 2012) including in it species with viscid to glutinous pileus and dry, often bulbous stipe, or dry-capped, stout species with a yellow KOH reaction, i.e. the species of sect. Phlegmacioides (Niskanen et al. 2012a).

Molecular techniques have been in use for more than a decade, and the sequencing of ITS regions even from older Cortinarius specimens is possible. Furthermore, type studies are essential for a stable and consistent application of names in Cortinarius where currently a large percentage of the Cortinarius sequences are incorrectly named or without a name in the public sequence databases (e.g., Niskanen et al. 2011a). While several papers present sequences for individuals or groups of species, for example Garnica et al. (2009) and Niskanen et al. (2011a), the only large study is that of Frøslev et al. (2007) where 52 types of Cortinarius sect. Calochroi were sequenced. The present study focuses on Cortinarius subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project are to: i) study type material to determine which taxa have already 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.

MATERIALS AND METHODS

RESULTS

Phylogenetic analyses

The 50 % majority rule phylogram resulting from the BI analysis is shown in Fig. 1. Most species are supported by 1.00 PP (or slightly less). Six species received support below 0.95 PP: C. castaneicolor, C. collocandoides, C. flavescentipes (= C. balteatocumatilis s.auct.), C. herpeticus, C. infractiflavus and C. multiformis. Three species, C. balteatoalutaceus, C. pseudocephalixus and C. subfoetens did not form monophyletic groups in our phylogenetic analysis. Most of these nine species represent species that differ from their closest relatives by less than 10 substitutions and indel positions in ITS regions, but they all, however, have low intraspecific variation.

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Fig. 1

The Bayesian 50 % majority-rule consensus tree inferred from ITS regions. PP > 0.50 are indicated above branches. In blue colour and bold are new species described in this study as well as new combinations made and in orange and bold neo- and epitypes designated in this study. In pink are species described by French authors, in red species described by other European authors, in green species described by North American authors and in black other specimens included in this study.

In a majority of the species, the intraspecific variation is from 0 to 1 substitutions or indel positions. In some species it is 2 substitutions and/or indel positions. Similarly, in a majority of the species, the interspecific difference is more than 10 substitutions and indel positions, but in some species it is only four, i.e. in the species pair C.claricolor/C. rex-claricolorum. All the species, however, have a clear barcoding gap between intra- and interspecific variation. In the following species the intraspecific variation was more than two substitutions and/or indel positions, but no clear grouping was obtained in the analysis of ITS sequences or the number of specimens was too low for making conclusions: C. albescens/C. gentianeus/C. volvatus complex, C. aureofulvus s.auct./C. orichalceus var. xanthocephalus complex, C. herpeticus/C. violaceonitens complex, C. misermontii/C. olidoamarus var. valentinus/C. subaccedens/C. van-campiae complex, C. pallidirimosus, C. porphyropus and C. scaurus.

A total of 236 types representing 154 species were successfully sequenced. Of these, 114 species are described only once whereas 40 species had one or more synonyms. The species with the largest number of synonyms are C.largus (14 synonyms), C. pseudonaevosus (6 synonyms), C. talus (5 synonyms), C. crassus (4 synonyms) and C.varius (4 synonyms). All the names of the types are listed in alphabetical order in Table 1, followed by the current name.

Several infrageneric groups with three or more species were supported by 1.00 PP: /Calochroi & Fulvi, /Claricolores, /Cyanites, /Dionysae (s.lat. 0.56 PP), /Glaucopodes, /Infracti, /Multiformes, /Phlegmacium, /Purpurascentes and /Scauri. In addition, clade /Arguti (0.98 PP) received some support and clades /Elastici & Percomes (0.94 PP, s.l. 0.53 PP), /Phlegmacioides (0.77 PP) and /Variosimiles (0.74 PP) low support.

NEW SPECIES AND COMBINATIONS

Species with an isolated position

Cortinarius cremeiamarescens Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805786; Fig. 2a, ​a,33a

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Fig. 2

Photo of: a. C. cremeiamarescens TN06-273; b. C. flavivelatus IK98-885; c. C. kytoevuorii TN05-158; d. C. flavipallens T. Kekki 368; e. C. boreidionysae IK97-1220; f. C. cruentipellis IK01-053; g. C. caesiophylloides TEB277-09. — Photos: a, c. Kare Liimatainen; b, e, f. I. Kytövuori; d. T. Kekki; g. T.E. Brandrud.

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Fig. 3

Spores of: a. C. cremeiamarescens TN04-768; b. C. flavivelatus IK98-885; c. C. kytoevuorii TN05-158; d. C. ochribubalinus IK93-641; e. C. subfraudulosus IK11-006, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 μm.

Etymology. The name refers to the colour of the basidiomata and bitter taste of pileus cuticle.

= Cortinarius caesiostramineus Rob. Henry sensu Brandrud et al. 1990, Jeppesen et al. 2012, p.p.

Type. SWEDEN, Gotland, Alskog and När parish, Ollajvs Nature Reserve, mesic to damp spruce forest with some Pinus, Quercus and Corylus, 27 Sept. 2011, I. Kytövuori 11-014 (holotype H; isotype NY). GenBank KF732493.

Pileus 3.5–5.5 cm broad, hemispherical to convex, then expanded, very finely innately fibrillose, cream-coloured to pale ochraceous yellow. Lamellae emarginate, almost crowded, pale greyish brown. Stipe 6.5–9.5 cm long, 0.7–1.4 cm thick at apex, 1–2.3 cm wide at base, with slightly marginate bulb, at first white, becoming pale brownish yellow with age. Universal veil white, sparse, at bulb margin. Context white. Odour indistinct. Taste: pileus cuticle bitter. Exsiccata: pileus ochraceous clay-colour to ochraceous yellow to warm ochraceous brown especially at the centre, stipe pale grey to brown.

In MLZ: Spores 7.0–7.8–8.8 × 4.3–4.7–5.0 μm, av. = 7.5–8.3 × 4.6–4.9 μm, Q = 1.54–1.67–1.87, Qav. = 1.59–1.74 (9 specimens, 260 spores, Fig. 3a), citriform to narrowly fusoid, beaked, thin-walled, fairly finely, densely, and often sharply verrucose, slightly to moderately dextrinoid. Basidia 24–34 × 6.5–8 μm (80 basidia), 4-spored, narrowly clavate, very thin-walled, colourless, more or less filled with blood red drops. Lamellar trama hyphae yellow, filled with small blood red drops, larger globose and worm-like guttules fairly scanty (especially more scanty than in C. gentianeus). Stipe apex hyphae almost colourless to yellow, smooth, full of small golden yellow to blood red drops, larger globose and worm-like guttules fewer. Pileipellis: epicutis strongly gelatinous, hyphae 2–6 μm wide, very thin-walled and difficult to define, filled with small to medium-sized blood red guttules, mostly evenly distributed in the hyphae. Hypoderm present, pale yellow. In C. gentianeus the epicutis hyphae are full of very long, worm-like, foamy guttules and the blood red layer is much thicker than in C. cremeiamarescens.

Ecology & Distribution — In hemiboreal and southern boreal conifer-dominated forests on rich to calcareous soil. Known from southern Europe and western North America, British Columbia and Alaska. Fruits from late August to late October.

Other specimens examined. FINLAND, Varsinais-Suomi, Parainen, Lemlahdensaari, Fallskogen, Pinus sylvestris heath forest with some Picea abies on sandy soil, by the calcareous dust road, 20 Oct. 2006, K. Liimatainen & T. Niskanen 06-273; Uusimaa, Espoo, Luukkaa outdoor recreation area, N of Haukkalampi, mesic, partly grass-herb-spruce forest with some Populus tremula, Betula and Pinus sylvetris, 9 Sept. 2004, K. Liimatainen & T. Niskanen 04-768, H6029461; Kirkkonummi, Dorgarn, Meiko-Trehörningen nature reserve, semi-open spruce forest with some hardwood bushes, 27 Sept. 2007, I. Kytövuori 07-1722, H6001575 (H); Siuntio, Lappträsk, grass-herb-spruce forest with some Pinus, Betula, Populus and Corylus, 24 Aug. 2000, I. Kytövuori 00-027 (H); Etelä-Savo, Mäntyharju, Juolasvesi, Hietaniemi, Sojonkangas, fairly rich spruce-pine forest with abundant Betula and Populus tremula, 29 Sept. 1994, I. Kytövuori 94-1177b, H6035734 (H). – SWEDEN, Västergötland, Undenäs, c. 1.5 km NNW of Sätra bruk, herb-rich Picea abies forest with some Betula, Populus and Pinus, 6 Sept. 2003, T. Niskanen et al. 03-1255, H7018363 (H); F03-1163, H7018395 (H); Närke. Hidinge, Garphyttans Nationalpark, herb-rich Picea abies forest with Corylus, Populus tremula, Betula and Quercus, 26 Sept. 2004, T. Niskanen et al. 03-1255, H7017775 (H).

