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Mycological Progress ISSN 1617-416XVolume 13Number 3 Mycol Progress (2014) 13:867-879DOI 10.1007/s11557-014-0970-6

Two new species of Cortinarius, subgenusTelamonia, sections Colymbadini andUracei, from Europe

Bálint Dima, Kare Liimatainen, TuulaNiskanen, Ilkka Kytövuori & DimitarBojantchev

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ORIGINAL ARTICLE

Two new species of Cortinarius, subgenus Telamonia, sectionsColymbadini 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 fromEuropean Quercus forests and one new combination is madebased on molecular and morphological data. Cortinariusuraceomajalis is a vernal species currently only known fromHungary, Bulgaria and Italy, but likely is common throughoutcentral and south-eastern Europe.Cortinarius uraceonemoralisis a widely distributed species in Europe. The new com-bination Cortinarius nolaneiformis is based on Hydrocybenolaneiformis Velen. and is a widespread vernal species inEurope. Cortinarius uraceomajalis and C. nolaneiformis arepreliminarily placed in sect. Colymbadini, characterized by hav-ing a positive (yellow) UV reaction, while C. uraceonemoraliswith a UV negative reaction is placed in sect.Uracei. A neotypeis selected for C. colymbadinus and C. uraceus to stabilizethe nomenclature. Taxonomic novelties: Cortinariusuraceomajalis Dima, Liimat., Niskanen & Bojantchev,Cortinarius uraceonemoralis Niskanen, Liimat., Dima,Kytöv., Bojantchev & H. Lindstr., Cortinarius nolaneiformis(Velen.) Dima, Niskanen & Liimat.

Keywords Agaricales . DNA . ITS . Taxonomy . UVfluorescence

Introduction

Molecular studies based on the ITS1 and ITS2 regions of therDNA, in combination with morphological and ecologicaldata, have been applied successfully for species delimitationin 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 Cortinariusbased largely on morphological characters (e.g., Bidaudet al. 1994; Consiglio et al. 2007; Consiglio 2012) is notsupported 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 betaxonomic synonyms while many others, even common ones,remain undescribed (Niskanen et al. 2008, 2013a, b). Themonophyletic subgenus Telamonia s. str. (Garnica et al.2005; Niskanen 2008; Peintner et al. 2004.) is comprised ofa very large number of species and is taxonomically themost difficult group of Cortinarii.

Subgenus Telamonia, section Uracei contains species withdark brown basidiomata which become blackish on drying,amygdaloid spores that are strongly verrucose and stronglydextrinoid, and occurrence on rich to calcareous soils. Thesection was first introduced by Kühner and Romagnesi(1953), later validated by Melot (1990) and later placed insubgenus Telamonia (Brandrud et al. 1989; Consiglio 2012).Bidaud et al. (2012) placed the section in subgenusHydrocybe(= Telamonia p. p.). The number of species included in sectionUracei ranges from three (Niskanen et al. 2008) to over 60taxa (Bidaud et al. 2012).

B. Dima (*) :K. Liimatainen : T. NiskanenDepartment of Biosciences, Plant Biology, University of Helsinki,P.O. Box 65, 00014 Helsinki, Finlande-mail: cortinarius1@gmail.com

K. Liimatainene-mail: kare.liimatainen@helsinki.fi

T. Niskanene-mail: tuula.niskanen@cortinarius.fi

I. KytövuoriBotanical Museum, University of Helsinki, P.O Box 7,00014 Helsinki, Finlande-mail: ilkka.kytovuori@funga.fi

D. Bojantchev345 Shipwatch Lane, Hercules, CA 94547, USAe-mail: dimitar@pontix.comURL: www.MushroomHobby.com

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Section Colymbadini Melot (1990) is generally character-ized by their early appearance (spring to summer), the oliva-ceous, metallic tinges on the basidiomata, and the strikinglyyellow UV fluorescence. Usually only the type species,C. colymbadinus, is included (e.g., Brandrud et al. 1998;Consiglio 2012), but, depending on different authors, alsoC. isabellinus and/or C. zinziberatus (e.g., Moser 1978) areplaced here. The presence of positive UV fluorescence ofC. colymbadinus (and C. isabellinus) led most of the authorsto place the section in subgenus Leprocybe or Cortinarius(e.g., Bidaud et al. 2005; Brandrud et al. 1989; Consiglio2012), where other UV positive species are found (e.g., sec-tion Leprocybe), but with completely different morphology.

Phylogenetic studies indicate that the species in sectionsUracei and Colymbadini are closely related and belong to thesubgenus Telamonia s. str. (Niskanen 2008; Niskanen et al.2013c) forming a still unresolved complex together withsection Cinnabarini (Ammirati et al. 2013). Here, wedescribe two new species belonging to sections Uracei andColymbadini based on morphological and molecular data andmake one new combination. Also neotypes are selected forC. uraceus and C. colymbadinus to resolve the taxonomic andnomenclatural problems within these groups.

Materials and methods

A total of 47 specimens from Finland, Sweden,Hungary, Bulgaria, Germany, Italy, Switzerland, Austria, andFrance were studied, including the type specimens ofC. fulvoisabellinus, C. pardinipes, C. pseudouraceus,C. rigidipes, C. umbonatoides, C. viridipes and Hydrocybenolaneiformis (see Table 1). Specimens are deposited in thepublic herbaria BP, H, S, IB, NY, PC, PRM, and UC. Personalcollections of László Albert (AL), Bálint Dima (DB), DimitarBojantchev (DBB) and Karl Soop (KS) were also used.Herbarium acronyms follow Thiers (continuously updated).The abbreviation of IK refers to Ilkka Kytövuori. Authorabbreviations of the species are based on the current versionof MycoBank (Robert et al. 2005).

