Two new species of Cortinarius, subgenus Telamonia, sections Colymbadini and Uracei, from Europe Bálint Dima, Kare Liimatainen, Tuula Niskanen, Ilkka Kytövuori & Dimitar Bojantchev Mycological Progress ISSN 1617-416X Volume 13 Number 3 Mycol Progress (2014) 13:867-879 DOI 10.1007/s11557-014-0970-6 1 23 Your article is protected by copyright and all rights are held exclusively by German Mycological Society and Springer-Verlag Berlin Heidelberg. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Mycol Progress (2014) 13:867–879 DOI 10.1007/s11557-014-0970-6 ORIGINAL ARTICLE Two new species of Cortinarius, subgenus Telamonia, sections Colymbadini and Uracei, from Europe Bálint Dima & Kare Liimatainen & Tuula Niskanen & Ilkka Kytövuori & Dimitar Bojantchev Received: 18 December 2013 / Revised: 11 February 2014 / Accepted: 14 February 2014 / Published online: 12 March 2014 # German Mycological Society and Springer-Verlag Berlin Heidelberg 2014 Abstract 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. B. Dima (*) : K. Liimatainen : T. Niskanen Department of Biosciences, Plant Biology, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland e-mail: cortinarius1@gmail.com K. Liimatainen e-mail: kare.liimatainen@helsinki.fi T. Niskanen e-mail: tuula.niskanen@cortinarius.fi I. Kytövuori Botanical Museum, University of Helsinki, P.O Box 7, 00014 Helsinki, Finland e-mail: ilkka.kytovuori@funga.fi D. Bojantchev 345 Shipwatch Lane, Hercules, CA 94547, USA e-mail: dimitar@pontix.com URL: www.MushroomHobby.com Keywords Agaricales . DNA . ITS . Taxonomy . UV fluorescence Introduction Molecular studies based on the ITS1 and ITS2 regions of the rDNA, in combination with morphological and ecological data, have been applied successfully for species delimitation in the Agaricales including the large genus Cortinarius (e.g., Eberhardt and Beker 2010; Garnica et al. 2011; Frøslev et al. 2007; Niskanen et al. 2011; 2013a; Suárez-Santiago et al. 2009). Traditional subgeneric classification of Cortinarius based largely on morphological characters (e.g., Bidaud et al. 1994; Consiglio et al. 2007; Consiglio 2012) is not supported by recent molecular studies (e.g., Garnica et al. 2005; Peintner et al. 2004) and should be mostly discarded. A great number of Cortinarius species have been found to be taxonomic synonyms while many others, even common ones, remain undescribed (Niskanen et al. 2008, 2013a, b). The monophyletic subgenus Telamonia s. str. (Garnica et al. 2005; Niskanen 2008; Peintner et al. 2004.) is comprised of a very large number of species and is taxonomically the most difficult group of Cortinarii. Subgenus Telamonia, section Uracei contains species with dark brown basidiomata which become blackish on drying, amygdaloid spores that are strongly verrucose and strongly dextrinoid, and occurrence on rich to calcareous soils. The section was first introduced by Kühner and Romagnesi (1953), later validated by Melot (1990) and later placed in subgenus Telamonia (Brandrud et al. 1989; Consiglio 2012). Bidaud et al. (2012) placed the section in subgenus Hydrocybe (= Telamonia p. p.). The number of species included in section Uracei ranges from three (Niskanen et al. 2008) to over 60 taxa (Bidaud et al. 2012). Author's personal copy 868 Section Colymbadini Melot (1990) is generally characterized by their early appearance (spring to summer), the olivaceous, metallic tinges on the basidiomata, and the strikingly yellow UV fluorescence. Usually only the type species, C. colymbadinus, is included (e.g., Brandrud et al. 1998; Consiglio 2012), but, depending on different authors, also C. isabellinus and/or C. zinziberatus (e.g., Moser 1978) are placed here. The presence of positive UV fluorescence of C. colymbadinus (and C. isabellinus) led most of the authors to place the section in subgenus Leprocybe or Cortinarius (e.g., Bidaud et al. 2005; Brandrud et al. 1989; Consiglio 2012), where other UV positive species are found (e.g., section Leprocybe), but with completely different morphology. Phylogenetic studies indicate that the species in sections Uracei and Colymbadini are closely related and belong to the subgenus Telamonia s. str. (Niskanen 2008; Niskanen et al. 2013c) forming a still unresolved complex together with section Cinnabarini (Ammirati et al. 2013). Here, we describe two new species belonging to sections Uracei and Colymbadini based on morphological and molecular data and make one new combination. Also neotypes are selected for C. uraceus and C. colymbadinus to resolve the taxonomic and nomenclatural problems within these groups. Materials and methods A total of 47 specimens from Finland, Sweden, Hungary, Bulgaria, Germany, Italy, Switzerland, Austria, and France were studied, including the type specimens of C. fulvoisabellinus, C. pardinipes, C. pseudouraceus, C. rigidipes, C. umbonatoides, C. viridipes and Hydrocybe nolaneiformis (see Table 1). Specimens are deposited in the public herbaria BP, H, S, IB, NY, PC, PRM, and UC. Personal collections of László Albert (AL), Bálint Dima (DB), Dimitar Bojantchev (DBB) and Karl Soop (KS) were also used. Herbarium acronyms follow Thiers (continuously updated). The abbreviation of IK refers to Ilkka Kytövuori. Author abbreviations of the species are based on the current version of MycoBank (Robert et al. 2005). DNA was extracted from dried material (a piece of lamella) with the NucleoSpin Plant kit (Macherey-Nagel, Düren, Germany). Primers ITS 1F and ITS 4 (White et al. 1990; Gardes and Bruns 1993) were used to amplify ITS regions and the same primer pairs were used in direct sequencing. For problematic material the primer combinations ITS 1F/ITS 2 and ITS 3/ITS 4 were also used. PCR amplification and sequencing followed Niskanen et al. (2009). Sequences were assembled and edited with Sequencher 4.1 (Gene Codes, Ann Arbor, Mich., USA). BLAST queries of the public databases (GenBank: http://www.ncbi.nlm.nih.gov/and UNITE: http://unite.ut.ee/), were used to check for identical or similar sequences. For the phylogenetic analysis, ITS Mycol Progress (2014) 13:867–879 sequences of the studied species and morphologically similar species were included. Two species from subgenus Phlegmacium were chosen as outgroup taxa, as in Niskanen et al. (2011). The alignment of 81 ITS sequences