Additional specimens. CANADA, British Columbia, Interior Cedar Hemlock Forest, mycorrhizal root tip of Betula papyrifera, isolate UBCOCS153F, GenBank EF218754. – USA, Alaska, Delta Junction, source: boreal forest, soil 0–20 cm, GenBank JN889840.

Notes — Cortinarius cremeiamarescens was previously confused taxonomically with C. gentianeus Rob. Henry (= C. caesiostramineus sensu Jeppesen et al. 2012 p.p.). However, the latter species is larger, typically has paler, more whitish or greyish exsiccata and larger (7.7–8.5–9.3 × 4.8–5.2–5.4 μm, Q = 1.54–1.65–1.76), amygdaloid to citriform, less thin-walled, more dextrinoid, and more verrucose spores, and much more abundant large, blood red, foamy, worm-like guttules in the pileipellis and lamellar trama. Cortinarius cremeiamarescens formed a well-supported clade in our phylogenetic analysis (1.00 PP). The ITS sequences of the species are identical and it differs from its sister species C. gentianeus by 17 substitutions and indel positions. Further relationships with other species of Cortinarius were not resolved in our analysis.

Cortinarius flavivelatus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805863; Fig. 2b, ​b,33b

Etymology. The name refers to the yellow universal veil.

Type. SWEDEN, Norrbotten, Pajala, Junosuando, Nature Reserve Area between Sarvikero and Tulemajoki, dryish Picea abies heath forest with Pinus, Betula, and with open meadows, 15 Aug. 1998, I. Kytövuori 98-885 (holotype H; isotype NY). GenBank KF732528.

Pileus 5–8 cm broad, hemispherical to convex, then plano-convex, viscid, finely innately fibrillose, olive brown to ochraceous brown to brown at the centre, lighter at the margin, with hygrophanous streaks. Lamellae emarginate, crowded, first distinctly pale bluish, later pale brown with a bluish tint. Stipe 6–10 cm long, 1–1.8 cm thick at apex, 1.5–2.5 cm wide at base, clavate to almost cylindrical, whitish, with a bluish tint at the apex. Universal veil yellow, forming girdles on stipe, somewhat viscid. Context white in pileus and lower part of the stipe, bluish at stipe apex. Odour indistinct. KOH-reaction negative in all parts. Exsiccata: pileus warm yellowish to reddish brown, stipe whitish.

In MLZ: Spores 9.1–9.8–10.9 × 5.0–5.4–5.9 μm, Q = 1.67–1.82–1.98 (1 specimen, 60 spores, Fig. 3b), amygdaloid-fusoid to slightly, narrowly citriform, with a shallow suprahilar depression and somewhat beaked apex, fairly finely, separately verrucose, slightly dextrinoid. Basidia 27–38 × 7.5–8 μm (40 basidia), 4-spored, clavate, pale sand brown, with few dark red brown granules, almost hyaline when mature. Lamellar trama hyphae: with abundant dark granules and chips, sometimes in small mounds. Stipe apex hyphae pale yellowish sand brown with small brown granules, outermost hyphae entangled, narrow, with more or less abundant blackish red granules. Pileipellis: epicutis strongly gelatinous, uppermost hyphae 5–10 μm wide, ochre brown, finely to strongly spirally incrusted, mostly not granulose, lower down dark-granulose. Hypoderm well developed, red brown. The upper part of the hypoderm and the transition hyphae towards the epicutis strongly incrusted and with mounds of blackish brown granules. Evenly distributed small granules present (sometimes absent) between the mounds.

Ecology & Distribution — In northern boreal coniferous forests. Known from Sweden, Norrbotten. No sequences of this species exist in public databases.

Notes — Based on morphology and molecular data C. flavivelatus is a sister species of C. pini Brandrud. Cortinarius pini, however, has white, sometimes ochraceous white universal veil and much larger spores (10.7–11.7–12.9 × 6.3–6.8–7.3 μm). In ITS regions the difference between the species is 17 substitutions and indel positions.

Cortinarius kytoevuorii Niskanen & Liimat., sp. nov. — MycoBank MB805865; Fig. 2c, ​c,33c

Etymology. The species is named in honour of Ilkka Kytövuori, a mycologist from Finland.

Type. FINLAND, Koillismaa, Kuusamo, Oulanka, Ampumavaara, S slope, old, grass-herb Picea abies forest with some Betula, Pinus sylvestris and Populus tremula, on calcareous soil, 17 Sept. 2005, T. Niskanen & K. Liimatainen 05-158, H6029355 (holotype H; isotype NY). GenBank KF732529.

Pileus 6–9 cm broad, hemispherical to convex, then expanded, finely innately fibrillose, yellow brown to brown, with hygrophanous streaks. Lamellae emarginate, almost crowded to medium spaced, at first pale brownish grey, becoming more brown with age. Stipe 6–9 cm long, 1.2–1.5 cm thick at apex, 2–2.5 cm wide at base, with fairly narrow, marginate bulb, at first white, becoming pale brownish yellow with age. Context whitish to pale yellow. Odour indistinct. KOH reactions: in pileipellis brown (no reddish tints); in context, mycelium and bulb margin negative. Exsiccata: pileus dull, dark red brown overall, stipe almost con-colorous with pileus.

In MLZ: Spores: 7.5–8.5–9.5 × 5.0–5.3–5.4 μm, Q = 1.54–1.61–1.72 (1 specimen, 60 spores, Fig. 3c), amygdaloid-ellipsoid, very strongly, separately, ± sharply verrucose (C. porphyropus like but more dextrinoid), slightly to moderately dextrinoid. Basidia: 24–32 × 7.5–9 μm (20 basidia), 4-spored, clavate. Lamellar trama hyphae: pale pellucid yellowish, guttulate, smooth. Stipe apex hyphae colourless to pale straw-coloured, smooth, guttulate. Pileipellis: epicutis in overall view orange, (very) weakly gelatinous, hyphae 5–10 μm wide, finely to strongly spirally incrusted, many of the uppermost ones abundantly with intercellular, unevenly distributed, orange red granules, granule mounds and concretions. Large-celled hypoderm present, yellowish and with scanty small orange granules in the upper part, lower down colourless.

Ecology & Distribution — In coniferous forests, on calcareous soil. Fruits in autumn.

Notes — Cortinarius kytoevuorii is reminiscent of C. glaucopus but is more slender, has a yellow brown to brown pileus, and lacks bluish tints in basidiomata. It is most easily recognised by the orange red granules and concretions in the uppermost hyphae of the pileipellis when mounted in MLZ. The sister species C. subrugulosus Bidaud & Armada has a more southern distribution. The northernmost known collection is from Sweden, Öland under Fagus (Kytövuori 98-2578 (H)). Furthermore, the spores are shorter (7.3–7.9–8.6 × 5.0–5.3–5.7 μm), relatively broader (Qav. = 1.5), less verrucose, and more strongly dextrinoid, and in the pileipellis small yellow to blood-red guttules are seen in MLZ. In our phylogenetic analysis C. kytoevuorii formed a well-supported clade (1.00 PP) with C. subrugulosus, but further relationships were not solved. The difference in ITS regions between the two sister species is 11 substitutions and indel positions.

Cortinarius ochribubalinus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805866; Fig. 3d

Etymology. The name refers to the colour of the basidiomata.

Type. FINLAND, Uusimaa, Espoo, Nuuksio, the open-air territory of Pirttimäki, between the main road and the lake, opposite the parking area, fairly rich grass-herb forest with Populus tremula, Betula, Alnus incana, Quercus, Corylus, Prunus padus, Salix spp., and some old pines and young spruce, 2 Sept. 1993, I. Kytövuori 93-641, H6032734 (holotype H; isotype NY). GenBank KF732530.

Pileus 5–8 cm broad, convex, soon plano-convex, sometimes with a broad umbo, very finely fibrillose, centre ochraceous, whitish towards margin. Lamellae emarginate, medium spaced, at first very pale brownish grey, later pale brown. Stipe 6–10 cm long, 0.8–1.3 cm thick at apex, 1.5–2 cm at base, clavate, at first white, becoming pale brownish yellow with age. Universal veil white, on pileus margin thin, forming thin belts on the stipe. Context white. Odour pleasant. Exsiccata: pileus warm yellowish brownish at centre, pale leather-coloured to almost whitish at margin, stipe concolorous with the pileus margin.