DNAwas 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 regionsand the same primer pairs were used in direct sequencing. Forproblematic material the primer combinations ITS 1F/ITS 2and ITS 3/ITS 4 were also used. PCR amplification andsequencing followed Niskanen et al. (2009). Sequences wereassembled and edited with Sequencher 4.1 (Gene Codes,Ann Arbor, Mich., USA). BLAST queries of the publicdatabases (GenBank: http://www.ncbi.nlm.nih.gov/andUNITE: http://unite.ut.ee/), were used to check for identicalor similar sequences. For the phylogenetic analysis, ITS

sequences of the studied species and morphologicallysimilar species were included. Two species from subgenusPhlegmacium were chosen as outgroup taxa, as in Niskanenet al. (2011).

The alignment of 81 ITS sequences was produced with theprogram MUSCLE (Edgar 2004) under default settings. Thealignment comprised 683 nucleotides (including gaps). Thealignment is available at TreeBASE under S15311 (http://www.treebase.org/treebase-web/home.html). Bayesianinference (BI) was performed with MrBayes 3.1.2 (Ronquistand Huelsenbeck 2003). The best substitution model for thealignment was estimated by both the Akaike informationcriterion and the Bayesian information criterion withjModelTest version 0.1.1 (Posada 2008). A GTR model, in-cluding a gamma shape parameter, was chosen. Two indepen-dent runs with four chains in each were performed for4,000,000 generations sampling every 100th generation. Alltrees sampled before stationarity were discarded with a 25 %safety margin (burn-in of 10,000 trees [1,000,000 genera-tions]). Sampled trees from both runs were combined in a50 % majority rule consensus phylogram and posterior prob-abilities (PP) were calculated. The analysis was run withcomputer clusters of the CSC, IT Center for Science, Espoo,Finland.

Macroscopic characteristics were observed from fresh ma-terials collected by the authors, including specimens in allstages of development. Colour codes from the Munsell soilcolour charts (Munsell 2009) were used to describe the col-ours of exsiccatae. Colour photographs of fresh specimens areprovided for all species except C. colymbadinus. The speciespresented here have mostly dark brownish to blackishexsiccatae, but these colours can develop in different ways.Based on our observations, rotting basidiomes or those thatare not dried properly can develop secondary blackening ofthe pileus centre. Basidiomes that are partially dried in thefield often remain much lighter in colour, when dried, thanthose dried in prime fresh condition.

Microscopic characteristics were observed from dried ma-terials mounted in Melzer’s reagent (MLZ, 0.5 g (1.2 %)iodine, 1.5 g (3.6 %) potassium iodide, 20 cm3 chloral-hydrate, 20 cm3 distilled water) and measurements were madeon a monitor with the aid of a Zeiss Laboval 2/I/C microscopeusing 100× oil immersion lens, and connected to a ALPHADCM 130E camera and equipped with ScopePhoto 3.0.4software. Twenty spores were measured (excluding apiculusand ornamentation) from one basidioma per each collection(70–120 spores per species), from the cortina remnants or topof the stipe. Length and width were measured from the samespore, and the length/width ratios (Q-value) were calculatedfor individual spores. Spore drawings are provided to comparethe shape and ornamentation of the spores between the spe-cies. The hyphae of the lamellar trama and basidia (ten basidiaper collection, 30–50 basidia per species) were examined and

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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)

857042 Czech Republic, Central Bohemia,SE of Praha

PRM KJ206486

C. nolaneiformis DB886 Hungary, Vas, Szalafő BP KJ206487

C. nolaneiformis DB1611 Hungary, Vas, Szalafő BP KJ206488

C. nolaneiformis DB2287 Hungary, Vas, Szalafő BP KJ206489

C. nolaneiformis DB4056 Hungary, BAZ, Bükkzsérc BP 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) AB01-10-106 France, Ain, Thézillieu PC KJ206497

C. pseudouraceus IK94-1364 Sweden, Ög, Ödenshög H 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 IK94-1830 France, Ain, Oyonnax H KJ206505

C. rigidipes IK95-1873 Germany, BW, Heiligenbronn H KJ206506

C. umbonatoides Moënne-Locc. & Reumaux (holotype) PML2196 France, HS, Avernioz PC KJ206507

C. uraceomajalis Dima, Liimat.,Niskanen & Bojantchev (holotype)

DB1623 Hungary, Heves, Recsk BP KJ206508

C. uraceomajalis 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)

TN04-1116 Italy, Sardinia, Gavoi H KJ206515

C. uraceonemoralis CFP1478 Italy, Süd-Tirol, Kaltern S KJ206516

C. uraceonemoralis DB4322 Hungary, Budapest BP KJ206517

C. uraceonemoralis IK98-1838 Sweden, Ög, Ödenshög H KJ206518

C. uraceonemoralis TN03-1264 Sweden, Öl, Vickleby H KJ206519

C. uraceonemoralis ORS-ERDO99-15-1

Hungary, Vas, Szalafő BP KJ206520

C. uraceonemoralis 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 IK01-002 Finland, V, Västenfjärd H KJ206526

C. viridipes M.M. Moser ex M.M. Moser (holotype) MM1949/0030 Austria, Tirol, Mühlbachtal IB 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), HSHaute-Savoie (France), KE Komárom-Esztergom County (Hungary), L Luzern (Switzerland), Switzerl Switzerland, VeVeszprém County (Hungary)

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measured from the pieces of lamellae. In addition, thepileipellis structure was studied from both radial freehandsections and scalps from midway to the pileus centre. Themeasurements were made from scalp preparations. The mi-croscopic descriptions are almost exclusively based on ITS-sequenced material. For checking the UV fluorescence, longwave UVat wavelength 366 nmwas used on dried specimens.

