Turkish Journal of Botany Turk J Bot (2018) 42: 327-334 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1710-8

Cortinarius conicoumbonatus ( subgen. Telamonia sect. Hinnulei): a new species from spruce- forests of the East Black Sea Region of Turkey

1, 2 Ertuğrul SESLİ *, Kare LIIMATAINEN 1 Department of Biology Education, Faculty of Education, Karadeniz Technical University, Trabzon, Turkey 2 Jodrell Laboratory, Royal Botanic Gardens, Kew, Surrey, United Kingdom

Received: 04.10.2017 Accepted/Published Online: 02.01.2018 Final Version: 03.05.2018

Abstract: Cortinarius conicoumbonatus is described as new to science based on the material collected from spruce-beech forests of Maçka, Trabzon, Turkey. The new species belongs to subgenus Telamonia section Hinnulei. A full description, in situ and micromorphological illustrations, a phylogenetic tree, and a short discussion are provided.

Key words: , mushroom phylogeny

1. Introduction Reumaux (Niskanen et al., 2006) and C. badioflavidus Cortinarius (Pers.) Gary is the largest of Ammirati, Beug, Niskanen, Liimat. & Bojantchev (Li et al., ectomycorrhizal fungi, containing more than 2000 2016). The present paper describes a new species based on described species worldwide and more than 120 taxa in morphological and molecular data. Turkey. Most of the species are associated with Pinaceae and Fagaceae, especially with pine, spruce, birch, oak, 2. Materials and methods beech, and southern beech in Turkey (Sesli, 2006; Kirk 2.1. Sampling and morphological studies et al., 2008; Sesli and Denchev, 2008; Kaya et al., 2009; The basidiomata were collected from the Sevinç Demirel et al., 2010; Uzun et al., 2013; Liimatainen et neighborhood of Maçka, Trabzon, Turkey, on 3 October al., 2014; Akata et al., 2015; Sesli et al., 2015; Sesli and 2014. The photographs were taken in situ and one Moreau, 2015). The genusCortinarius can be divided basidioma was used to obtain a . Other into four subgenera based on macromorphology of the basidiomata were dried with an electric dryer for future basidiomata: Cortinarius (Pers.) Gary, Myxacium (Fr.) microscopic studies. Very thin freehand sections of the P.Kumm., Phlegmacium (Fr.) Wünsche, and Telamonia (Fr.) pileus surface and lamellae of exsiccatae were obtained Wünsche (Niskanen and Kytövuori, 2012). However, this with a razor blade, treated in potassium hydroxide, stained basic taxonomic division is not supported by phylogenetic with Congo red, and photographed under a Zeiss Axio studies (Garnica et al., 2005; Harrower et al., 2011), but Imager A2 trinocular research microscope. Microscopic can be used for purposes of identification (Brandrud et structures, including basidiospores, basidia, marginal al., 1989; Niskanen et al., 2008a). The subgenus Telamonia cells, and pileipellis, were measured with Axio Imager has been divided into a number of sections including the section Hinnulei Melot, to which this new species belongs. software using Melzer’s reagent. The description was made Section Hinnulei is characterized by its yellowish brown according to Smith (1939), Ammirati (1989), Moser et al. to ochre brown and strongly hygrophanous basidiomata; (1995), Bidaud et al. (1997), Høiland and Holst-Jensen pungent to raphanoid odor; generally whitish universal (2000), Moser (2002), Bidaud et al. (2006), and Clémençon veil; and coarsely verrucose, subglobose to ellipsoid (2009). The type material is deposited in the Fungarium of basidiospores (Bidaud et al., 1997, 2006; Niskanen et al., KATO, Fatih Faculty of Education, Karadeniz Technical 2008b). The number of species placed in Hinnulei varies University, Trabzon, Turkey. from 85 (Bidaud et al., 1997) to five (Niskanen et al., 2008a) 2.2. DNA extraction, PCR amplification, and sequencing taxa. So far only two species in the section Hinnulei have Total DNA was extracted from the pileus context of the been studied using molecular data, C. hinnuleoarmillatus dried specimens (Table 1) employing a modified protocol * Correspondence: [email protected] 327 SESLİ and LIIMATAINEN / Turk J Bot based on that of Murray and Thompson (1980). A 3. Results portion of each specimen was blended with the aid of a 3.1. Molecular analysis micropestle in 600 µL of CTAB buffer (CTAB 2%, NaCl ITS rDNA analysis obtained significant support for most 1.4 M, EDTA pH 8.0 20 mM, Tris-HCl pH 8.0 100 mM). sections of Cortinarius in the work of Niskanen et al. The resulting mixture was incubated for 15 min at 65 °C. (2013), but some of them were not well supported (e.g., A similar volume of chloroform : isoamyl alcohol (24 : 1) Uracei/Colymbadini/Cinnabarini). Some clades, such as was added and carefully mixed with the samples until their Furvolaesi, received