In MLZ: Spores 12.5–13.4–14.3 × 7.5–7.9–8.2 μm, Q = 1.57–1.69–1.84 (1 specimen, 60 spores, Fig. 3d), amygdaloid, strongly verrucose, most strongly so at the apex, very much like those in C. riederi group or C. violaceus, moderately dextrinoid. Basidia 39–48 × 9–12 μm (20 basidia), light sand brown, with large to small blood black granules. Lamellar trama hyphae with blood black substance and/or same-coloured granules, dark chips almost lacking. Stipe apex hyphae yellow, with colourless guttules, coloured granules lacking, but outermost hyphae sand brown to somewhat redder, with few to abundant blood red, small granules. Velum/Cortina hyphae sand brown to somewhat redder, with few to abundant blood red, small granules. Pileipellis: epicutis somewhat gelatinous, hyphae 3–10 μm wide, near the surface ochraceous brown, finely, densely incrusted, mostly not granulose, lower down strongly granulose with small to large blackish brown granules in mounds and long, sausage-shaped clusters and concretions. Hypoderm well developed, red brown, hyphae mostly granulose in the upper part.

Ecology & Distribution — With deciduous trees, possible hosts Populus, Corylus or Quercus. Known from South Finland. No sequences of this species exist in public databases.

Notes — Based on our molecular studies C. ochribubalinus does not have any known, close relatives. It holds an isolated position in the phylogenetic tree and differs in ITS regions by more than 20 substitutions and indel positions from the closest species C. patrickensis (M.M. Moser) Niskanen, Liimat., Kytöv., Bojantchev & Ammirati. Morphologically, it is reminiscent of species in /Arguti. The spores are large and similar to those of C. fraudulosus and C. subfraudulosus, but more strongly verrucose. The smell of the lamellae is also different, pleasant in C. ochrobubalinus and often earthy-raphanoid in C. fraudulosus and C. subfraudulosus. In addition, the universal veil is sparse where as C. fraudulosus and C. subfraudulosus have more abundant sometimes even floccose universal veil remnants on stipe.

/ARGUTI

Cortinarius patrickensis (M.M. Moser) Niskanen, Liimat.,

Kytöv., Bojantchev & Ammirati, comb. nov. — MycoBank MB805883

Basionym. Cortinarius fraudulosus var. patrickensis M.M. Moser, Mycotaxon 74, 1: 10. 2000.

Type. USA, California, Humboldt Co., Patrick’s Point State Park prope Trinidad, in coniferous forest (Picea sitchensis, Pseudotsuga menziesii), 25 Nov. 1995, M. Moser 95/617 (holotype IB). GenBank KF732307.

Other specimens examined. SWEDEN, Medelpad, Borgsjö, Julåsen, SE sloping, partly paludified, rich grass-herb-spruce forest with some pines and birches, 15 Sept. 1995, I. Kytövuori 95-1400, H7019584 (H). – USA, Six Rivers Nat’l forest, off Hwy 299, 1004 m asl, mixed forest (Notholithocarpus densiflorus, Quercus kelloggii, Pinus spp., Pseudotsuga menziesii, etc.), 20 Nov. 2010, D. Bojantchev DBB39406 (UC).

Notes — Cortinarius patrickensis is a typical member of /Arguti, it has a whitish to pale brown basidiomata and large spores, and the placement is also supported by molecular data. It was first described as a variety of C. fraudulosus but based on our genetic and morphological data it should be treated as a species. In our phylogenetic analysis C. patrickensis grouped together with C. rosargutus Chevassut & Rob. Henry (0.86 PP). Also with the pairwise comparison the closest species is C. rosargutus from which it differs by 5 substitutions and indel positions in ITS regions. Both taxa, C. patrickensis and C. rosargutus, include several identical or almost identical sequences and have a clear barcoding gap (no overlap between intra- and interspecific variation). A description of the species is provided in Moser & Ammirati (2000). Typical for the species are amygdaloid, large spores 12.2–13.3–14.7 × 7.0–7.7–8.4 μm, Q = 1.62–1.73–1.84, and green apple-like to strong green corn-like odour. The spores are narrower than those of C. rosargutus (Qav. = 1.60). Cortinarius patrickensis grows in coniferous forests, often on calcareous soil. It was previously only known from California, USA but is here reported for the first time from Europe in Sweden. Species might be occasional in suitable habitats, at least in Europe and North America, but has most likely been misidentified as either C. fraudulosus or C. rosargutus.

Cortinarius subfraudulosus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805867; Fig. 3e

Etymology. The name refers to the affinity of C. fraudulosus.

Type. NORWAY, Oppland, Lunner, Skøyenåsen, Picea abies forest with Corylus on calcareous soil, 3 Sept. 2011, I. Kytövuori 11-006 (holotype H; isotype NY). GenBank KF732564.

Illustrations — Brandrud et al. (1990: pl. A07 as C. argutus subsp. fraudulosus), Abarenkov et al. (2010: photo under the accession no. UDB011261).

Pileus 4–10 cm broad, hemispherical to convex, then plano-convex, finely fibrillose, white to ochraceous white when young, with age becoming pale brownish. Lamellae emarginate, medium spaced to almost distant, first white to very pale brownish grey, later pale brown. Stipe 5–10 cm long, 1–2 cm thick at apex, 1.5–3 cm wide at base, clavate to slightly bulbous, sometimes slightly rooting, whitish, becoming brownish with handling and with age. Universal veil white, forming distinct girdles on stipe, sometimes floccose. Context white, but becomes very slowly black when bruised. Odour weak, a combination of earthy and raphanoid. Exsiccata: pileus dirty greyish to yellow brown at the centre, whitish to yellowish brown towards the margin, stipe greyish white to pale brown.

In MLZ: Spores 12.0–13.6–15.0 × 7.0–7.8–8.4 μm, av. = 12.5–14.4 × 7.6–8.2 μm, Q = 1.57–1.72–1.88, Qav. = 1.64–1.75 (4 specimens, 100 spores, Fig. 3e), amygdaloid (to weakly citriform), with a narrow apex, moderately to fairly strongly verrucose, warts anastomosing, not high, moderately dextrinoid. Basidia 38–55 × 10–12 μm (2 specimens, 60 basidia), almost colourless to pale sand brownish, clear or with few small, brown granules. Lamellar trama hyphae full of dark granules or chips or blood black particles. Stipe apex hyphaeyellow to reddish brown, outermost ones with abundant large red brown to blood blackish granule masses or opaque particles. Velum/Cortina hyphae with abundant large red brown to blood blackish granule masses or opaque particles. Pileipellis: epicutis fairly weakly gelatinous, hyphae at the surface 3–10 μm wide, finely incrusted, ochraceous brownish, mostly not granulose, lower down somewhat granulose or not. In the transition between epicutis and hypoderm up to 15 μm wide, dark red brown, strongly incrusted hyphae with large, blackish granula mounds, intracellular concretions and/or red brown, 3–15 μm wide extracellular pigment mounds on the hyphae, with few evenly distributed small granules. Hypoderm well developed, red brownish to very dark red brown, somewhat granulose or not.

Ecology & Distribution — In hemiboreal and southern boreal coniferous forests, calcicolous. Known from Fennoscandia and Estonia. Fruiting in autumn, in September.

Other specimens examined. ESTONIA, Hiiumaa, Kõrgessaare, Kõpu, old spruce forest with Pinus, Betula, Quercus, Populus tremula and Corylus, on calcareous soil, 15 Sept. 2001, T. Niskanen & I. Kytövuori (H); Pühalepa, Kallaste pank, spruce forest with Pinus, Betula, Populus tremula and Corylus, on calcareous soil, 23 Sept. 2008, J. Vesterholt & J. Vauras 26655F (TUR-A). – SWEDEN, Östergötland, Väversunda parish, E sloping, very rich spruce grass-herb forest with hardwood bushes, 26 Sept. 1988, I. Kytövuori 88-2016 (H); Uppland, Börstils sn, Marieberg, Täpporna, in rich spruce forest, 11 Sept. 1986, T.E. Brandrud et al. CFP481, F44801 (S); Södermanland, Mörkö parish, Oaxen, dryish spruce grass-herb forest with Pinus, Betula, Populus tremula and Salix spp., on calcareous soil, 27 Sept. 1998, I. Kytövuori 98-2244 (H), 98-2245 (H).

Additional specimen. ESTONIA, Saare, Kihelkonna, Mäebe, in coniferous forest, 21 Sept. 2009, V. Liiv TU106607, UNITE No. UDB011261 as C. fraudulosus (TU(M)).

Notes — The species has earlier been called C. fraudulosus in northern Europe. Cortinarius fraudulosus, however, has been described by Britzelmayr (1885) from Siebentischwald, Bavaria, Germany. In our phylogenetic tree the northern material and the more southern European material formed two separate, well-supported clades which differ from one another by 9 substitutions and indel positions in their ITS regions. The sequences of C. subfraudulosus have one base polymorphisms and the maximum pairwise distance is zero. Therefore, we here describe the northern species as new.