Results of phylogenetic analyses

The two new species,C. uraceomajalis andC. uraceonemoralis,were strongly supported in our phylogenetic analysis (PP 0.96and 1.00). Cortinarius uraceomajalis was placed in thesection Colymbadini together with C. colymbadinus andC. nolaneiformis but with a low support (PP 0.50).Cortinarius uraceonemoralis belongs to section Uraceiand the relationship with the closest sister species C. rigidipesand C. uraceus was well supported (PP 0.96). The intraspe-cific variation in both species is 0 to 1 substitutions or indelpositions, and they differ from their closest relatives by morethan 12 substitutions and indel positions.

The sequencing of the end of the ITS2 region ofC. viridipes type material did not succeed and the partialsequence was left out of the analysis. The successfully se-quenced part, however, was identical with the ITS sequencefrom the neotype of C. uraceus, and, therefore, they areconsidered 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. uraceomajalisformed a clade, but with a low support. The species in thisgroup, however, share one characteristic, which is lackingfrom the other species of sections Uracei and Cinnabarini s.l., they all have a positive (yellow) UVreaction. In addition, allthree species fruit in spring and early summer and have rathersmall (< 9.5×4.5–5.5 μm), amygdaloid, and moderately tostrongly 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

(S, neotype designated here). Mycobank No. MBT177200;GenBank No. JX127302.

Pileus 20–70 mm, hemispherical, then low convex to al-most plane, with a low umbo, diffracted towards margin, finelyfibrillose when young, hygrophanous, chocolate brown to oli-vaceous yellow brown, with a characteristic, metallic, oliva-ceous yellow shine, which persists on drying. Lamellae medi-um spaced, emarginate, yellow-brown to cinnamon brownwhen young, later darker, with yellow, often uneven edge.

Stipe 40–100 mm long, 4–10 mm thick at apex, cylindricalto somewhat clavate, silky-fibrillose, slightly glossy, greyishyellow to brown, towards base dark brown, base with yellowmycelial felt, cortina sparse. Universal veil greenish yellow,sparse, initially sock-like, then forming a thin ring-like orindistinct zone. Context in pileus thin, pale brownish, instipe often hollow, with hygrophanous streaks, brown,darker towards base. Basal mycelium greenish yellow.Odour indistinct or slightly raphanoid. Exsiccata pileus darkbrown (10YR 3/3), dark yellowish brown (10YR 3/4,10YR 4/4) to very dark brown (10YR 2/2), becoming verydark greyish brown (10YR 3/2) to black (10YR 2/1) towardscentre. Stipe olive brown (2.5Y 4/3, 2.5Y 4/4), base of stipe(basal mycelium?) yellow (2.5Y 8/8). UV observations: posi-tive almost on the whole basidiomata, especially the stipe andthe 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, strong-ly to very strongly verrucose, moderately to fairly stronglydextrinoid. Basidia 4-spored, 22–31×7–9 μm, clavate, oliva-ceous yellow in MLZ. Lamellar edge fertile, with numerous,clavate sterile cells, concolorous with the basidia. Lamellartrama hyphae olivaceous yellow in MLZ, finely encrusted(rarely with spot-like incrustation). Pileipellis duplex, in over-all view pale yellowish brown, epicutis thin to moderatelythick, hyphae 3–8 μm wide, hyaline or with yellowish browncontent, slightly encrusted or smooth. Hypoderm weakly de-veloped, elements 10–22 μm wide, hyaline, smooth. Clampconnections present.

ITS-regions (including 5.8S region): Maximum pairwisedistance of the sequences is 0 (includes two polymorphic sites).Cortinarius colymbadinus differs from C. uraceomajalis by 13substitutions and indel positions and from C. nolaneiformis by21 substitutions and indel positions.

Ecology and distribution: In mesic Picea forests, mostly onrich soil. Producing basidiomes from summer to autumn,often fruiting early in the season. Widely distributed andoccasional in Europe. Sequenced collections are from north-ern Europe (Finland and Sweden) and central Europe (Austria,Hungary). Collection (DB2535) represents the first record inHungary.

Differential diagnosis:Cortinarius colymbadinus is a fairlyslender species with characteristic olivaceous yellow, metallictints on the basidiomata, especially on drying. It grows in

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coniferous forests and has strong, chrome yellow UV fluores-cence on the stipe and lamellae edge. The early appearance ofthe species is also characteristic. Cortinarius colymbadinus isa widely used name, at least in the Nordic countries for thespecies treated here. Also, the description by Fries (1838) fitsthe species. Therefore, we propose the collection depicted anddescribed in Brandrud et al. (1998) as a neotype to stabilize theinterpretation and nomenclature of this species.

Cortinarius isabellinus (Batsch) Fr. is often synonymisedwithC. colymbadinus (e.g. Brandrud et al. 1998). The originalplate ofAgaricus isabellinusBatsch, however, shows a funguswith bluish tint in the context, reminiscent to that ofC. biformis (Batsch 1783). This fungus does not resembleC. colymbadinus.