was produced with the program MUSCLE (Edgar 2004) under default settings. The alignment comprised 683 nucleotides (including gaps). The alignment is available at TreeBASE under S15311 (http:// www.treebase.org/treebase-web/home.html). Bayesian inference (BI) was performed with MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). The best substitution model for the alignment was estimated by both the Akaike information criterion and the Bayesian information criterion with jModelTest version 0.1.1 (Posada 2008). A GTR model, including a gamma shape parameter, was chosen. Two independent runs with four chains in each were performed for 4,000,000 generations sampling every 100th generation. All trees sampled before stationarity were discarded with a 25 % safety margin (burn-in of 10,000 trees [1,000,000 generations]). Sampled trees from both runs were combined in a 50 % majority rule consensus phylogram and posterior probabilities (PP) were calculated. The analysis was run with computer clusters of the CSC, IT Center for Science, Espoo, Finland. Macroscopic characteristics were observed from fresh materials collected by the authors, including specimens in all stages of development. Colour codes from the Munsell soil colour charts (Munsell 2009) were used to describe the colours of exsiccatae. Colour photographs of fresh specimens are provided for all species except C. colymbadinus. The species presented here have mostly dark brownish to blackish exsiccatae, but these colours can develop in different ways. Based on our observations, rotting basidiomes or those that are not dried properly can develop secondary blackening of the pileus centre. Basidiomes that are partially dried in the field often remain much lighter in colour, when dried, than those dried in prime fresh condition. Microscopic characteristics were observed from dried materials mounted in Melzer’s reagent (MLZ, 0.5 g (1.2 %) iodine, 1.5 g (3.6 %) potassium iodide, 20 cm3 chloralhydrate, 20 cm3 distilled water) and measurements were made on a monitor with the aid of a Zeiss Laboval 2/I/C microscope using 100× oil immersion lens, and connected to a ALPHA DCM 130E camera and equipped with ScopePhoto 3.0.4 software. Twenty spores were measured (excluding apiculus and ornamentation) from one basidioma per each collection (70–120 spores per species), from the cortina remnants or top of the stipe. Length and width were measured from the same spore, and the length/width ratios (Q-value) were calculated for individual spores. Spore drawings are provided to compare the shape and ornamentation of the spores between the species. The hyphae of the lamellar trama and basidia (ten basidia per collection, 30–50 basidia per species) were examined and Author's personal copy Mycol Progress (2014) 13:867–879 869 Table 1 Cortinarius sequences produced in this study Species Voucher Locality Herb. GenBank accession number C. colymbadinus DB2535 Hungary, Vas, Szalafő BP KJ206482 C. colymbadinus DB4571 Austria, Süd-Tirol, Iselsberg BP KJ206483 C. colymbadinus IK99-295 Finland, Pep, Ylitornio H KJ206484 C. fulvoisabellinus Rob. Henry (holotype) RH1891 France PC KJ206485 C. nolaneiformis (Velen.) Dima, Niskanen & Liimat. (holotype of Hydrocybe nolaneiformis) C. nolaneiformis 857042 PRM KJ206486 DB886 Czech Republic, Central Bohemia, SE of Praha Hungary, Vas, Szalafő BP KJ206487 C. nolaneiformis DB1611 Hungary, Vas, Szalafő BP KJ206488 C. nolaneiformis C. nolaneiformis DB2287 DB4056 Hungary, Vas, Szalafő Hungary, BAZ, Bükkzsérc BP BP KJ206489 KJ206490 C. nolaneiformis KS-CO699 Sweden, Srm, Nacka KS KJ206491 C. pardinipes Romagn. (holotype) 66-201 France PC KJ206492 C. pardinipes TN03-239 Finland, KP, Veteli H KJ206493 C. pardinipes IK00-030 Finland, PH, Virrat H KJ206494 C. pardinipes TN05-073 Finland, EH, Ruovesi H KJ206495 C. pardinipes TN04-534 Finland, PeP, Tornio H KJ206496 C. pseudouraceus Bidaud & Reumaux (holotype) C. pseudouraceus AB01-10-106 IK94-1364 France, Ain, Thézillieu Sweden, Ög, Ödenshög PC H KJ206497 KJ206498 C. pseudouraceus IK98-195 Finland, PeP, Tornio H KJ206499 C. pseudouraceus IK01-049 Finland, V, Förby H KJ206500 C. pseudouraceus IK01-054 Finland, V, Lohja H KJ206501 C. pseudouraceus TN03-1567 Sweden, Öl, Ismantorp H KJ206502 C. pseudouraceus TN04-828 Finland, U, Porvoo H KJ206503 C. rigidipes M.M. Moser (holotype) MM1962/0062 Switzerl, L, Doppelschwand IB KJ206504 C. rigidipes C. rigidipes IK94-1830 IK95-1873 France, Ain, Oyonnax Germany, BW, Heiligenbronn H H KJ206505 KJ206506 C. umbonatoides Moënne-Locc. & Reumaux (holotype) PML2196 France, HS, Avernioz PC KJ206507 C. uraceomajalis Dima, Liimat., Niskanen & Bojantchev (holotype) C. uraceomajalis DB1623 Hungary, Heves, Recsk BP KJ206508 DB1624 Hungary, Heves, Recsk BP KJ206509 C. uraceomajalis DB2283 Hungary, Nógrád, Mátranovák BP KJ206510 C. uraceomajalis DB2291 Hungary, GyMS, Sopron BP KJ206511 C. uraceomajalis DB2990 Hungary, KE, Kesztölc BP KJ206512 C. uraceomajalis DB2303 Hungary, Ve, Bakonygyepes BP KJ206513 C. uraceomajalis DBB51403 Bulgaria, Pavel Bania, Sredna Gora UC KJ206514 C. uraceonemoralis Niskanen, Liimat., Dima, Kytöv., Bojantchev & H. Lindstr. (holotype) C. uraceonemoralis TN04-1116 Italy, Sardinia, Gavoi H KJ206515 CFP1478 Italy, Süd-Tirol, Kaltern S KJ206516 C. uraceonemoralis C. uraceonemoralis DB4322 IK98-1838 Hungary, Budapest Sweden, Ög, Ödenshög BP H KJ206517 KJ206518 C. uraceonemoralis TN03-1264 Sweden, Öl, Vickleby H KJ206519 C. uraceonemoralis Hungary, Vas, Szalafő BP KJ206520 C. uraceonemoralis ORS-ERDO 99-15-1 DBB56726 Bulgaria, Pavel Bania, Sredna Gora UC KJ206521 C. uraceus Fr. (neotype) TN04-872 Finland, V, Lohja H KJ206522 C. uraceus CFP730 Sweden, Jmt, Ragunda S KJ206523 C. uraceus IK97-1519 Finland, Kn, Puolanka H KJ206524 C. uraceus IK98-1607 Finland, V, Lohja H KJ206525 C. uraceus C. viridipes M.M. Moser ex M.M. Moser (holotype) IK01-002 MM1949/0030 Finland, V, Västenfjärd Austria, Tirol, Mühlbachtal H IB KJ206526 KJ206527 For acronyms of biogeographical provinces of Finland and Sweden, see Knudsen and Vesterholt (2008): pp. 29–35. Other abbreviations are as follows: BAZ Borsod-Abaúj-Zemplén County (Hungary), BW Baden-Württemberg (Germany), GyMS Győr-Moson-Sopron County (Hungary), HS Haute-Savoie (France), KE Komárom-Esztergom County (Hungary), L Luzern (Switzerland), Switzerl Switzerland, Ve Veszprém County (Hungary) Author's personal copy 870 measured from the pieces of lamellae. In addition, the pileipellis structure was studied from both radial freehand sections and scalps from midway to the pileus centre. The measurements were made from scalp preparations. The microscopic descriptions are almost exclusively based on ITSsequenced material. For checking the UV fluorescence, long wave UVat wavelength 366 nm was used on dried specimens. Results of phylogenetic analyses The two new species, C. uraceomajalis and C. uraceonemoralis, were strongly supported in our phylogenetic analysis (PP 0.96 and 1.00). Cortinarius uraceomajalis was placed in the section Colymbadini together with C. colymbadinus and C. nolaneiformis but with a low support (PP 0.50). Cortinarius uraceonemoralis belongs to section Uracei and the relationship with the closest sister species C. rigidipes and C. uraceus was well supported (PP 0.96). The intraspecific variation in both species is 0 to 1 substitutions or indel positions, and they differ from their closest relatives by more than 12 substitutions and indel positions. The sequencing of the end of the ITS2 region of C. viridipes type material did not succeed and the partial sequence was left out of the analysis. The successfully sequenced part, however, was identical with the ITS sequence from the neotype of C. uraceus, and, therefore, they are considered the same species. Taxonomic descriptions Cortinarius section Colymbadini Melot Type species: Cortinarius colymbadinus Fr. In our phylogenetic analysis, three European species, C. colymbadinus, C. nolaneiformis and C. uraceomajalis formed a clade, but with a low support. The species in this group, however, share one characteristic, which is lacking from the other species of sections Uracei and Cinnabarini s. l., they all have a positive (yellow) UV reaction. In addition, all three species fruit in spring and early summer and have rather small (< 9.5×4.5–5.5 μm), amygdaloid, and moderately to strongly verrucose spores. Cortinarius colymbadinus Fr., Epicr. syst. mycol.: 289 (1838) (Fig. 2a) Cortinarius isabellinus (Batsch) Fr. s. Brandrud et al. (1998) Cortinarius zinziberatus (Scop. : Fr.) Fr. s. Moser (1978). Illustration: Brandrud et al. (1998), D55. Typus: Sweden, Jämtland, Ragunda, Kullstabodarna, in rich spruce forest, 16 August 1992, T. E. Brandrud, H. Lindström, H. Marklund & S. Muskos, Neotype CFP1130 Mycol Progress (2014) 13:867–879 (S, neotype designated here). Mycobank No. MBT177200; GenBank No. JX127302. Pileus 20–70 mm, hemispherical, then low convex to almost plane, with a low umbo, diffracted towards margin, finely fibrillose when young, hygrophanous, chocolate brown to olivaceous yellow brown, with a characteristic, metallic, olivaceous yellow shine, which persists on drying. Lamellae medium spaced, emarginate, yellow-brown to cinnamon brown when young, later darker, with yellow, often uneven edge. Stipe 40–100 mm long, 4–10 mm thick at apex, cylindrical to somewhat clavate, silky-fibrillose, slightly glossy, greyish yellow to brown, towards base dark brown, base with yellow mycelial felt, cortina sparse. Universal veil greenish yellow, sparse, initially sock-like, then forming a thin ring-like or indistinct zone. Context in pileus thin, pale brownish, in stipe often hollow, with hygrophanous streaks, brown, darker towards base. Basal mycelium greenish yellow. Odour indistinct or slightly raphanoid. Exsiccata pileus dark brown (10YR 3/3), dark yellowish brown (10YR 3/4, 10YR 4/4) to very dark brown (10YR 2/2), becoming very dark greyish brown (10YR 3/2) to black (10YR 2/1) towards centre. Stipe olive brown (2.5Y 4/3, 2.5Y 4/4), base of stipe (basal mycelium?) yellow (2.5Y 8/8). UV observations: positive almost on the whole basidiomata, especially the stipe and the edge of the lamellae (chrome yellow), cap paler yellowish. Spores: 7.5–9.5×4.5–5.5 μm, Q=(1.5)1.6–1.8(1.9), av. 8.1–8.7×4.8–5.3 μm, Q(av.)=1.63–1.79, amygdaloid, strongly to very strongly verrucose, moderately to fairly strongly dextrinoid. Basidia 4-spored, 22–31×7–9 μm, clavate, olivaceous yellow in MLZ. Lamellar edge fertile, with numerous, clavate sterile cells, concolorous with the basidia. Lamellar trama hyphae olivaceous yellow in MLZ, finely encrusted (rarely with spot-like incrustation). Pileipellis duplex, in overall view pale yellowish brown, epicutis thin to moderately thick, hyphae 3–8 μm wide, hyaline or with yellowish brown content, slightly encrusted or smooth. Hypoderm weakly developed, elements 10–22 μm wide, hyaline, smooth. Clamp connections present. ITS-regions (including 5.8S region): Maximum pairwise distance of the sequences is 0 (includes two polymorphic sites). Cortinarius colymbadinus differs from C. uraceomajalis by 13 substitutions and indel positions and from C. nolaneiformis by 21 substitutions and indel positions. Ecology and distribution: In mesic Picea forests, mostly on rich soil. Producing basidiomes from summer to autumn, often fruiting early in the season. Widely distributed and occasional in Europe. Sequenced collections are from northern Europe (Finland and Sweden) and central Europe (Austria, Hungary). Collection (DB2535) represents the first record in Hungary. Differential diagnosis: Cortinarius colymbadinus is a fairly slender species with characteristic olivaceous yellow, metallic tints on the basidiomata, especially on drying. It grows in Author's personal copy Mycol Progress (2014) 13:867–879 coniferous forests and has strong, chrome yellow UV fluorescence on the stipe and lamellae edge. The early appearance of the species is also characteristic. Cortinarius colymbadinus is a widely used name, at least in the Nordic countries for the species treated here. Also, the description by Fries (1838) fits the species. Therefore, we propose the collection depicted and described in Brandrud et al. (1998) as a neotype to stabilize the interpretation and nomenclature of this species. Cortinarius isabellinus (Batsch) Fr. is often synonymised with C. colymbadinus (e.g. Brandrud et al. 1998). The original plate of Agaricus isabellinus Batsch, however, shows a fungus with bluish tint in the context, reminiscent to that of C. biformis (Batsch 1783). This fungus does not resemble