low support using the multigenic emulsion. It was then centrifuged for 10 min at 13,000 × approach (Niskanen et al., 2013) and were found to g, and the DNA in the supernatant was precipitated with a be polyphyletic with ITS rDNA data alone. ITS rDNA volume of isopropanol. After centrifuging the supernatant sequences from the new species C. conicoumbonatus for an additional 15 min at the same speed, the pellet was (Table 1) are identical and are closely related to C. washed in 70% cold ethanol, centrifuged again for 2 min, hinnuleus Fr. (I) (PP 0.86, BP 85), and to a lesser extent to and dried. It was then resuspended in 200 µL of ddH2O. other species in section Hinnulei (PP 0.90, BP 56), such as PCR amplification was performed with the primers ITS1F C. hinnuleoarmillatus and C. badioflavidus. Therefore, C. and ITS4 (White et al., 1990; Gardes and Bruns, 1993) for conicoumbonatus (Figure 2) is formally placed in section the ITS region, while LR0R and LR5 (Vilgalys and Hester, Hinnulei. 1990; Cubeta et al., 1991) were used to amplify the 28S 3.2. rDNA region. PCR reactions were performed with BioTaq Cortinarius conicoumbonatus E. Sesli, Liimat. & K. (Bioline) polymerase under a program consisting of a hot Demirel, sp. nov. (Figures 2 and 3) start at 95 °C for 5 min, followed by 35 cycles at 94 °C, 54 MycoBank: MB823044 °C, and 72 °C (45, 30, and 45 s, respectively) and a final Diagnosis: Basidiomata gregarious in mixed spruce- step at 72 °C for 10 min. PCR products were checked in 1% beech forests, small to medium in size; pileus conic to umbonate, margin incurved, cream colored, light ocher agarose gels, and the products of positive reactions were to beige brown, reddish-brown spotted, surface smooth sequenced with one or both PCR primers. Chromatograms to fibrous, strongly hygrophanous; concolorous with were checked searching for putative reading errors, which the pileus, generally long and curved, covered with a white were corrected. veil from the middle to the base; lamellae light ocher- 2.3. Sequence alignment and phylogenetic analysis brown to reddish or rust-brown; odor sharp, unpleasant; BLAST (Altschul et al., 1997) was used to select the most basidiospores broadly ellipsoid to obovoid, moderately to closely related ITS rDNA sequences from INSD public strongly verrucose, on average 7.9 × 5.2 µm. It differs in databases. Sequences came mainly from Peintner et al. the ITS from the other known species of section Hinnulei (2003), Kytövuori et al. (2005), Niskanen et al. (2006, by 8 substitutions and indel positions (1.5%). 2013), Ammirati et al. (2013), Dima et al. (2014), and Holotype: Turkey, Trabzon, Maçka, Sevinç Brandrud et al. (2015). Sequences first were aligned in neighborhood, 40°51′09.04″N, 39°37′13.33″E, 868 m alt., MEGA 5.0 software (Tamura et al., 2011) with its Clustal in mixed spruce-beech forest, 03.10.2014, leg. E. Sesli W application and then corrected manually. The ITS (KATO - Fungi 3455). GenBank numbers: MF696139 (ITS alignment was 607 nucleotides long and included 261 region), MF696141 (ITS region), MF696140 (LSU region). variable sites. The 28S rDNA region was not employed Etymology: The epithet is derived from the conic to in the analyses. The aligned data set was subjected to umbonate shape of the pileus. MrModeltest 2.3 (Nylander, 2004) in PAUP* 4.0b10. Pileus 25–50 mm wide, hemispherical when young, Model GTR+G+I was selected and implemented in later conic-umbonate or obtusely umbonate, with a sharp umbo in age; margin incurved and covered with cottony MrBayes 3.1 (Ronquist and Huelsenbeck, 2003), where velar remnants, the edges sometimes upturned, sometimes a Bayesian analysis was performed (two simultaneous split from the edge to the disc; surface smooth to fibrous, runs, six chains, temperature set to 0.2, sampling every sometimes furfuraceus, strongly hygrophanous, gray- 100th generation) until convergence parameters were met beige, tan, cream colored, light ocher to beige brown, after 8.04M generations, the standard deviation having reddish-brown spotted. Lamellae notched, broadly fell below 0.01. Finally, a full search for the best-scoring attached, thick, broad; light ocher-brown, dark orange- maximum likelihood tree was performed in RAxML brown to reddish or rust-brown; edges even or rarely (Figure 1) using the standard search algorithm (2000 crenate. Stipe 50–100 × 5–10 mm; solid when young bootstrap replications). The significance threshold was and slightly hollow when mature; cylindrical to clavate; set above 0.95 for posterior probability (PP) and 70% generally curved, tapered towards the base; creamy, gray- bootstrap proportions (BP) (Stamatakis, 2006). beige, light ocher to beige brown, dark reddish when