Cortinarius zinziberatus (Scop. : Fr.) Fr. is known as an-other early species, and the name has been applied toC. colymbadinus in Central Europe (e.g., Moser 1978). Thisfact was verified in our phylogenetic studies, since two col-lections identified as C. zinziberatus from Austria (one fromMoser, see UDB001095) had identical ITS sequences with theproposed neotype of C. colymbadinus (Fig. 1.). Cortinariuszinziberatus would be an older name for this species, but theoriginal description of Agaricus zinziberatus Scop. includes asmall-sized species (pileus only up to 2.7 cm) with brightyellow pileus and lamellae, and the smell was reported similarto that of ginger (Scopoli 1772). None of these characters arein correspondence with C. colymbadinus treated here. Fries(1821) sanctioned the name Agaricus zinziberatus and keptthe original idea of the bright yellow colours and smell.Further contradiction is the habitat which was neithermentioned by Scopoli (1772) in the original description,nor by Fries (1821). Later on Fries (1838), placed both speciesin Cortinarius, and noted the high degree of similarity withC. colymbadinus, and he published it as a rare species offrondose woods. Based on our data, however, C. colymbadinusoccurs exclusively in coniferous forests. According to the abovementioned contradictions, we are in agreement with Melot(1986) that Agaricus zinziberatus is impossible to interpret, andtherefore cannot be used for C. colymbadinus described here.The identity of C. zinziberatus f. flavoannulatus M.M. Moserremains unresolved, since we were not able to sequence the typematerial (MM 1949/0041, herb. IB).

Collections examined: Austria. Süd-Tirol: Lienz, Iselsbergin coniferous forest (Picea abies) in calcareous soil, 28 July2011, L. Albert, B. Dima, DB4571 (BP). - Hungary. Vas:Szalafő inPicea 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 Populustremula, 18 August 2004, I. Kytövuori, IK99-295 (H). -Sweden. Jämtland: Ragunda, Kullstabodarna, in rich Piceaabies forest, 16 August 1992, T. E. Brandrud, H. Lindström,H. Marklund, S. Muskos, CFP1130 Neotype (S).

Additional specimens:Austria.Tirol: oberhalb Patscher-Alm,under Picea abies, 19 June 1998, M. Moser, IB19980007 (IB),UNITE no. UDB001095 (asC. zinziberatus). Italy. 9May 1999,E. Campo, A. Coan MCVE19650, GenBank no. JF907865.Sweden. Uppland: Uppsala, Nåsten, in mixed forest, 27August 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 May1936, J. Velenovský, Holotype 857042 (PRM), GenBank no.KJ206486.

Cortinarius umbonatoides Moënne-Locc. & Reumaux inBidaud et al. (2008): Atlas des Cortinaires 17: 1178, nom.inval.

Typus: France, Haute-Savoie, Avernioz, in calcareousPicea forest, 26 June 1991, P. Moënne-Loccoz, HolotypePML2196 (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 todark 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 25–75mm long, 5–10mm thick atapex, 5–7 mm thick at base, cylindrical to clavate, sometimestapering downwards, white fibrillose when young, later withbrownish longitudinal streaks, pale (ochraceous) brown whendry, 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 distinctlyyellowish (difficult to observe). Odour distinct, somewhatreminiscent of cucumber or of other raw vegetables, slightlyraphanoid. Exsiccata: pileus very dark grey (10YR 3/1), verydark greyish brown (10YR 3/2), very dark brown (10YR 2/2)to black (10YR 2/1), stipe dark greyish brown (10YR 4/2) tovery dark greyish brown (10YR 3/2), base of stipe (basalmycelium?) yellow (2.5Y 8/8). UV observations: positive,lower part and the base of stipe strong yellow, pileus, lamellaeand 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, occasionallyovoid to weakly ellipsoid, somewhat thick-walled, moderatelyto fairly strongly verrucose, moderately to fairly weaklydextrinoid.

Basidia 4-spored, 22–36×6–10.5 μm, clavate, pale oliva-ceous in MLZ. Lamellar edge fertile, with scattered to

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Fig. 1 The Bayesian 50 % majority-rule consensus tree inferred from ITS regions. PP>0.50 are indicated above branches

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numerous, clavate sterile cells, pale olivaceous in MLZ.Lamellar trama hyphae pale olivaceous, finely and denselyencrusted. Pileipellis duplex, in overall view brown to yel-lowish brown, epicutis thin to moderately thick, hyphae3–8 μm wide, hyaline or with brownish yellow content,smooth or slightly encrusted. Hypoderm with distinct,10–20 μm wide, cylindrical to ellipsoid elements, withhyaline, smooth walls. Clamp connections present.

ITS-regions (including 5.8S region): Maximum pairwisedistance of the sequences two substitutions and indel posi-tions.Cortinarius nolaneiformis differs fromC. colymbadinusby 21 substitutions and indel positions and fromC. uraceomajalis by 22 substitutions and indel positions.

Ecology and distribution: According to our present knowl-edgeC. nolaneiformis grows both with coniferous (Picea) andseveral deciduous trees, especially with Fagus, but also withQuercus, Carpinus and Corylus, on rich to calcareous soils.The type collection of this species was found in Quercusforest, but some ITS identical collections from Europe andIran have been collected in conifer mixed Fagus forests orother types of mixed forests. Cortinarius nolaneiformis seemsto be widespread, but overlooked and most likely confusedwith C. uraceomajalis.