/CALOCHROI & FULVI

Cortinarius flavipallens Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805868; Fig. 2d, ​d,44a

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Fig. 4

Spores of: a. C. flavipallens IK08-1729; b. C. boreicyanites CFP931; c. C. boreidionysae IK97-1220; d. C. cruentipellis IK98-2503; e. C. luteiaureus IK07-247a, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 μm.

Etymology. The name refers to the colour of the pileus.

Type. FINLAND, Kainuu, Kajaani, Hietalahti, Torakangas NNW of Korpitaipale, by the power line, fairly damp grass-herb-spruce forest with Pinus, Betula and Populus tremula, on calcareous soil, 13 Sept. 2008, I. Kytövuori 08-1729, H6032745 (holotype H; isotype NY). GenBank KF732554.

Pileus 4–8 cm broad, hemispherical to convex, soon expanded, pale ochraceous to pale brown. Lamellae emarginate, crowded, pale grey to pale greyish brown. Stipe 4–6 cm long, 1–1.5 cm thick at apex, 2–2.5 cm wide at base, with a marginate bulb, white. Universal veil whitish to pale brown at bulb margin. Context white. Odour indistinct. KOH reactions on pileus and bulb margin red, on basal tomentum and rhizomorphs slowly and/or weakly vinaceous pink. Exsiccata: pileus pale ochre to ochre brown, darkest at the centre, paler towards the margin, stipe greyish white to pale brown.

In MLZ: Spores 9.5–10.7–11.6 × 5.4–6.0–6.3 μm, av. = 10.3–11.0 × 5.9–6.1, Q = 1.65–1.77–1.85, Qav. = 1.73–1.80 (3 specimens, 120 spores, Fig. 4a), amygdaloid-fusoid to weakly citriform, with a shallow suprahilar depression and mostly a somewhat blunt apex, moderately verrucose, warts anastomosing or not, slightly to moderately dextrinoid. Basidia 26–38 × 8–10 μm (60 basidia), 4-spored, clavate, very pale yellowish, clear, few slightly granulose-guttulate. Lamellar trama hyphae pale yellowish, smooth, sometimes guttulate, some colourless crystals solitary or in roundish masses, not in branch-like fascicles. Pileipellis: epicutis strongly gelatinous, hyphae on the surface 3–5 μm wide, very pale ochre, smooth to very finely, densely, spirally incrusted, walls distinctly visible, also some hyphae with yellow, foamy contents. Lower down equally wide to slightly wider, strongly incrusted hyphae, often in bundles. Hypoderm absent. In KOH the gelatinized hyphae pale vinaceous pink.

Ecology & Distribution — In boreal Picea abies dominated forests on calcareous soil. So far only known from Finland but might be widely distributed, since an ITS sequence (FJ039640) from a specimen collected in British Columbia in western Canada differed only by 4 substitutions and indel positions from Finnish material and might be conspecific.

Other specimens examined. FINLAND, Laatokan Karjala, Parikkala, Vaaranperä, Soininmäki, S part, Soininjoki, fairly old, SE sloping spruce forest with some Pinus, Betula and Populus tremula, 17 Sept. 2009, A. Ahola H6032393 (H); Perä-Pohjanmaa, Tervola, Peura, Raemäki, in Picea dominated forest on calcareous soil, 5 Sept. 2011, T. Kekki 368 (H).

Notes — Cortinarius flavipallens with its pale ochraceous colours is in appearance between C. metarius Kauffman (= C. barbarorum Bidaud, Moënne-Locc. & Reumaux) and C. caesiocinctus Kühner, without the bright yellow pileus of the former or the bluish grey one of the latter. In addition, the KOH reaction in the mycelium is slower and weaker than in the former two species. Cortinarius piceae Frøslev, T.S. Jeppesen & Brandrud and C. corrosus Fr. differ from C. flavipallens by their negative KOH reaction in the basal mycelium. The ITS sequences of C. flavipallens were identical and it formed a well-supported clade in our phylogenetic analysis. However, the relationships with other species of /Calochroi p.p. were not resolved.

Cortinarius luteicolor (A.H. Sm.) Ammirati, Bojantchev, Niskanen & Liimat., stat. nov. & nom. nov. — MycoBank MB805896

Etymology. The name refers to the yellowish colours of the pileus and lamellae.

Basionym. Cortinarius orichalceus var. olympianus f. luteifolius A.H. Sm., Lloydia 7: 185. 1944.

Type. USA, Washington, Olympic Mts, near Lake Angeles, 19 Sept. 1941, A.H. Smith 16970, barcode 10389 (holotype MICH). GenBank KF732368.

Other specimens examined. USA, California, Sierra Nevada, Yosemite Nat’l Park, Hwy 120, 2433 m asl, Pinus contorta, Pseudotsuga menziesii, Abies concolor, A. magnifica, etc., 16 Nov. 2011, D. Bojantchev DBB46740 (UC), 14 Nov. 2010, D. Bojantchev DBB38211 (UC).

Additional specimens. CANADA, British Columbia, VMS13, GenBank FJ-717511 as ‘Cortinarius sp.’ (UBC). – USA, Oregon, Clackamas Co., Bull Run Watershed, Tsuga heterophylla, Pseudotsuga menziesii, 11 Nov. 1995, M.M. 95/500/ J.F. Ammirati 11701, GenBank EU057024 as ‘sp.’; Washington, Chelan Co., Chatter Creek, Tsuga heterophylla, Pseudotsuga menziesii, 6 Oct. 2007, J. Birkebak JMB10-06-2007-03 (UBC), GenBank HM068562 as ‘C. cupreorufus’.

Notes — ITS sequence analysis shows C. luteicolor as a well-delimited species within clade /Laeticolores. Morphologically, it is not similar to any of the species we have studied. Initially, the species was described as a form of C. orichalceus var. olympianus (= C. pseudocupreorufus (A.H. Sm.) Niskanen, Liimat. & Ammirati). In the ITS regions it, however, differs from C. pseudocupreorufus by more than 25 substitutions and indel positions. Furthermore, Smith (1944) noticed several differences among these taxa: “This form (= C. luteicolor) differs from the typical form (= C. pseudocupreorufus) in lacking the faint lilac tint in the apex of stipe, in the pileus not becoming dark vinaceous red but instead merely dull cinnamon brown when fresh, and in the brighter yellow lamellae. The pilei of the herbarium specimens are also paler, but the bulb is deep purplish as in typical material of the variety (= C. pseudocupreorufus)”. Based on morphology and molecular data we conclude that C. luteicolor should be treated as a species. Description of the species is presented in Smith (1944). Typical for the species are at first rich dull yellow or yellow tinged pileus gradually becoming dull cinnamon, pale yellow gills, and subalmond-shaped spores (9–11.5 6–7 μm). Cortinarius luteicolor grows in coniferous forests (Pseudotsuga, Tsuga) and is currently known from Pacific northwest of North America, from California and Washington, USA, and British Columbia, Canada.

Cortinarius pseudocupreorufus (A.H. Sm.) Niskanen, Liimat. & Ammirati, stat. nov. & nom. nov. — MycoBank MB805886

Etymology. The name refers to the affinity to C. cupreorufus.

Basionym. Cortinarius orichalceus var. olympianus A.H. Sm., Lloydia 7, 3: 184. 1944.

Type. USA, Washington, Olympic National Park, Olympic Hot Springs, under conifers, 2 Oct. 1941, A.H. Smith 17513, barcode 10390 (holotype MICH). GenBank KF732367.

Notes — In our phylogenetic analysis C. pseudocupreorufus is placed as a sister species of C. cupreorufus Brandrud from which it differs by 10 substitutions and indel positions in ITS regions. The spores of C. pseudocupreorufus are 9–11 × 5–6.5 μm and somewhat amygdaloid, while the spores of C. cupreorufus are broader, 10–11.5 × 6.5–7.5 μm, and amygdaloid-citriform. Based on morphology and molecular data we conclude that C. pseudocupreorufus should be treated as a species. Description of the species is presented in Smith (1944). The species is so far only known from the type locality.

/CYANITES

Cortinarius boreicyanites Kytöv., Liimat., Niskanen & A.F.S. Taylor, sp. nov. — MycoBank MB805870; Fig. 4b

Etymology. The name refers to the affinity with C. cyanites and distribution in boreal zone.

Holotype. SWEDEN, Jämtland, Ragunda sn, Ragunda, Böle, at the riverside, in birch forest on rich soil (Betula, Picea), 24 July 1990, Brandrud et al. CFP931 (S). GenBank KF732296.