C. colymbadinus. Cortinarius zinziberatus (Scop. : Fr.) Fr. is known as another early species, and the name has been applied to C. colymbadinus in Central Europe (e.g., Moser 1978). This fact was verified in our phylogenetic studies, since two collections identified as C. zinziberatus from Austria (one from Moser, see UDB001095) had identical ITS sequences with the proposed neotype of C. colymbadinus (Fig. 1.). Cortinarius zinziberatus would be an older name for this species, but the original description of Agaricus zinziberatus Scop. includes a small-sized species (pileus only up to 2.7 cm) with bright yellow pileus and lamellae, and the smell was reported similar to that of ginger (Scopoli 1772). None of these characters are in correspondence with C. colymbadinus treated here. Fries (1821) sanctioned the name Agaricus zinziberatus and kept the original idea of the bright yellow colours and smell. Further contradiction is the habitat which was neither mentioned by Scopoli (1772) in the original description, nor by Fries (1821). Later on Fries (1838), placed both species in Cortinarius, and noted the high degree of similarity with C. colymbadinus, and he published it as a rare species of frondose woods. Based on our data, however, C. colymbadinus occurs exclusively in coniferous forests. According to the above mentioned contradictions, we are in agreement with Melot (1986) that Agaricus zinziberatus is impossible to interpret, and therefore cannot be used for C. colymbadinus described here. The identity of C. zinziberatus f. flavoannulatus M.M. Moser remains unresolved, since we were not able to sequence the type material (MM 1949/0041, herb. IB). Collections examined: Austria. Süd-Tirol: Lienz, Iselsberg in coniferous forest (Picea abies) in calcareous soil, 28 July 2011, L. Albert, B. Dima, DB4571 (BP). - Hungary. Vas: Szalafő in Picea abies plantation, 24 September 2006, I. Siller, DB2535 (BP). - Finland. Perä-Pohjanmaa: Ylitornio, Palorommas, eutrophic, submesic spruce and pine forest (Picea abies and Pinus sylvestris) with Betula and Populus tremula, 18 August 2004, I. Kytövuori, IK99-295 (H). Sweden. Jämtland: Ragunda, Kullstabodarna, in rich Picea abies forest, 16 August 1992, T. E. Brandrud, H. Lindström, H. Marklund, S. Muskos, CFP1130 Neotype (S). 871 Additional specimens: Austria. Tirol: oberhalb Patscher-Alm, under Picea abies, 19 June 1998, M. Moser, IB19980007 (IB), UNITE no. UDB001095 (as C. zinziberatus). Italy. 9 May 1999, E. Campo, A. Coan MCVE19650, GenBank no. JF907865. Sweden. Uppland: Uppsala, Nåsten, in mixed forest, 27 August 2005, A.F.S. Taylor, AT2005071 (UPS), UNITE no. UDB002195. Cortinarius nolaneiformis (Velen.) Dima, Niskanen & Liimat., comb. nov. MycoBank No. MB807865 (Figs. 2b and 3a) Basionym: Hydrocybe nolaneiformis Velen. Novitates Mycologicae: 116 (1939). Typus: Czech Republic, Central Bohemia, SE of Praha, near Řičany, near Hrusice, on soil in Quercus forest, 19 May 1936, J. Velenovský, Holotype 857042 (PRM), GenBank no. KJ206486. Cortinarius umbonatoides Moënne-Locc. & Reumaux in Bidaud et al. (2008): Atlas des Cortinaires 17: 1178, nom. inval. Typus: France, Haute-Savoie, Avernioz, in calcareous Picea forest, 26 June 1991, P. Moënne-Loccoz, Holotype PML2196 (PC). GenBank no. KJ206507. Pileus 15–55 mm, conical to hemispherical, then convex, often umbonate, margin slightly striate, even, later undulate, smooth, glossy, strongly hygrophanous, dark greyish brown to dark brown, pale yellowish to ochraceous brown when dry, margin blackish brown when old. Lamellae medium spaced to fairly distant, emarginate, moderately broad, edge whitish when young, often uneven, brown to dark brown. Stipe 25–75 mm long, 5–10 mm thick at apex, 5–7 mm thick at base, cylindrical to clavate, sometimes tapering downwards, white fibrillose when young, later with brownish longitudinal streaks, pale (ochraceous) brown when dry, cortina very sparse. Universal veil yellowish, very sparse. Context occasionally hollow, uniformly brown when moist, greyish-whitish in pileus and most of the stipe when dry, brown at base. Basal mycelium whitish to more distinctly yellowish (difficult to observe). Odour distinct, somewhat reminiscent of cucumber or of other raw vegetables, slightly raphanoid. Exsiccata: pileus very dark grey (10YR 3/1), very dark greyish brown (10YR 3/2), very dark brown (10YR 2/2) to black (10YR 2/1), stipe dark greyish brown (10YR 4/2) to very dark greyish brown (10YR 3/2), base of stipe (basal mycelium?) yellow (2.5Y 8/8). UV observations: positive, lower part and the base of stipe strong yellow, pileus, lamellae and context dull yellowish brown. Spores 7.5–9(9.3)×4.5–5.5 μm, Q=1.5–1.9, av. 8.1–8.6× 4.8–5.1 μm, Q(av.)=1.62–1.72, amygdaloid, occasionally ovoid to weakly ellipsoid, somewhat thick-walled, moderately to fairly strongly verrucose, moderately to fairly weakly dextrinoid. Basidia 4-spored, 22–36×6–10.5 μm, clavate, pale olivaceous in MLZ. Lamellar edge fertile, with scattered to Author's personal copy 872 Mycol Progress (2014) 13:867–879 Fig. 1 The Bayesian 50 % majority-rule consensus tree inferred from ITS regions. PP>0.50 are indicated above branches Author's personal copy Mycol Progress (2014) 13:867–879 873 Fig. 2 Spore drawings of a Cortinarius colymbadinus (CFP1130, neotype), b C. nolaneiformis (DB4056), c C. uraceomajalis (DB1623, holotype), d C. uraceus (MM1949/0030, C. viridipes holotype), e C. uraceonemoralis (CFP1478). Drawings by T. Niskanen. Scale bar=10 μm numerous, clavate sterile cells, pale olivaceous in MLZ. Lamellar trama hyphae pale olivaceous, finely and densely encrusted. Pileipellis duplex, in overall view brown to yellowish brown, epicutis thin to moderately thick, hyphae 3–8 μm wide, hyaline or with brownish yellow content, smooth or slightly encrusted. Hypoderm with distinct, 10–20 μm wide, cylindrical to ellipsoid elements, with hyaline, smooth walls. Clamp connections present. ITS-regions (including 5.8S region): Maximum pairwise distance of the sequences two substitutions and indel positions. Cortinarius nolaneiformis differs from C. colymbadinus by 21 substitutions and indel positions and from C. uraceomajalis by 22 substitutions and indel positions. Fig. 3 Photos of a Cortinarius nolaneiformis (DB1611), b C. uraceomajalis (DB1623, holotype), c C. uraceus (TN04-872, neotype), d C. uraceonemoralis (TN04-1116, holotype). Photographs a, b by Bálint Dima and c, d by Kare Liimatainen Ecology and distribution: According to our present knowledge C. nolaneiformis grows both with coniferous (Picea) and several deciduous trees, especially with Fagus, but also with Quercus, Carpinus and Corylus, on rich to calcareous soils. The type collection of this species was found in Quercus forest, but some ITS identical collections from Europe and Iran have been collected in conifer mixed Fagus forests or other types of mixed forests. Cortinarius nolaneiformis seems to be widespread, but overlooked and most likely confused with C. uraceomajalis. Differential diagnosis: Cortinarius nolaneiformis is a medium-sized (sometimes fairly small), dark brown Telamonia, with similar appearance to other brown species Author's personal copy 874 of the subgenus, but the spring to summer occurrence and yellow UV fluorescence make this fungus quite unique among others. It can be confused with the other spring species UV positive C. uraceomajalis, but C. uraceomajalis has somewhat lighter brown pilei, generally smaller (av. 7.8–8.1×4.6– 4.7 μm), narrower (Qav. > 1.7) and more roughly verrucose, more dextrinoid spores, and a different pattern of UV fluorescence at the base of stipe. Even though we found one ITS identical collection from Sweden which has been found in August, we regard C. nolaneiformis as a mainly early season species. Basidiomata found in August can be mixed with other autumn species in similar habitats, e.g., C. rigidipes, which, however, has larger spores (8.2–10.2×5.0–6.3 μm) and negative UV fluorescence. Based on rDNA ITS sequences the closest relatives of C. nolaneiformis are C. colymbadinus and C. uraceomajalis. Both species appear relatively early in the season and have a positive UV fluorescence. In Bidaud et al. (2008) C. umbonatoides, a species similar to C. nolaneiformis, was described. The purpose of describing C. umbonatoides was to replace the name C. umbonatus (Velen.) Rob. Henry, because Bidaud et al. (2008) regarded the combination of Henry (1947) as a homonym of C. umbonatus Cleland & J.R. Harris, which they thought had been described in 1946. Nevertheless, the paper of Cleland and Harris (1948) was published one year later than that of Henry’s, thus, according to the ICN Art 53.1, the name described by the two Australian workers is illegitimate (Gasparini 2004, 2006). Therefore C. umbonatus (Velen.) Rob. Henry in fact is a valid name and C. umbonatoides was described superfluously. Hence, C. umbonatus (Velen.) Rob. Henry could be the oldest name for C. nolaneiformis. However, Hydrocybe umbonata Velen. cannot be the same species as C. nolaneiformis, since in the original description (Velenovský 1921) the spores were reported as ellipsoid, and 9–10 μm long, and a relationship with C. armeniacus (Schaeff.) Fr. and C. erugatus (Weinm.) Fr. was presumed. Cortinarius nolaneiformis, however, does not resemble either of them and the spores are amygdaloid, and shorter (7.5–9.3 μm). In addition, there is no type material available for H. umbonata for comparison with our taxon. Therefore, we conclude that H. umbonata (= C. umbonatus) and C. nolaneiformis are different species, and the latter can be used as the oldest name for the species treated here. Collections examined: Czech Republic. Central Bohemia: SE of Praha, near Řičany, near Hrusice, on soil in Quercus forest, 19 May 1936, J. Velenovský, 857042 Holotype of Hydrocybe nolaneiformis (PRM). - France. Haute-Savoie, Avernioz, on calcareous Picea forest, 26 June 1991, P. Moënne-Loccoz, PML2196 Holotype of Cortinarius umbonatoides (PC). Hungary. Borsod-Abaúj-Zemplén: Bükkzsérc under Quercus, Carpinus and Fagus on calcareous soil, 09 June 2010, L. Albert & B. Dima, DB4056 (BP); Vas: Szalafő, under Picea abies and Fagus sylvatica on rich soil, 30 May 2004, B. Mycol Progress (2014) 13:867–879 Dima & I. Siller, DB886 (BP), 07 May 2005, B. Dima & I. Siller, DB1611 (BP), under Fagus sylvatica and Picea abies on rich soil, 27 May 2006, DB2287 (BP). - Sweden. Södermanland: Nacka, Hällasgården, K. Soop, KS-CO699 (Soop pers. herb.). Additional specimens: Hungary. Vas: Orfalu under Fagus sylvatica, Quercus spp., Pinus sylvestris on rich soil, B. Dima 28 May 2010, DB3972 (BP). Sweden. Södermanland: Hellasgården, under Quercus, Corylus, 25. August 2004, Juan Santos, AT2004032 (UPS), UNITE no. UDB000709. Iran. GenBank no. FR852014; Nowshahr, with Fagaceae, UNITE no. UDB005369 (Bahram et al. 2011). Cortinarius uraceomajalis Dima, Liimat., Niskanen & Bojantchev spec. nov. MycoBank no. MB807866 (Figs. 2c and 3b) Etymology: The epithet refers to the C. uraceus-like appearance and to the month May (= “majalis”), the main fruiting period of the species. Typus: Hungary, Heves County, Recsk, Quercus cerris dominated thermophilous deciduous forest mixed with some Q. petraea and Carpinus betulus, on rich soil, 22 May 2005, B. Dima, M. Németh & Á. Sági, Holotype DB1623 (BP), Isotype (H), GenBank No. KJ206508. Pileus 20–60 mm, conical to hemispherical then convex, with distinct to low umbo, surface smooth, somewhat glossy, margin slightly fibrillose and striate, yellowish brown to brown, sometimes dark brown, pale ochraceous yellow when dry, strongly hygrophanous. Lamellae emarginate, medium spaced, moderately broad, edge finely to distinctly undulate, paler at least when young, pale yellowish (ochraceous) brown to brown. Stipe 30–80 mm long, 5–10 mm thick at apex, 4–9 mm at base, cylindrical to tapering towards base, surface smooth to finely fibrillose, pale ochraceous yellow at apex, darkening downwards with age, brown to dark brown, sometimes blackish brown at base or completely pale whitish, cortina very sparse. Universal veil very sparse, yellowish. Context whitish to pale ochraceous in pileus and in cortex of the upper half of stipe, darkening to brown below the middle part, dark brown at base. Basal mycelium whitish to more distinctly yellowish (difficult to observe). Odour similar to raw vegetables, mostly peas, especially in the older and dried out basidiomata, sometimes slightly raphanoid. Exsiccata: pileus dark brown (10YR 3/3), very dark greyish brown (10YR 3/2) to black (10YR 2/1), stipe greyish brown (10YR 5/2), dark brown (10YR 3/3) to very dark greyish brown (10YR 2/2), basal mycelium yellow (2.5Y 8/8). UV observations: positive at lower part of the stipe (bright to chrome yellow), and at the base (occasionally orange), other parts of basidiomata are negative. Spores (7.2)7.5–8.5×(4.2)4.5–5 μm, Q=1.57–1.85, av. 7.8– 8.1×4.6–4.7 μm, Q(av.)