Differential diagnosis: Cortinarius nolaneiformis is amedium-sized (sometimes fairly small), dark brownTelamonia, with similar appearance to other brown species

Fig. 3 Photos of aCortinarius nolaneiformis(DB1611), b C. uraceomajalis(DB1623, holotype), c C. uraceus(TN04-872, neotype), dC. uraceonemoralis (TN04-1116,holotype). Photographs a, b byBálint Dima and c, d by KareLiimatainen

Fig. 2 Spore drawings of aCortinarius colymbadinus(CFP1130, neotype), bC. nolaneiformis (DB4056), cC. uraceomajalis (DB1623,holotype), d C. uraceus(MM1949/0030, C. viridipesholotype), e C. uraceonemoralis(CFP1478). Drawings by T.Niskanen. Scale bar=10 μm

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of the subgenus, but the spring to summer occurrence andyellowUV fluorescencemake this fungus quite unique amongothers. It can be confused with the other spring species UVpositive C. uraceomajalis, but C. uraceomajalis has some-what 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 fluores-cence at the base of stipe. Even though we found one ITSidentical collection from Sweden which has been found inAugust, we regard C. nolaneiformis as a mainly early seasonspecies. Basidiomata found in August can bemixed with otherautumn species in similar habitats, e.g., C. rigidipes, which,however, has larger spores (8.2–10.2×5.0–6.3 μm) and neg-ative UV fluorescence. Based on rDNA ITS sequences theclosest relatives of C. nolaneiformis are C. colymbadinus andC. uraceomajalis. Both species appear relatively early in theseason and have a positive UV fluorescence.

In Bidaud et al. (2008) C. umbonatoides, a species similartoC. nolaneiformis, was described. The purpose of describingC. umbonatoides was to replace the name C. umbonatus(Velen.) Rob. Henry, because Bidaud et al. (2008) regardedthe combination of Henry (1947) as a homonym ofC. umbonatus Cleland & J.R. Harris, which they thoughthad been described in 1946.

Nevertheless, the paper of Cleland and Harris (1948) waspublished one year later than that of Henry’s, thus, accordingto the ICN Art 53.1, the name described by the two Australianworkers is illegitimate (Gasparini 2004, 2006). ThereforeC. umbonatus (Velen.) Rob. Henry in fact is a valid nameand C. umbonatoides was described superfluously. Hence,C. umbonatus (Velen.) Rob. Henry could be the oldest namefor C. nolaneiformis. However, Hydrocybe umbonata Velen.cannot be the same species as C. nolaneiformis, since in theoriginal description (Velenovský 1921) the spores were re-ported as ellipsoid, and 9–10 μm long, and a relationship withC. armeniacus (Schaeff.) Fr. and C. erugatus (Weinm.) Fr.was presumed. Cortinarius nolaneiformis, however, does notresemble either of them and the spores are amygdaloid, andshorter (7.5–9.3 μm). In addition, there is no type materialavailable for H. umbonata for comparison with our taxon.Therefore, we conclude thatH. umbonata (=C. umbonatus) andC. nolaneiformis are different species, and the latter canbe used as the oldest name for the species treated here.

Collections examined: Czech Republic. Central Bohemia:SE of Praha, nearŘičany, near Hrusice, on soil inQuercus forest,19 May 1936, J. Velenovský, 857042 Holotype of Hydrocybenolaneiformis (PRM). - France. Haute-Savoie, Avernioz, oncalcareous 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 Piceaabies and Fagus sylvatica on rich soil, 30 May 2004, B.

Dima & I. Siller, DB886 (BP), 07 May 2005, B. Dima & I.Siller, DB1611 (BP), under Fagus sylvatica and Picea abieson rich soil, 27 May 2006, DB2287 (BP). - Sweden.Södermanland: Nacka, Hällasgården, K. Soop, KS-CO699(Soop pers. herb.).

Additional specimens: Hungary. Vas: Orfalu under Fagussylvatica, Quercus spp., Pinus sylvestris on rich soil, B. Dima28 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 ap-

pearance and to the month May (= “majalis”), the mainfruiting period of the species.

Typus: Hungary, Heves County, Recsk, Quercus cerrisdominated thermophilous deciduous forest mixed with someQ. 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 tobrown, sometimes dark brown, pale ochraceous yellow whendry, 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 mmlong, 5–10 mm thick at apex, 4–9 mm at base, cylindrical totapering towards base, surface smooth to finely fibrillose, paleochraceous yellow at apex, darkening downwards with age,brown to dark brown, sometimes blackish brown at base orcompletely pale whitish, cortina very sparse.Universal veil verysparse, yellowish. Context whitish to pale ochraceous in pileusand in cortex of the upper half of stipe, darkening to brownbelow the middle part, dark brown at base. Basal myceliumwhitish to more distinctly yellowish (difficult to observe).Odour similar to raw vegetables, mostly peas, especially in theolder and dried out basidiomata, sometimes slightly raphanoid.

Exsiccata: pileus dark brown (10YR 3/3), very darkgreyish brown (10YR 3/2) to black (10YR 2/1), stipe greyishbrown (10YR 5/2), dark brown (10YR 3/3) to very darkgreyish brown (10YR 2/2), basal mycelium yellow (2.5Y8/8). UV observations: positive at lower part of the stipe(bright to chrome yellow), and at the base (occasionallyorange), 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, narrowlyamygdaloid (to ovoid-ellipsoid), somewhat thick-walled, moder-ately to strongly (roughly) verrucose, especially at apex,

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moderately to fairly strongly dextrinoid. Basidia 4-spored, 19–30×(5.5)6–9(10)μm, clavate, pale olivaceous inMLZ.Lamellaredge fertile, with scattered, clavate sterile cells, pale olivaceous inMLZ. Lamellar trama hyphae pale olivaceous, almost smooth tofinely encrusted in MLZ. Pileipellis duplex, in overall viewyellowish brown, epicutis thin to moderately thick, hyphae 4–9 μmwide, hyaline or with yellowish brown content, smooth orsome encrusted. Hypoderm well-developed, with 12–30 μmwide, ovoid, ellipsoid to cylindrical, hyaline, smooth-walledelements. Clamp connections present.