Pileus 4–10 cm broad, hemispherical to convex, then expanded, distinctly innately fibrillose, bluish grey when young, later pale greyish brown. Lamellae emarginate, almost crowded, greyish blue when young, later brownish violet. Stipe 5–10 cm long, 1–2 cm thick at apex, 2.5–4 cm wide at base, clavate to bulbose, greyish blue. Universal veil pale greyish brown, abundant, forming girdles on stipe. Context violet when young, later in pileus and bulb white to brownish white, marbled hygrophanous, turning vinaceous red on exposure. Odour indistinct. Exsiccata: pileus reddish brown at the centre, greyish brown at the margin, stipe pale bluish greyish, brown at the base.

In MLZ: Spores 9.1–9.8–10.4 × 5.4–5.8–6.3 μm, av. = 9.7–9.9 × 5.6–5.9 μm, Q = 1.58–1.69–1.80, Qav. = 1.66–1.73 (3 specimens, 120 spores, Fig. 4b), amygdaloid, with a blunt apex, fairly finely to moderately verrucose, many with few small golden yellow guttules, slightly dextrinoid. Basidia 36–45 × 7–9 μm (60 basidia), 4-spored, narrowly clavate, with a long, narrow pedicel, with blood red guttules, commonly with yellow foamy contents. Lamellar trama hyphae 4–10 μm wide, colourless to yellowish brown, smooth, with abundant small to large worm-like blood red guttules. Stipe apex hyphae 3–8 μm wide, pale yellowish, smooth, with small to large blood red guttules, the outermost ones c. 3 μm wide, ochraceous brown, mostly not guttulate. Pileipellis: epicutis very weakly gelatinous, uppermost hyphae 4–8(–13) μm wide, ochraceous brown, very thinly spot-like incrusted, not or weakly guttulate with small, golden yellow guttules, lower hyphae up to 10 μm wide, brown, smooth, with abundant small to large worm-like, blood red guttules. Hypoderm not developed.

Ecology & Distribution — In boreal mixed forests of Picea, Betula and Populus in northern Europe. In Scotland also collected with Helianthemum. Fruiting in autumn.

Other specimens examined. FINLAND, Pohjois-Savo, Kuopio, Vehmersalmi, submesic, mixed forest (Betula, Picea, Pinus), in grassy places, 1 Aug. 2003, T. Niskanen et al. 03-112 (H). – GREAT BRITAIN, Scotland, Grampian, Braemar, Morrone Wood, with Helianthemum on rich calcareous soil, 8 Aug. 2010, A. Taylor 2010203 (UPS).

Notes — ITS sequence analysis shows C. boreicyanites as a well-delimited species within clade /Cyanites (Fig. 1). The ITS sequences of the species are identical and differ by more than 30 substitutions and indel positions from those of C. cyanites Fr. and C. violaceorubens Moënne-Locc. & Reumaux. Typical for C. boreicyanites are a pale greyish brown pileus, exsiccata with reddish brown pileus at the centre, greyish brown at the margin, and spores on average 9.7–9.9 × 5.6–5.9 μm. The species is so far only known from the middle boreal zone of Sweden, Finland and Scotland. Cortinarius cyanites has a more southern distribution extending from the hemiboreal zone of Finland and Sweden to France. In addition, the pileus is more bluish. Cortinarius violaceorubens has a more brownish pileus, dark dirty violaceous brown exsiccata, and larger spores (av. 9.9–11.0 × 6.3–6.6 μm). It grows in Picea dominated forests, often on rich soil.

/DIONYSAE s.l

Cortinarius boreidionysae Kytöv., Liimat., Niskanen & Dima, sp. nov. — MycoBank MB805894; Fig. 2e, ​e,44c

Etymology. The name refers to the affinity with C. dionysae and the northern distribution.

Type. FINLAND, Perä-Pohjanmaa, Tervola, Peura, Raemäki, grass-herb-spruce forest with spring-fed depressions, calcareous soil, 11 Sept. 1997, I. Kytövuori 97-1220 (holotype H; isotype NY). GenBank KF732488.

Pileus 5–10 cm broad, hemispherical to convex, then expanded, glutinous, innately fibrillose, mustard brown to cocoa brown with a somewhat silky shining centre and olive yellowish tint on the margin when young. Lamellae emarginate, crowded, violaceous when young, later violet grey to pale brownish grey. Stipe 3–9 cm long, 1–1.8 cm thick at apex, 2–2.5 cm wide at base, with a marginate bulb, when young pale violaceous, becoming yellowish. Universal veil yellow at bulb margin. Context in pileus white, in stipe violaceous, in bulb at first whitish, later brownish yellowish. Odour faintly farinaceous. Exsiccata: pileus uniformly orange brown, sometimes with a faint greyish tint, stipe somewhat paler.

In MLZ: Spores 8.4–9.3–10.2 × 5.2–5.8–6.1 μm, av. = 8.9–9.5 × 5.3–5.8 μm, Q = 1.54–1.67–1.80, Qav. = 1.63–1.67 (5 specimens, 160 spores, Fig. 4c), strongly citriform, strongly beaked, moderately, sharply verrucose, often with small coloured guttules, faintly to moderately dextrinoid. Basidia 23–32 × 7–9 μm (60 basidia), 4-spored, clavate, some with brownish yellow, foamy contents. Lamellar trama hyphae yellow, granulose-guttulate, blood red guttules absent. Stipe apex hyphae yellowish to yellow, smooth, with golden yellow small drops and large, blood red, worm-like guttules abundant in the outermost hyphae. Pileipellis: epicutis strongly gelatinous, uppermost hyphae 3–11 μm wide, mostly densely, spirally incrusted with small to large blood red guttules, lower down with a thick layer of somewhat wider hyphae filled with small to large or very large, worm-like, foamy, blood red guttules. Hypoderm well developed, yellow to red brownish.

Ecology & Distribution — In northern boreal Picea abies dominated forests on calcareous soil, rare. Fruiting in autumn.

Other specimens examined. FINLAND, Oulun Pohjanmaa, Kiiminki, Juuvansydänmaa, S part of Iso Juuvankangas, W of the lake Iso Juuvanjärvi, grass-herb-spruce forest with some Betula, Populus tremula and Pinus, on calcareous soil, 17 Aug. 2007, M. Toivonen & I. Kytövuori 07-245, H6000476 (H); Perä-Pohjanmaa, Tervola, Peura, Raemäki, grass-herb-spruce forest with spring-fed depressions, calcareous soil, 21 Aug. 1998, I. Kytövuori 98-1316, H6035751 (H), 21 Aug. 2007, M. Toivonen & I. Kytövuori 07-491, H6000724 (H); church village, Ala-Kirvesmaa, rich grass-herb-spruce forest with some Betula, Populus and Pinus, on calcareous soil, 24 Aug. 2007, I. Kytövuori 07-768, H6001001 (H); Petäjämaa, Kivimaa, 6 Sept. 2012, I. Kytövuori (H); Tornio, Korkiamaa, Runteli, rich grass-herb-spruce forest with hardwood bushes and some pines, slightly paludified depressions, calcareous soil, 12 Aug. 1995, I. Kytövuori 95-210, H6035732 (H), 10 Sept. 1997, I. Kytövuori 97-1170 (H), 20 Aug. 1998, I. Kytövuori 98-1279, H6035740 (H), 22 Aug. 2007, I. Kytövuori 07-626, H6000859 (H); Kuusamo, Oulanka National Park, Ampumavaara, mesic to damp, herb-rich Picea abies forest, with some Pinus, Populus and Betula, 19 Sept. 2002, T. Niskanen et al. 02-876, H6033134 (H).

Notes — Cortinarius boreidionysae reminds one of C. olivaceodionysae A. Ortega, Vila & Fdez.-Brime, but the latter has paler, olive brown to yellowish brown pileus, somewhat larger, less distinctly and less regularly citriform spores, and a more southern distribution (up to the hemiboreal zone). The ITS sequences of C. boreidionysae are identical and it formed a well-supported clade in our phylogenetic analysis. It belongs to /Dionysae but further relationships were not solved. Based on ITS sequence comparison the closest species is C. olivaceodionysae from which it differs in ITS regions by 9 substitutions and indel positions, although two of them include intragenomic base polymorphisms.