=1.70–1.73, amygdaloid, narrowly amygdaloid (to ovoid-ellipsoid), somewhat thick-walled, moderately to strongly (roughly) verrucose, especially at apex, Author's personal copy Mycol Progress (2014) 13:867–879 moderately to fairly strongly dextrinoid. Basidia 4-spored, 19– 30×(5.5)6–9(10) μm, clavate, pale olivaceous in MLZ. Lamellar edge fertile, with scattered, clavate sterile cells, pale olivaceous in MLZ. Lamellar trama hyphae pale olivaceous, almost smooth to finely encrusted in MLZ. Pileipellis duplex, in overall view yellowish brown, epicutis thin to moderately thick, hyphae 4– 9 μm wide, hyaline or with yellowish brown content, smooth or some encrusted. Hypoderm well-developed, with 12–30 μm wide, ovoid, ellipsoid to cylindrical, hyaline, smooth-walled elements. Clamp connections present. ITS-regions (including 5.8S region): All sequences are identical. Cortinarius uraceomajalis differs from C. colymbadinus by 13 substitutions and indel positions and 22 from C. nolaneiformis. Ecology and distribution: In thermophilous deciduous forests on rich to calcareous soils, possibly associated exclusively with Quercus spp. So far only known from Hungary, Bulgaria and Italy. Fruiting from April to June. Differential diagnosis: Cortinarius uraceomajalis is a vernal, medium-sized, brown Telamonia which becomes dark brown to blackish on drying, and has yellowish basal mycelium (especially seen on exsiccate). It occurs typically in warm oak or mixed deciduous forests, under Quercus spp., and produces fruitbodies from April to June. Cortinarius nolaneiformis looks similar in the field (fruiting in spring as well), but is rarely found under Quercus and seems to prefer Fagus, Fagus-Carpinus and Picea forests, often mixed with deciduous trees. It has yellow UV fluorescence on the basal mycelium as well, however, the spores are somewhat longer and broader (7.5–9×4.5–5.5 μm, Qav. = 1.6–1.7), less verrucose and less dextrinoid. Cortinarius uraceonemoralis is rarely found in springtime, but then it can be confused with C. uraceomajalis. The former, however, has completely negative UV fluorescence of the basidiomata and somewhat larger spores. Cortinarius rigidipes M.M. Moser can also be morphologically similar, but it fruits in autumn, does not have any UV fluorescence, and the spores are distinctly larger (8.2– 10.2×5.0–6.3 μm) and more strongly verrucose. Species in sections Brunnei and Disjungendi can be distinguished by their occurrence in autumn, the subglobose to ellipsoid spores and the lack of UV fluorescence. Based on the painting and the description of Bidaud et al. (2012), C. fulvoisabellinus Rob. Henry could also be a candidate for C. uraceomajalis. However, we have sequenced the rDNA ITS region of the holotype of C. fulvoisabellinus (Henry no. 1891, in herb. PC), and based on the result, the species belongs to the section Hinnulei. Collections examined: Bulgaria. Pavel Bania: Sredna Gora, Fagus sylvatica (immediate vicinity), Quercus cerris and Q. petraea 50 m away, 23 May 2012, D. Bojantchev DBB51403 (UC). Hungary. Heves: Recsk under Quercus cerris, Q. petraea and Carpinus betulus on rich soil, 22 May 2005, B. Dima, M. Németh & Á. Sági, DB1623 Holotype 875 (BP), Isotype (H); Heves: Recsk under Quercus cerris, Q. petraea and Carpinus betulus on rich soil, 22 May 2005, B. Dima, DB1624 (BP); Nógrád: Mátranovák (Nyírmádpuszta) under Quercus cerris and Q. petraea on rich soil, 26 May 2006, L. Albert & B. Dima, DB2283 (BP); GyőrMoson-Sopron: Sopron under Quercus petraea on rich soil, 27 May 2006, L. Albert & B. Dima, DB2291 (BP); Veszprém: Bakonygyepes under Quercus spp. and Carpinus betulus on calcareous soil, 4 June 2006, M. Németh, DB2303 (BP); Komárom-Esztergom: Kesztölc under Quercus cerris and Q. petraea on calcareous soil, 17 May 2008, B. Dima, DB2990 (BP). Additional specimens: Hungary. Pest: Budakeszi, under Quercus petraea and Q. cerris, 1 June 2013, B. Dima, DB5020 (BP). Italy. 25 April 1995, G. Zecchin, MCVE8011, GenBank No. JF907947 (as C. isabellinus). Section Uracei Kühner & Romagnesi ex Melot Type species: Cortinarius uraceus Fr. Basidiomata dark brown to black, often with greenish or yellowish tints. Universal veil very sparse. Spores amygdaloid to weakly ellipsoid, somewhat thick-walled, moderately to fairly strongly verrucose. Based on our phylogenetic analysis the section includes three European species, C. uraceus, C. uraceonemoralis, and C. rigidipes, and presumably also C. nauseosouraceus Niskanen, Liimat. & Ammirati from western North America, but the latter with low support. Cortinarius uraceus Fr., Epicr. syst. mycol.: 309 (1838) (Figs. 2d and 3c) Cortinarius viridipes M.M. Moser ex M.M. Moser, in Gams, Kl. Krypt.-Fl., Ed. 3, 2b/2: 324 (1967) Illustrations: Fries (1867–1884: pl. 162). Typus: Finland, Varsinais-Suomi, Lohja, Esker of Lohja, outdoor recreation area by road 41, dry pine (Pinus sylvestris) heath forest on sandy soil, with lime dust effect coming from the factory near by, 19 September 2004, I. Kytövuori & T. Niskanen, Neotype TN04-872 (H, neotype designated here), MycoBank No. MBT177203; GenBank No. KJ206522. Pileus 30–65 mm, conical to hemispherical, then low convex with an umbo, smooth, glossy, brown to very dark brown, often narrowly to 1/3 pellucid-striate, strongly hygrophanous. Lamellae medium spaced, emarginate, gill edge concolorous, brown to dark brown. Stipe 60–100×4–9 mm (apex), 5–12 mm (base), cylindrical, rarely clavate, greyish white fibrillose, often with a greenish tint, especially at the top, becoming later brown to dark brown, cortina sparse. Universal veil very sparse, difficult to detect, greyish, often with a greenish tint. Context: dark brown. Basal mycelium white. Odour: indistinct. Exsiccata: basidiomata uniformly brown (7.5YR 4/2, 7.5YR 4/3, 7.5YR 5/2), very dark brown (10YR 2/2), very Author's personal copy 876 dark grey (7.5YR 3/1) to very dark greyish brown (10YR 3/2), on pileus some part black (7.5YR 2.5/1), stipe sometimes dark reddish grey (5YR 4/2). Base of stipe (basal mycelium?) pinkish white (7.5 YR 8.5/2) to very pale brown (10YR 8/2). UV observations: all parts of the basidiomata are negative. Spores (7.7)8–9.5×(4.7)5–5.5(5.7) μm, Q=1.5–1.8, av. 8.5–8.7 ×5.1–5.4 μm, Q(av.)