ITS-regions (including 5.8S region): All sequencesare identical. Cortinarius uraceomajalis differs fromC. colymbadinus by 13 substitutions and indel positions and22 from C. nolaneiformis.

Ecology and distribution: In thermophilous deciduous for-ests on rich to calcareous soils, possibly associated exclusivelywithQuercus spp. So far only known from Hungary, Bulgariaand Italy. Fruiting from April to June.

Differential diagnosis: Cortinarius uraceomajalis is a ver-nal, medium-sized, brown Telamonia which becomes darkbrown to blackish on drying, and has yellowish basal myce-lium (especially seen on exsiccate). It occurs typically inwarmoak or mixed deciduous forests, under Quercus spp., andproduces fruitbodies from April to June. Cortinariusnolaneiformis looks similar in the field (fruiting in spring aswell), but is rarely found under Quercus and seems to preferFagus, Fagus-Carpinus and Picea forests, often mixed withdeciduous trees. It has yellow UV fluorescence on the basalmycelium as well, however, the spores are somewhat longerand broader (7.5–9×4.5–5.5 μm, Qav. = 1.6–1.7), less verru-cose and less dextrinoid. Cortinarius uraceonemoralis is rare-ly found in springtime, but then it can be confused withC. uraceomajalis. The former, however, has completely neg-ative UV fluorescence of the basidiomata and somewhatlarger spores. Cortinarius rigidipes M.M. Moser can also bemorphologically similar, but it fruits in autumn, does not haveany UV fluorescence, and the spores are distinctly larger (8.2–10.2×5.0–6.3 μm) and more strongly verrucose. Species insections Brunnei and Disjungendi can be distinguished bytheir occurrence in autumn, the subglobose to ellipsoid sporesand 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 candi-date for C. uraceomajalis. However, we have sequenced therDNA ITS region of the holotype of C. fulvoisabellinus(Henry no. 1891, in herb. PC), and based on the result, thespecies belongs to the section Hinnulei.

Collections examined: Bulgaria. Pavel Bania: SrednaGora, Fagus sylvatica (immediate vicinity), Quercus cerrisand Q. petraea 50 m away, 23 May 2012, D. BojantchevDBB51403 (UC). Hungary. Heves: Recsk under Quercuscerris, Q. petraea and Carpinus betulus on rich soil, 22 May2005, B. Dima, M. Németh & Á. Sági, DB1623 Holotype

(BP), Isotype (H); Heves: Recsk under Quercus cerris,Q. petraea and Carpinus betulus on rich soil, 22 May 2005,B . Dima , DB1624 (BP) ; Nógrád : Má t r anovák(Nyírmádpuszta) under Quercus cerris and Q. petraea on richsoil, 26May 2006, L. Albert & B. Dima, DB2283 (BP);Győr-Moson-Sopron: Sopron underQuercus petraea on rich soil, 27May 2006, L. Albert & B. Dima, DB2291 (BP); Veszprém:Bakonygyepes under Quercus spp. and Carpinus betulus oncalcareous soil, 4 June 2006, M. Németh, DB2303 (BP);Komárom-Esztergom: Kesztölc under Quercus cerris andQ. petraea on calcareous soil, 17 May 2008, B. Dima,DB2990 (BP).

Additional specimens: Hungary. Pest: Budakeszi, underQuercus 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 amygdaloidto weakly ellipsoid, somewhat thick-walled, moderately tofairly strongly verrucose. Based on our phylogenetic analysisthe section includes three European species, C. uraceus,C. uraceonemoralis, and C. rigidipes, and presumably alsoC. nauseosouraceus Niskanen, Liimat. & Ammirati fromwestern 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, inGams, 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 fromthe 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 lowconvex with an umbo, smooth, glossy, brown to very darkbrown, often narrowly to 1/3 pellucid-striate, stronglyhygrophanous.

Lamellaemedium spaced, emarginate, gill edge concolorous,brown to dark brown.

Stipe 60–100×4–9 mm (apex), 5–12 mm (base), cylindri-cal, rarely clavate, greyish white fibrillose, often with a green-ish tint, especially at the top, becoming later brown to darkbrown, cortina sparse. Universal veil very sparse, difficult todetect, greyish, often with a greenish tint.

Context: dark brown. Basal myceliumwhite.Odour: indis-tinct. Exsiccata: basidiomata uniformly brown (7.5YR 4/2,7.5YR 4/3, 7.5YR 5/2), very dark brown (10YR 2/2), very

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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 darkreddish grey (5YR 4/2). Base of stipe (basal mycelium?)pinkish white (7.5 YR 8.5/2) to very pale brown (10YR8/2). UV observations: all parts of the basidiomata arenegative.

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 toweakly ellipsoid, somewhat thick-walled, moderately to fairlystrongly verrucose, especially at the apex, moderately to fairlystrongly dextrinoid. Basidia 4-spored, 22–34×7–9 μm, cla-vate, with pale sepia to olivaceous content in MLZ. Lamellaredge fertile, with scattered to numerous, clavate sterile cells,olivaceous in MLZ. Lamellar trama hyphae pale sepia to paleolivaceous, almost smooth to finely encrusted. Pileipellisduplex, in overall view pale yellowish brown, epicutis thinto moderately thick, hyphae 4–9 μm wide, hyaline or withpale brownish yellow content, walls finely encrusted or morerarely smooth. Hypoderm well-developed, with 15–25 μmwide, cylindrical to elongate elements, hyaline, walls smoothor finely encrusted. Clamp connections present.