Cortinarius olivaceodionysae has been called C. dionysae Rob. Henry in northern Europe. Cortinarius dionysae, however, has been described from France (Henry 1933) as a species with a distinctly farinaceous smell and collected under Fagus – both characters in contradiction with our material of C. olivaceodionysae. To confirm the identity of C. dionysae we tried to sequence the type material but unfortunately we did not succeed. In our phylogenetic analysis all the taxa with a strong farinaceous smell, C. dionysae sensu Frøslev & Garnica, C. dionysae var. avellanus Rob. Henry ex Bidaud & Carteret and C. palazonianus Vila, A. Ortega & Fdez.-Brime, formed a well-supported subclade (1.00 PP) inside the clade /Dionysae. Of these, C. dionysae sensu Frøslev & Garnica and C. dionysae var. avellanus are potential candidates for the real C. dionysae. The former is collected under Fagus, but is so far only recorded from Germany and the spores are smaller than given in the original description. The latter is found in mixed forest in France but the spores fit well to the description. Further studies with French material are needed to confirm the identity of C. dionysae.

Cortinarius cruentipellis Kytöv., Liimat., Niskanen & Dima, sp. nov. — MycoBank MB805872; Fig. 2f, ​f,44d

Etymology. The name refers to the large and abundant blood red drops/guttules in the pileipellis.

= Cortinarius luteoimmarginatus Rob. Henry sensu Jeppesen et al. (2012).

= Cortinarius polymorphus Rob. Henry s.auct. p.p.

Type. SWEDEN, Öland, Långlöt, Åstad, Nitares hägn, grassy pasture with Corylus and Juniperus, 13 Sept. 2003, I. Kytövuori, K. Liimatainen & T. Niskanen 03-1451 (holotype H; isotype NY). GenBank KF732539.

Pileus 3–7.5 cm broad, hemispherical to convex, then expanded, unevenly wavy, olivaceous yellowish brown at centre, olivaceous to ochraceous yellow towards margin. Lamellae emarginate, almost crowded, pale grey when young, later pale greyish brown. Stipe 3–5 cm long, 0.5–1.4 cm thick at apex, 1.5–2.5 cm wide at base, with a marginate bulb, when young whitish, becoming slightly yellow with age. Universal veil ochraceous yellow at bulb margin. Context whitish. Odour indistinct. Exsiccata: pileus orange brown to dirty red brown at the centre, yellowish brown at the margin, stipe concolorous with the centre.

In MLZ: Spores 9.5–10.4–11.6 × 5.7–6.1–6.6 μm, av. = 10.3–10.5 × 6.0–6.2 μm, Q= 1.59–1.71–1.85, Qav. = 1.69–1.73 (5 specimens, 160 spores, Fig. 4d), often strongly citriform, beaked, moderately, sharply verrucose, fairly faintly to moderately dextrinoid. Basidia 23–34 × 7.5–9.5 μm (50 basidia), 4-spored, clavate, some with yellow foamy contents. Lamellar trama hyphae yellowish, many with greenish yellowish, oily guttules. Blood red guttules absent. Stipe apex hyphae pale yellowish to greyish brownish to reddish brownish, smooth to finely or more strongly incrusted, somewhat granulose-guttulate, large blood red and smaller golden yellow guttules present in the outermost hyphae. Pileipellis: epicutis distinctly gelatinous, uppermost hyphae 4–10 μm wide, thin-walled, finely densely, spirally incrusted, mostly not guttulate, below these a thick layer of smooth to somewhat incrusted hyphae with abundant very long sausage-like guttules with blood red foamy contents. Hypoderm present, yellow brownish.

Ecology & Distribution — Very rare in temperate to hemiboreal grass-herb forests with deciduous trees, mostly Corylus and Quercus, on calcareous soil. In addition, it occurs in half-open deciduous vegetation in wooded pastures and parks. In Northwest Europe it is known in Denmark, Norway, Sweden and Estonia. Fruits in autumn.

Other specimens examined. ESTONIA, Hiiumaa, Pühalepa, Sarve, dryish Corylus forest with Juniperus, Quercus and Populus, stony calcareous soil, 16 Sept. 2001, I. Kytövuori & T. Niskanen (Kytövuori 01-055, H). – NORWAY. Oslo, Bygdøy, Dronginberget south, deciduous forest, 26 Sept. 2004, T.E. Brandrud (Niskanen F04-983) (H7017763); Akershus, Asker, Spirodden, rich calcareous forest, Corylus, 8 Sept. 2011, I. Kytövuori 11-008 (H). – SWEDEN, Öland, Algutsrum, Gråbor Fornborg, rich grass-herb forest with Corylus, Quercus, Populus tremula and Picea, on sedimentary limestone, 30 Sept. 1988, I. Kytövuori 98-2503 (H); Öland, Torslunda, 1.5 km S from Borg, Corylus forest with some Picea, Betula and Quercus, 12 Sept. 2003, T. Niskanen D03-1393 & I. Kytövuori & K. Liimatainen, H7018687 (H).

Notes — Cortinarius cruentipellis is a rather small, olivaceous tinted Phlegmacium of deciduous forests with citriform spores and strongly blood red pileipellis in MLZ. Of the other deciduous forest species with red MLZ reaction in pileus cuticle, C. gracilior (M.M. Moser) M.M. Moser is small and yellowish and has exsiccata with pale stramineous pileus and whitish stipe, C. leonicolor Reumaux (= C. anserinus (Velen.) Rob. Henry s.auct.) has much larger spores, and C. subdecolorans Langl. & Reumaux (= C. multiformium Cons. & Moënne-Locc.) pileipellis hyphae filled by mostly small guttules. Based on the ITS sequence analysis C. cruentipellis is a well-supported species (1.00 PP) belonging to the clade /Dionysae (0.82 PP). It has no close known relatives, and in the ITS regions it differs by more than 30 substitutions and indel positions from the closest species.

Jeppesen et al. (2012) used the name C. luteoimmarginatus Rob. Henry for this species. In the original description of C. luteoimmarginatus (Henry 1939, as C. multiformis var. luteoimmarginatus) the species is mentioned to have a clavate stipe, like the one of C. cliduchus, and this is also illustrated by Henry in Bidaud et al. (2012) whereas our species has a marginate bulb (Jeppesen et al. 2012). In addition, the spores are 11–11.5 × 5.5–6.5 μm, generally longer than those of our species and not overlapping with the average size of the spores (10.4 × 6.1 μm) or with the measurements of Jeppesen et al. (2012) (10–11 × 5.5–6.5 μm). In the original description no type has been designated but later Henry (1985) suggested a type (no. 1006) and a heterotype (no. 1014) for the species. The type of C. luteoimmarginatus, however, was not located in Paris (PC). A collection numbered as 1014 was found, but it was labelled as C. privignofulvus and represented a species of subg. Telamonia. Since the original description of C. luteoimmarginatus does not fit to our species we here describe it as new.

/GLAUCOPODES

Cortinarius subrubrovelatus (Bidaud) Kytöv., Liimat., Niskanen & Dima, comb. nov. — MycoBank MB805884

Basionym. Cortinarius glaucopus var. subrubrovelatus Bidaud, Atlas des Cortinaires 17, 2: 1237. 2008.

Holotype. FRANCE, Ain, Farges, in mixed forest of Populus, Corylus, Fagus and Abies, 1 Nov. 1997, A. Bidaud 97-11-431 (PC). GenBank KF732317.

Other specimens examined. FINLAND, Varsinais-Suomi, Lohja, Jalassaari, herb-rich Picea abies forest with some Corylus avellana, 19 Sept. 2004, I. Kytövuori & T. Niskanen 04-881, H6031330 (H); Paimio, Huso, Kalkkimäki, below the old limestone quarries, moist spruce grass-herb forest with hardwood bushes, 6 Oct. 1985, I. Kytövuori 85-1553 (H). – GERMANY, Bayern, Oberjoch, conifer forest with Picea abies, 4 Oct. 2001, S. Garnica 4498 (TUB 011414), GenBank AY174787.

Additional specimens. ESTONIA, Saare, Lümanda, in garden, with spruce and pine, V. Liiv 2009-10-29 (as ‘C. pini’ TU106663/UDB011262).

Notes — ITS sequence analysis shows C. subrubrovelatus as a well-defined species (1.00 PP) in /Glaucopodes. Typical for C. subrubrovelatus is pale brown to red brown pileus, bluish lamellae, and small, relatively broad spores (7.0–7.9–8.8 × 4.5–5.0–5.4 μm, av. = 7.7–8.0 × 4.8–5.2 μm, Q = 1.44–1.56–1.70, Qav. = 1.52–1.61 (3 specimens, 180 spores)). From its closest relatives C. subfoetens and C. glaucopus it differs by 2 and 4 substitutions and/or indel positions. Cortinarius subfoetens has larger, more fusoid and narrower, and less dextrinoid (8.2–8.9–9.7 × 5.0–5.3–5.4 μm, Q = 1.58–1.69–1.82) spores, and the spores of C. glaucopus are relatively narrower as seen in the Q-values (7.3–8.1–9.1 × 4.5–5.0–5.4 μm, Q = 1.54–1.64–1.82). The latter also has somewhat more reddish brown colours deep in the pileipellis and less incrusted upper hyphae than those of C. subrubrovelatus. The differences in the ITS region between the three species are not large but since all three groups are represented by several collections and also have morphological differences we conclude that C. subrubrovelatus should be treated as a species. Cortinarius subrubrovelatus grows in coniferous forests on calcareous soil and is known from Central Europe and hemiboreal zone of northern Europe.