= 1.61–1.66, amygdaloid to weakly ellipsoid, somewhat thick-walled, moderately to fairly strongly verrucose, especially at the apex, moderately to fairly strongly dextrinoid. Basidia 4-spored, 22–34×7–9 μm, clavate, with pale sepia to olivaceous content in MLZ. Lamellar edge fertile, with scattered to numerous, clavate sterile cells, olivaceous in MLZ. Lamellar trama hyphae pale sepia to pale olivaceous, almost smooth to finely encrusted. Pileipellis duplex, in overall view pale yellowish brown, epicutis thin to moderately thick, hyphae 4–9 μm wide, hyaline or with pale brownish yellow content, walls finely encrusted or more rarely smooth. Hypoderm well-developed, with 15–25 μm wide, cylindrical to elongate elements, hyaline, walls smooth or finely encrusted. Clamp connections present. ITS-regions (including 5.8S region): Maximum pairwise distance of the sequences is 2. Cortinarius uraceus differs from C. rigidipes by 12 substitutions and indel positions and from C. uraceonemoralis by 17 substitutions and indel positions. Ecology and distribution: In mesic Picea forests, mostly on rich to calcareous soil. Fruiting from late summer to autumn but one collection from Austria made in July. Considered occasional in hemiboreal-boreal zones. Known from northern and central Europe, and from western North America but the exact distribution and frequency is unclear due to the confusion with similar species, especially with C. rigidipes. Outside boreal coniferous forests, Cortinarius uraceus might be much rarer than can be inferred from the literature. Differential diagnosis: Cortinarius uraceus is a slender, medium-sized, very dark brown species of rich coniferous forests with a very sparse veil and often greenish tinged stipe. It is hard to separate from the closely related C. rigidipes, but C. rigidipes has somewhat larger spores (8.2–10.2 × 5– 6.3 μm), and grows in pure broadleaved forests or in deciduous forest mixed with conifers (e.g., Abies, Picea). The conifer associated C. pseudouraceus Bidaud & Reumaux and C. pardinipes Romagn. are both rather C. uraceus-like species, but the former has distinctly larger spores (10–13×6.2– 8 μm) while the latter has somewhat shorter and broadly ellipsoid spores (7.5–8.5×5.5–6 μm). Fries (1838) described C. uraceus as a dark, conifer-associated species reminiscent of C. glandicolor and with an olivaceous tint at the stipe apex. A painting of the species was published in Fries (1867–1884). The name C. uraceus has been used at least for the species discussed here, but it has also included C. rigidipes (e.g., Brandrud et al. 1994) and most likely also Mycol Progress (2014) 13:867–879 C. uraceonemoralis. This conifer associated species fits best the description of Fries and is, therefore, proposed as the neotype for the species. The morphological and molecular study of the type material of C. viridipes shows that it is a later synonym of C. uraceus. Collections examined: Austria. Tirol: Mühlbachtal, Matrei, under Picea abies, among Hylocomium splendens, 5 July 1949, M. Moser, MM 1949/0030 Holotype of C. viridipes (IB). Finland. Kainuu: Puolanka, Väyrylä, Körölä, grass-herb Picea abies forest with some Pinus and hardwood bushes, 15 September 1997, I. Kytövuori, IK97-1519 (H). PeräPohjanmaa: Tornio, Korkeamaa, Runteli nature reserve area, grass-herb mesic Picea abies forest with Betula, Populus tremula and some Pinus sylvestris, 30 August 2004, K. Liimatainen & T. Niskanen, TN04-557 (H). VarsinaisSuomi: Lohja, in dry pine (Pinus sylvestris) heath forest on sandy soil with lime dust effect, coming from the factory near by, 19 September 2004, Ilkka Kytövuori & Tuula Niskanen, TN04-872 Neotype (H); Lohja, Vihti, in moist Picea abies forest with Betula, Pinus sylvestris, Alnus incana and Salix, 2 September 1998, I. Kytövuori, IK98-1607 (H); Västenfjärd, 25 September 2001, I. Kytövuori, IK01-002 (H). - Sweden. Jämtland: Ragunda, Kullstabodarna, in herbaceous Picea abies forest, 31 August 1988, T. E. Brandrud, H. Lindström, H. Marklund, S. Muskos, CFP730 (S). Cortinarius uraceonemoralis Niskanen, Liimat., Dima, Kytöv., Bojantchev & H. Lindstr. spec. nov. MycoBank no.: MB807867 (Figs. 2e and 3d) Etymology: The epithet refers to the taxonomic placement in the section Uracei (= “uraceo”) and to the appearance in deciduous forests (= “nemoralis”). Typus: Italy, Sardinia, Nuoro, Gavoi, Lago di Gusana, by the hotel Taloro, under Quercus ilex and Cistus sp. on calcareous ground, 4 November 2004, T. Niskanen & K. Liimatainen, Holotype TN04-1116 (H), Isotype (NY), GenBank No. KJ206515 Pileus 20–60 mm, conical to hemispherical then convex, with narrow to low umbo, surface smooth, glossy, strongly hygrophanous, dark brown to very dark brown, margin even to somewhat undulate, slightly striate, paler than the centre. Stipe 35–70 mm long, 5–10 thick at apex, 5–15 mm thick at base, cylindrical to clavate, almost bulbous, occasionally tapering downwards, sometimes flexuose, greyish white fibrillose then becoming glabrescent, pale yellowish to ochraceous brown at apex, with brown hygrophanous streaks lengthwise, brown to dark brown towards base, cortina very sparse. Universal veil whitish, sparse. Context in pileus thin, whitish to brown, in stipe thicker, whitish at apex, brown to blackish brown at base, marbled hygrophanous. Basal mycelium white. Odour slightly raphanoid. Exsiccata: basidiomata uniformly dark brown (7.5YR 3/2), very dark brown (7.5YR 2.5/2, 10YR 2/2) to very dark Author's personal copy Mycol Progress (2014) 13:867–879 greyish brown (10YR 3/2), on pileus some part black (10YR2/ 1). Stipe can also be brown (7.5YR 4/2) to very dark grey (7.5YR 3/1), sometimes black (7.5YR 2.5/1). Base of stipe (basal mycelium?) white (7.5YR 9/1) to pinkish grey (5YR 7/2), sometimes pink (7.5YR 7/3). UV observations: all part of the basidiomata is negative. Spores 7.5–9×(4.3)4.5–5.5 μm, Q=1.5–1.8, av. 8–8.3× 4.8–5 μm, Q(av.)