ITS-regions (including 5.8S region): Maximum pairwisedistance of the sequences is 2. Cortinarius uraceus differsfrom C. rigidipes by 12 substitutions and indel positions andfrom C. uraceonemoralis by 17 substitutions and indelpositions.

Ecology and distribution: In mesic Picea forests, mostly onrich to calcareous soil. Fruiting from late summer to autumnbut one collection from Austria made in July. Consideredoccasional in hemiboreal-boreal zones. Known from northernand central Europe, and from western North America but theexact distribution and frequency is unclear due to the confu-sion with similar species, especially withC. rigidipes. Outsideboreal coniferous forests, Cortinarius uraceusmight be muchrarer than can be inferred from the literature.

Differential diagnosis: Cortinarius uraceus is a slender,medium-sized, very dark brown species of rich coniferousforests with a very sparse veil and often greenish tinged stipe.It is hard to separate from the closely related C. rigidipes, butC. rigidipes has somewhat larger spores (8.2–10.2×5–6.3 μm), and grows in pure broadleaved forests or in decidu-ous forest mixed with conifers (e.g.,Abies, Picea). The coniferassociated C. pseudouraceus Bidaud & Reumaux andC. pardinipes Romagn. are both rather C. uraceus-like spe-cies, but the former has distinctly larger spores (10–13×6.2–8 μm) while the latter has somewhat shorter and broadlyellipsoid spores (7.5–8.5×5.5–6 μm). Fries (1838) describedC. uraceus as a dark, conifer-associated species reminiscentof C. glandicolor and with an olivaceous tint at thestipe apex. A painting of the species was published inFries (1867–1884). The name C. uraceus has been used atleast for the species discussed here, but it has also includedC. rigidipes (e.g., Brandrud et al. 1994) and most likely also

C. uraceonemoralis. This conifer associated species fits bestthe description of Fries and is, therefore, proposed as theneotype for the species. The morphological and molecularstudy of the type material of C. viridipes shows that it is alater synonym of C. uraceus.

Collections examined:Austria. Tirol: Mühlbachtal, Matrei,under Picea abies, among Hylocomium splendens, 5 July1949, M. Moser, MM 1949/0030 Holotype of C. viridipes(IB). Finland. Kainuu: Puolanka, Väyrylä, Körölä, grass-herbPicea abies forest with some Pinus and hardwood bushes, 15September 1997, I. Kytövuori, IK97-1519 (H). Perä-Pohjanmaa: Tornio, Korkeamaa, Runteli nature reserve area,grass-herb mesic Picea abies forest with Betula, Populustremula and some Pinus sylvestris, 30 August 2004, K.Liimatainen & T. Niskanen, TN04-557 (H). Varsinais-Suomi: Lohja, in dry pine (Pinus sylvestris) heath forest onsandy soil with lime dust effect, coming from the factory nearby, 19 September 2004, Ilkka Kytövuori & Tuula Niskanen,TN04-872 Neotype (H); Lohja, Vihti, in moist Picea abiesforest with Betula, Pinus sylvestris, Alnus incana and Salix, 2September 1998, I. Kytövuori, IK98-1607 (H); Västenfjärd,25 September 2001, I. Kytövuori, IK01-002 (H). - Sweden.Jämtland: Ragunda, Kullstabodarna, in herbaceous Piceaabies 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 indeciduous forests (= “nemoralis”).

Typus: Italy, Sardinia, Nuoro, Gavoi, Lago di Gusana, bythe hotel Taloro, under Quercus ilex and Cistus sp. on calcar-eous 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, stronglyhygrophanous, dark brown to very dark brown, margin evento somewhat undulate, slightly striate, paler than the centre.

Stipe 35–70 mm long, 5–10 thick at apex, 5–15 mm thickat base, cylindrical to clavate, almost bulbous, occasionallytapering downwards, sometimes flexuose, greyish whitefibrillose then becoming glabrescent, pale yellowish toochraceous brown at apex, with brown hygrophanous streakslengthwise, brown to dark brown towards base, cortina verysparse. Universal veil whitish, sparse.

Context in pileus thin, whitish to brown, in stipethicker, whitish at apex, brown to blackish brown atbase, 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

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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 (5YR7/2), sometimes pink (7.5YR 7/3). UV observations: all partof 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 amygda-loid, 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 brownishin MLZ. Lamellar edge fertile, with scattered, clavate sterilecells, olivaceous in MLZ. Lamellar trama hyphae oli-vaceous, finely and densely encrusted. Pileipellis du-plex, in overall view pale reddish brown to yellowishbrown, epicutis thin to moderately thick, hyphae 2–7 μmwide, hyaline or with yellowish brown content, wallsencrusted or smooth.

Hypoderm with thick, 15–35 μm wide, cylindrical to ellip-soid elements, hyaline or with pale yellowish brown content,smooth walls. Clamp connections present.

ITS-regions (including 5.8S region): Maximum pairwisedistance of the sequences is 0. Cortinarius uraceonemoralisdiffers from C. uraceus by 12 substitutions and indelpositions and from C. rigidipes by 20 substitutions andindel positions.

Ecology and distribution: In deciduous forest, especiallyunder Quercus spp. (including Q. ilex - holotype), but alsowith Fagus, Carpinus, Tilia and Corylus on thermophilous,calcareous soil. According to our data, C. uraceonemoralis iswidely distributed in Europe (Bulgaria, Hungary, Italy,Sweden). We found an identical sequence from anectomycorrhizal root tip in public databases from Iran(FR852021).