/ELASTICI & PERCOMES

Cortinarius luteiaureus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805895; Fig. 4e

Etymology. The name refers to the colour of the pileus.

Type. FINLAND, Oulun Pohjanmaa, Kiiminki, Juuvansydänmaa, S part of Iso Juuvankangas, W of the lake Iso Juuvanjärvi, grass-herb Picea abies forest with some Betula, Populus tremula and Pinus, on calcareous soil, 17 Aug. 2007, M. Toivonen & I. Kytövuori 07-247b, H6033617 (holotype H; isotype NY). GenBank KF732568.

Pileus 4–7 cm broad, convex, then plano-convex, with a low and broad umbo, viscid to glutinous, not fibrillose, yellow to brownish yellow. Lamellae emarginate, almost crowded, greyish white, later very pale greyish brown. Stipe 5–10 cm long, 1–1.5 cm thick at apex, 1.5–2 cm wide at base, almost cylindrical with clavate to subbulbous base, white. Universal veil yellow, forming girdles on stipe. Context white. Odour not recorded. Exsiccata: pileus yellow brownish, stipe whitish.

In MLZ: Spores 9.7–10.6–11.6 × 5.7–6.1–6.6 μm, Q = 1.62–1.74–1.92 (1 specimen, 60 spores, Fig. 4e), narrowly amygdaloid, with rounded apex, moderate to fairly strongly verrucose, moderately dextrinoid, often with dark red brown angular granules in the spore. Basidia 32–44 × 8–10 μm (20 basidia), 4-spored, clavate, sand brown, with fairly small granules and chips. Lamellar trama hyphae with moderate to very large sand brown to red brown granules. Stipe apex hyphae sand brown, densely granulose, outermost ones more orange or reddish, not granulose. Pileipellis: epicutis strongly gelatinous, uppermost hyphae 2–4 μm wide, ochraceous yellow to ochraceous brown, mostly not granulose, lower down 4–10 μm wide, full of small to very large dark red brown granules. Hypoderm absent.

Ecology & Distribution — In coniferous forest (Picea) on calcareous soil. Known from northern Finland, Oulun Pohjanmaa. No sequences of this species exist in public databases.

Notes — Cortinarius luteiaureus resembles somewhat a species which we assume could be called C. rhizophorus Bidaud & Cons. but we have not studied the type material of the species yet. Cortinarius cf. rhizophorus is larger, somewhat paler, has larger spores (10.2–11.3–12.5 × 6.1–6.5–7.0 μm) and occurs in temperate to hemiboreal broad-leaved and coniferous forests. Cortinarius varius (Schaeff.: Fr.) Fr. is also somewhat similar but has blue lamellae, stout, clavate stipe, wider spores, and hypoderm in the pileipellis. Cortinarius luteiaureus differs in ITS regions by more than 30 substitutions and indel positions from the closest species found with the BLAST. Based on our phylogenetic analysis it belongs to the large /Elastici & Percomes clade.

/INFRACTI

Cortinarius infractiflavus (M.M. Moser) Kytöv., Niskanen, Liimat., Bojantchev & Ammirati, stat. nov. & nom. nov. — MycoBank MB805887

Etymology. The name refers to the affinity to C. infractus and the yellowish pileus.

Basionym. Cortinarius infractus var. flavus M.M. Moser, Mycotaxon 72: 313. 1999.

Type. USA, Wyoming, Shoshone Natl. Forest, prope Brooks Lake Lodge, in subalpine coniferous forest (Picea engelmanii, Abies lasiocarpa), 12 Aug. 1997, M. Moser 97/169 (holotype IB). GenBank KF732327.

Other specimens examined. BULGARIA, Pirin Mt, Bansko locality, 1600 m asl, under Picea abies, 7 Oct. 2009, D. Bojantchev, DBB19634 (UC). – FINLAND, Sompion Lappi, Pelkosenniemi, Suvanto, Kalkkivaara, half-open dry Pinus sylvestris forest with Picea, Juniperus, Betula, Populus and Alnus, dolomite rock, 26 Aug. 1992, I. Kytövuori 92-1109 (H).

Additional specimen. CANADA, British Columbia SMI286, GenBank FJ039612 (UBC).

Notes — Cortinarius infractiflavus formed a well-supported clade (1.00 PP) in our phylogenetic analysis. It belongs to sect. Infracti and differs from the sister species C. infractus by 9 substitutions and indel positions in ITS regions. Morphologically, it can be distinguished from C. infractus by the yellowish colours of the pileus, paler gills, a nearly mild taste and larger spores (7.5–8.2–9.1 × 6.3–6.7–7.0 μm, Q = 1.16–1.23–1.30; C. infractus 7.3–8.0–8.8 × 5.7–6.1–6.6 μm, Q= 1.23–1.30–1.38). Cortinarius infractiflavus grows in boreal and mountainous conifer forests (Picea, Abies) and is widespread occurring from Wyoming, western USA to Finland and Bulgaria. Full description of the species can be found from Moser & Ammirati (1999).

/MULTIFORMES

Cortinarius caesiolamellatus(Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F.S. Taylor, comb. nov. — MycoBank MB805885

Basionym. Cortinarius rufoallutus var. caesiolamellatus Bidaud, Atlas des Cortinaires 16: 1095. 2006.

Type. FRANCE, Ain, Bugey, col de Bérentin, in young Picea abies plantation, on calcareous soil, 3 Oct. 1993, A. Bidaud PML4905 (holotype PC). GenBank KF732414.

= Cortinarius multiformis var. caesiophyllus Moënne-Locc., Atlas des Cortinaires 16: 1095. 2006. — Type. P. Moënne-Loccoz PML882 (holotype PC), France, Savoie, Arith, under Picea on calcareous soil, 3 June 1988. Gen-Bank KF732351.

Illustrations — Bidaud et al. (2006: pl. 581, 587).

Pileus 4–8 cm broad, hemispherical to plano-convex, sometimes radially rugulose towards the margin, viscid, red brown to ochraceous brown, rarely bluish brown, becoming paler ochraceous brown with age, often bi-coloured, outer part hygrophanous and darker (grey) brown. Lamellae emarginate, almost crowded, pale grey with a bluish tint when young, later pale brown. Stipe 4–8 cm long, 0.7–1.7 cm thick at apex, cylindrical, slender, 1.5–2.5 cm wide at base, mostly bulbous but the bulb often fairly small, fibrillose (not glossy like C. multiformis), whitish but often with a bluish tint at apex when very young, often becoming pale brown with age. Universal veil white, sparse, sometimes viscid (at bulb margin). Context in the pileus fairly thin, whitish, brownish below the pileus cuticle, in the stipe with a bluish tint or not when young. Odour faint to distinct of honey in the context of the stipe base. Exsiccata: pileus greyish to reddish brown, stipe very pale.

In MLZ: Spores 8.2–9.3–10.7 × 5.0–5.6–6.3 μm, av. = 8.5–9.9 × 5.2–5.9 μm, Q = 1.51–1.65–1.85, Qav. = 1.57–1.74 (7 specimens, 240 spores), ovoid-amygdaloid to narrowly ovoid-ellipsoid, with a fairly long, blunt, tapering apex, moderately to strongly verrucose, warts wide, anastomosing, slightly to fairly strongly dextrinoid. Basidia 25–34 × 7.5–9 μm (100 basidia), 4-spored, clavate, mostly with fairly wide base. Many basidia and subhymenium with yellowish, granulose/guttulate contents. Pileipellis: epicutis with a fairly thick, gelatinous layer with some 2–3 μm wide, erectentangled, smooth, obscure, colourless hyphae. Hypoderm present, fairly thin-walled, with pale brownish parietal pigment, some small brown spots present, large yellow to yellow brown spots absent, few spirally incrusted hyphae present deep in the pileipellis.

Ecology & Distribution — In montane to middle boreal, mesic coniferous forests, often on rich soil. In Europe apparently mainly/only under Picea, but in USA also found under Pinus. Known from Central and northern Europe, and Washington, USA, and considered occasional. The fruiting period is very long, once collected in June (France) and once in November (USA).