=1.58–1.68, amygdaloid, broadly amygdaloid, moderately to strongly verrucose, especially at apex, moderately to fairly strongly dextrinoid. Basidia 4-spored, 20–31×6.5–9 μm, clavate, olivaceous to olivaceous brownish in MLZ. Lamellar edge fertile, with scattered, clavate sterile cells, olivaceous in MLZ. Lamellar trama hyphae olivaceous, finely and densely encrusted. Pileipellis duplex, in overall view pale reddish brown to yellowish brown, epicutis thin to moderately thick, hyphae 2–7 μm wide, hyaline or with yellowish brown content, walls encrusted or smooth. Hypoderm with thick, 15–35 μm wide, cylindrical to ellipsoid elements, hyaline or with pale yellowish brown content, smooth walls. Clamp connections present. ITS-regions (including 5.8S region): Maximum pairwise distance of the sequences is 0. Cortinarius uraceonemoralis differs from C. uraceus by 12 substitutions and indel positions and from C. rigidipes by 20 substitutions and indel positions. Ecology and distribution: In deciduous forest, especially under Quercus spp. (including Q. ilex - holotype), but also with Fagus, Carpinus, Tilia and Corylus on thermophilous, calcareous soil. According to our data, C. uraceonemoralis is widely distributed in Europe (Bulgaria, Hungary, Italy, Sweden). We found an identical sequence from an ectomycorrhizal root tip in public databases from Iran (FR852021). Differential diagnosis: C. uraceonemoralis is a fairly dark brown medium-sized Telamonia in rich to calcareous deciduous forests with small, amygdaloid, distinctly verrucose and fairly strongly dextrinoid spores. Usually it shares this habitat with C. rigidipes, but it has distinctly larger (8.2–10.2×5.0– 6.3 μm) and somewhat more dextrinoid spores. C. uraceus can be similar, but it grows with conifers, and has on average somewhat larger spores (8.5–8.7×5.1–5.4 μm). The genetically unrelated but morphologically similar C. pseudouraceus Bidaud & Reumaux and C. badiolaevis Niskanen, Liimat., Mahiques, Ballarà & Kytöv. can be distuinguished from C. uraceonemoralis by the occurrence with conifers and by the spores: the former has significantly larger spores (on average over 10×6 μm) while the latter has somewhat smaller (7–8.5 × 4.5–5 μm, Qav = 1.51–1.62) and less verrucose spores. Cortinarius nolaneiformis and C. uraceomajalis are both early season species, however, one ITS identical collection (sequence not included in the phylogenetic analysis) of C. uraceonemoralis from Hungary was 877 found in May. Thus, fruitbodies of C. uraceonemoralis collected in spring can be confused in the field with C. uraceomajalis and C. nolaneiformis, but the latter two species have yellow UV fluorescence at the base of the stipe. Collections examined: Bulgaria. Pavel Bania: Sredna Gora, Fagus sylvatica (immediate vicinity), Quercus cerris and Q. petraea 50 m away, 10 November 2012, D. Bojantchev DBB56726 (UC). - Hungary. Vas: Szalafő, “Őserdő” Forest Reserve, in mixed forest with Quercus petraea, Carpinus betulus, Fagus sylvatica and Pinus sylvestris on rich soil, 22 September 2010, B. Dima & K. Takács, ORS-ERDO 99-15-1 (BP). Budapest: Budapest, János-hegy, in grass-rich deciduous forest under Quercus cerris on calcareous soil, 6 October 2010, B. Dima, DB4322 (BP). - Italy. Süd-Tirol: Kaltern, 14 October 2000, CFP1478 (S). Sardinia: Nuoro, Gavoi, Lago di Gusana, by the hotel Taloro, under Quercus ilex and Cistus sp. on calcareous soil, 4 November 2004, T. Niskanen & K. Liimatainen, TN04-1116 Holotype (H), Isotype (NY). Sweden. Östergötland: Ödenshög, SW of Omberg, in dryish grass-herb forest, under Fagus, Quercus robur and young Picea abies, 11 September 1998, I. Kytövuori, IK98-1838 (H). Öland: Vickleby, in mixed deciduous forest with Quercus, Tilia, Corylus, Fraxinus and some Betula, in parts damp, on mull soil, 9 September 2003, T. Niskanen, K. Liimatainen & I. Kytövuori, TN03- 1264 (H). Additional specimens: Hungary. Pest: Lepence, under Quercus petraea and Q. cerris, 2 June 2013, L. Albert, AL13/279 (Albert pers. herb.). Iran. GenBank no. FR852021 (Bahram et al. 2011). Key to the currently known European species of sections Colymbadini and Uracei 1 Basidiomata entirely without UV fluorescence …… 2 2 Associated with coniferous trees ……… C. uraceus 2* In deciduous and/or in mixed forests …………… 3 3 Spores 8.2–10.2×5.0–6.3 μm, on average >9×5.5 μm, strongly verrucose, strongly dextrinoid, in pure deciduous, but also in conifer mixed forests, in autumn ……......................………… C. rigidipes 3* Spores smaller, 7.5–9×4.5–5.5 on average 8.3×5 μm, less verrucose, less dextrinoid, mainly under Quercus in autumn but rarely also in spring … C. uraceonemoralis 1*Basidiomata becoming yellow (or orange) under UV, at least at the basal part of the stipe (carefully collected!); ……………………………………… 4 4 In warm deciduous forests, to date only found under Quercus spp.; lower third of the stipe becoming Author's personal copy 878 Mycol Progress (2014) 13:867–879 striking chrome yellow under UV (basal mycelium occasionally fluorescence orange), while other part of basidiomata is completely UV negative; spores small, 7.2–8.5×4.2–5 μm ……………… C. uraceomajalis 4* In coniferous and/or deciduous forests; UV fluorescence different; spores larger ………………..… 5 5 UV fluorescence strong yellow on the entire basidiomata, very strongly chrome yellow on the whole stipe and lamellae edge; in coniferous forests under Picea (and Pinus); spores strongly verrucose and strongly dextrinoid ………………. C. colymbadinus 5* UV fluorescence dull yellow on the stipe (hard to observe), but strong (egg) yellow at the lower third of the stipe (including basal mycelium); found mostly under Picea and Fagus, more rarely under Quercus, spores less verrucose, less dextrinoid …………. C. nolaneiformis Acknowledgments We are grateful to the curators of IB, PC, PRM and S. We thank László Albert for providing Hungarian collections of C. uraceomajalis and C. uraceonemoralis, and Irén Siller for providing collections of C. colymbadinus and C. nolaneiformis. Håkan Lindström and Karl Soop are thanked for providing the Swedish collection of C. nolaneiformis. We are grateful to Gábor M. Kovács (Eötvös Loránd University, Budapest) for the opportunity to study/sequence Hungarian collections of C. uraceomajalis and C. uraceonemoralis. Heino Vänskä is thanked for the revision of Latin names and Teuvo Ahti for the help with nomenclatoric questions. 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