Differential diagnosis: C. uraceonemoralis is a fairly darkbrown medium-sized Telamonia in rich to calcareous decidu-ous forests with small, amygdaloid, distinctly verrucose andfairly strongly dextrinoid spores. Usually it shares this habitatwith C. rigidipes, but it has distinctly larger (8.2–10.2×5.0–6.3 μm) and somewhat more dextrinoid spores. C. uraceuscan be similar, but it grows with conifers, and has on averagesomewhat larger spores (8.5–8.7×5.1–5.4 μm). The geneti-cally unrelated but morphologically similar C. pseudouraceusBidaud & Reumaux and C. badiolaevis Niskanen, Liimat.,Mahiques, Ballarà & Kytöv. can be distuinguished fromC. uraceonemoralis by the occurrence with conifers and bythe spores: the former has significantly larger spores (onaverage over 10×6 μm) while the latter has somewhat smaller(7–8.5×4.5–5 μm, Qav=1.51–1.62) and less verrucosespores. Cortinarius nolaneiformis and C. uraceomajalisare both early season species, however, one ITS identi-cal collection (sequence not included in the phylogenet-ic analysis) of C. uraceonemoralis from Hungary was

found in May. Thus, fruitbodies of C. uraceonemoraliscollected in spring can be confused in the field withC. uraceomajalis and C. nolaneiformis, but the latter twospecies have yellow UV fluorescence at the base of the stipe.

Collections examined: Bulgaria. Pavel Bania: SrednaGora, Fagus sylvatica (immediate vicinity), Quercus cerrisandQ. petraea 50 m away, 10 November 2012, D. BojantchevDBB56726 (UC). - Hungary. Vas: Szalafő, “Őserdő” ForestReserve, in mixed forest with Quercus petraea, Carpinusbetulus, Fagus sylvatica and Pinus sylvestris on rich soil, 22September 2010, B. Dima & K. Takács, ORS-ERDO 99-15-1(BP). Budapest: Budapest, János-hegy, in grass-rich deciduousforest underQuercus cerris on calcareous soil, 6 October 2010,B. Dima, DB4322 (BP). - Italy. Süd-Tirol: Kaltern, 14 October2000, CFP1478 (S). Sardinia: Nuoro, Gavoi, Lago di Gusana,by the hotel Taloro, under Quercus ilex and Cistus sp. oncalcareous soil, 4 November 2004, T. Niskanen & K.Liimatainen, TN04-1116 Holotype (H), Isotype (NY). -Sweden. Östergötland: Ödenshög, SW of Omberg, in dryishgrass-herb forest, under Fagus, Quercus robur and youngPicea abies, 11 September 1998, I. Kytövuori, IK98-1838(H). Öland: Vickleby, in mixed deciduous forest withQuercus, Tilia, Corylus, Fraxinus and some Betula, in partsdamp, on mull soil, 9 September 2003, T. Niskanen, K.Liimatainen & I. Kytövuori, TN03- 1264 (H).

Additional specimens: Hungary. Pest: Lepence, underQuercus 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 sectionsColymbadini and Uracei

1 Basidiomata entirely without UV fluorescence…… 2

2 Associated with coniferous trees……… C. uraceus2* In deciduous and/or inmixed forests…………… 3

3 Spores 8.2–10.2×5.0–6.3 μm, on average >9×5.5 μm,strongly verrucose, strongly dextrinoid, in puredeciduous, but also in conifer mixed forests, inautumn ……......................………… C. rigidipes

3* Spores smaller, 7.5–9×4.5–5.5 on average 8.3×5 μm,less verrucose, less dextrinoid, mainly underQuercus inautumn but rarely also in spring…C. uraceonemoralis

1*Basidiomata becoming yellow (or orange) under UV,at least at the basal part of the stipe (carefullycollected!); ……………………………………… 4

4 In warm deciduous forests, to date only found underQuercus spp.; lower third of the stipe becoming

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striking chrome yellow under UV (basal myceliumoccasionally fluorescence orange), while other part ofbasidiomata is completely UV negative; spores small,7.2–8.5×4.2–5 μm ……………… C. uraceomajalis4* In coniferous and/or deciduous forests; UV fluo-rescence different; spores larger ………………..… 5

5 UV fluorescence strong yellow on the entirebasidiomata, very strongly chrome yellow on thewhole stipe and lamellae edge; in coniferous forestsunderPicea (andPinus); spores strongly verrucose andstrongly dextrinoid………………. C. colymbadinus

5* UV fluorescence dull yellow on the stipe (hard to ob-serve), but strong (egg) yellow at the lower third of thestipe (including basal mycelium); found mostly underPicea andFagus, more rarely underQuercus, spores lessverrucose, less dextrinoid…………. C. nolaneiformis

Acknowledgments We are grateful to the curators of IB, PC, PRM andS. We thank László Albert for providing Hungarian collections ofC. uraceomajalis and C. uraceonemoralis, and Irén Siller for providingcollections of C. colymbadinus and C. nolaneiformis. Håkan Lindströmand Karl Soop are thanked for providing the Swedish collection ofC. nolaneiformis. We are grateful to Gábor M. Kovács (Eötvös LorándUniversity, Budapest) for the opportunity to study/sequence Hungariancollections ofC. uraceomajalis andC. uraceonemoralis. HeinoVänskä isthanked for the revision of Latin names and Teuvo Ahti for the help withnomenclatoric questions. Reviewers are thanked for valuable commentswhich helped us to improve the manuscript. First author’s field work waspartly supported by the Hungarian Scientific Research Fund (OTKA,K79158) and the Directorate of Őrség National Park. This work wassupported by the Academy of Finland (project 129052) and the Ministryof Environment, Finland (YM38/5512/2009).

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