Other specimens examined. ESTONIA, Saaremaa, Mustjala, Võhma, dryish pine heath forest with some spruce, on calcareous soil, 16 Sept. 1993, I. Kytövuori 93-1336 (H). – FINLAND, Uusimaa, Kunnarla, Myllyoja, grass-herb forest with some pines and hardwood trees, 22 Sept. 1994, I. Kytövuori 94-852 (H); Pohjois-Savo, Kuopio, Vehmersalmi, Putroniemi, Roikanselän ranta-Mäkijärvi E, under Picea, 30 years old forest, 26 Sept. 2009, J. Ruotsalainen 8103F, H6011464 (H). – FRANCE, Oyonnax SE, Giron, Forêt de Champfromier, rich conifer forest with Abies alba, Picea abies and Fagus, 27 Oct. 1994, P. & I. Kytövuori 94-1897 (H); Savoie, Arith, under Picea on calcareous soil, 3 June 1988, P. Moënne-Loccoz PML882 (PC). – GERMANY, Baden-Württemberg, Thölendorf, Picea, 11 Oct. 1998, UL 98/122 (TUB 011841), GenBank AY669531 as ‘C. allutus’; Schwaben, Ehingen a.d. Donau, Kohlhau, Picea plantation on calcareous soil, 28 Sept. 2010, G. Schmidt-Stohn & T.E. Brandrud, TEB 428-10 (O, TUB). – NORWAY, Oslo, Steinbruvannet-Røverkollen, Picea forest of somewhat richer, low-herb type, 4 Sept. 2011, T.E. Brandrud 687-11 (O). – SWEDEN, Uppland, Uppsala, Nåsten, mixed forest, 27 Aug. 2005, A. Taylor 2005085, UNITE No. UDB002202 as ‘C. allutus’ (UPS); Vänge, Fiby urskog, virgin spruce forest with Pinus, Betula, Populus tremula, Colylus, Alnus glutinosa, etc., with fairly rich depressions, decaying logs very abundant, 8 Sept. 1994, P. & I. Kytövuori 94-309 (H). – USA, Washington, Grays Harbor Co., Ocean Shore State Park, under Pinus on sandy soil, 11 Nov. 2009, J.F. Ammirati & T. Niskanen 09-201 (H).

Notes — Cortinarius caesiolamellatus and C. caesiophylloides are easily distinguished from the related species in /Multiformes by their initially bluish tinged lamellae and often also stipe apex. Based on material seen so far and available descriptions, these two blue-gilled species are hardly distinguishable macromorphologically, but C. caesiolamellatus differs in slightly larger and more strongly verrucose spores. Furthermore, there seems to be a geographical differentiation in Europe; C. caesiolamellatus apparently being mainly a Central European species, whereas C. caesiophylloides is hitherto found only in northern Europe. If bluish tints have gone, C. caesiolamellatus can also be confused with conifer associated C. rufoallutus Rob. Henry ex Bidaud & Reumaux, C. multiformis Fr. and C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S Taylor & Sesli. The latter two, however, have less strongly ornamented spores and lack the yellow to yellow brown large spots deep in the pileipellis. Cortinarius rufoallutus differs from C. caesiolamellatus by stouter appearance, small amygdaloid-fusoid, finely verrucose spores and abundant strongly spirally incrusted red brown hyphae deep in the pileipellis.

In our phylogenetic analysis C. caesiolamellatus forms a well-supported group (1.00 PP) and belongs together with C. caesiophylloides and C. pallidirimosus in a strongly supported (1.00 PP) subclade in clade /Multiformes (1.00 PP). The species was originally described twice in Bidaud et al. (2006), as a variety of both C. rufoallutus (var. caesiolamellatus) and C. multiformis (var. caesiophyllus). Based on the molecular and morphological data neither of those relationships is supported. Cortinarius caesiolamellatus clearly represents a distinct species and is here combined at species level.

Cortinarius caesiophylloidesKytöv., Liimat., Niskanen, Brandrud & Frøslev, sp. nov. — MycoBank MB805873; Fig. 2g, ​g,55a

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Fig. 5

Spores of: a. C. caesiophylloides TN05-016; b. C. melleicarneus IK01-053; c. C. pallidirimosus IK95-585; d. C.talimultiformis AT2004096, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 μm.

Etymology. The name refers to the bluish tints in lamellae and affinity with C. caesiolamellatus (= Cortinarius multiformis var. caesiophyllus).

Type. FINLAND, Etelä-Savo, Joutsa, Koivuranta, W of Rakkolanselkä, fairly young, mesic to damp, Picea abies-dominated forest with some Betula and Pinus sylvestris, 30 Aug. 2005, T. Niskanen et al. 05-016, H6029792 (holotype H; isotype NY). GenBank KF732572.

Pileus 4–8 cm broad, hemispherical to plano-convex, viscid, apricot to redbrown-coloured, with hygrophanous streaks or outer zone, hygrophanous areas somewhat darker umber to less vivid (grey) brown, becoming paler ochraceous brown to ochraceous yellow with age, sometimes almost whitish ochre at centre. Lamellae emarginate, almost crowded, pale grey with a bluish tint when young, later pale brown. Stipe 5–11 cm long, 1–1.5 cm thick at apex, 1.5–3.5 cm wide at base, with a more or less distinct marginate bulb, whitish but usually with a bluish tint at apex when very young, often becoming pale brown with age, but not as pronounced as with C. multiformis. Universal veil white, sparse to hardly visible, at bulb margin. Context white, slightly bluish at the apex of the stipe when young. Odour faint to distinct of honey in the context of the stipe base. Exsiccata: pileus pale dirty brown, stipe somewhat lighter.

In MLZ: Spores 8.6–9.7–10.9 × 5.2–5.8–6.3 μm, av. = 9.0–10.2 × 5.5–6.0 μm, Q = 1.54–1.68–1.85, Qav. = 1.63–1.73 (4 specimens, 240 spores, Fig. 5a), amygdaloid, with fairly narrow apex, sometimes almost ellipsoid, finely to moderately verrucose, warts often rounded-confluent, slightly to fairly strongly dextrinoid; with a bit narrower apex, thinner wall and finer verrucosity than in C. multiformis. Basidia 24–37 × 7–9 μm (60 basidia), 4-spored, clavate, almost colourless. Lamellar trama hyphae very pale yellowish, smooth, concolorous guttulate. Stipe apex hyphae very pale yellow, smooth, somewhat concolorous guttulate. Pileipellis: epicutis with a clear gelatinous layer with sparse, erect-entangled, 2–3 μm wide, smooth, colourless and very obscurely seen hyphae. Hypoderm present, hyphae fairly thin-walled, with pale yellowish brown amber-like parietal pigment, small brown encrust-spots present, large yellow brown spots absent, a few spirally incrusted (zebra-striped) hyphae seen deep in the cuticle.

Ecology & Distribution — In mesic coniferous forests, presumably with Picea, mainly in richer low-herb types, sometimes on calcareous soil. Known only from Fennoscandia and considered occasional. Fruits from August to September.

Other specimens examined. FINLAND, Kainuu, Paltamo, Oikarilankylä, Kivesvaara, old mesic spruce forest with some Betula, Pinus and Populus tremula, 11 Sept. 2008, I. Kytövuori 08-1554, H6032621 (H); Sompion Lappi, Pelkosenniemi, Suvanto, Kalkkivaara, steep SW slope, dry pine forest with Juniperus, Betula, Populus, Alnus incana, 26 Aug. 1992, I. Kytövuori 92-3044, H6032620 (H). – NORWAY, Nord-Trøndelag, Stjørdal, Beistadvollen, calcareous spruce forest (on karst surfaces), 13 Aug. 2009, T.E. Brandrud 277-09 (O).

Notes — Cortinarius caesiophylloides is a typical member of sect. Multiformes. Together with C. caesiolamellatus they are the only known species of the section with bluish tinted lamellae when young. Cortinarius caesiolamellatus differs from C. caesiophylloides by more strongly ornamented spores. When bluish tints are absent they can be confused with C. multiformis and C. talimultiformis, but in the latter two there are abundant, yellow to yellow-brown, large spots deep in the pileipellis. Those are practically lacking from C. caesiophylloides and C. caesiolamellatus. In C. multiformis the honey smell is either lacking or is very weak, and the stipe becomes strong brass brown with age.

Cortinarius caesiophylloides is strongly supported (1.00 PP) in our phylogenetic analysis. It belongs to /Multiformes (1.00 PP) and further, in a well-supported (1.00 PP) subclade with C. caesiolamellatus and C. pallidirimosus. The ITS sequences of C. caesiophylloides have one base polymorphism and the maximum pairwise distance is zero. The difference to C. pallidirimosus is 11 substitutions and indel positions.

Cortinarius melleicarneus Kytöv., Liimat., Niskanen & Brandrud, sp. nov. — MycoBank MB805874; Fig. 5b, ​b,66a