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AY669585 Cortinarius olivaceofuscus KJ206484 Cortinarius colymbadinus 1.00/99 Coly m NR 131819 Cortinarius colymbadinus TYPE 1.0/100 NR 131826 Cortinarius brunneovernus TYPE

0.66/54 KC608577 Cortinarius brunneovernus b a d EU266679 Cortinarius clarobrunneus Br u i 1.0/100 n 1.00/99 NR 131805 Cortinarius clarobrunneus TYPE nn e i /

1.0/100 EU266642 Cortinarius brunneus C i i nn a b

EU266641 Cortinarius brunneus +

0.98/62 0.71/56 KC608589 Cortinarius adustorimosus

NR 120163 Cortinarius cinnabarinus TYPE ar i 1.0/100 JX114944 Cortinarius cinnabarinus n

U56023 Cortinarius californicus i 1.00/90 JX114946 Cortinarius californicus

1.0/100 KP165544 Cortinarius uraceisporus NR 131878 Cortinarius uraceisporus TYPE

1.0/100 NR 131875 Cortinarius ferrugineovelatus TYPE 0.89/66 GQ159884 Cortinarius ferrugineovelatus KU041740 Cortinarius roseobasilis 0.99/100 KU041741 Cortinarius roseobasilis NR 120095 Cortinarius aavae TYPE

1.00/99 DQ117926 Cortinarius hinnuleus (II) 0.83/56 AY669667 ‘Cortinarius helvolus’

1.0/100 HQ604716 ‘Cortinarius casimiri’ FJ039550 ‘Cortinarius subsertipes’ 1.00/100 MF696139 Cortinarius conicoumbonatus sp. nov. MF696141 Cortinarius conicoumbonatus sp. nov. AY669665 Cortinarius hinnuleus (I) 0.86/85 Hinnulei AY083183 Cortinarius hinnuleus (I) 1.0/100 AY083184 Cortinarius hinnuleus (I) NR 131790 Cortinarius hinnuleoarmillatus TYPE 1.00/94 DQ499463 Cortinarius hinnuleoarmillatus KU041729 Cortinarius badioflavidus 1.00/88 KU041732 Cortinarius badioflavidus AY669658 Cortinarius umbrinolens 1.0/100 KX449461 Cortinarius umbrinolens KC608587 Cortinarius politus 1.0/100 NR 131829 Cortinarius politus TYPE NR 131791 Cortinarius pseudobovinus DQ497191 Cortinarius rubrovioleipes 0.67/58 0.98/93 DQ499466 Cortinarius boulderensis Boulderenses 0.99/98 NR 121207 Cortinarius boulderensis TYPE KT591596 Cortinarius fuscescens 1.0/100 NR 131879 Cortinarius fuscescens TYPE

1.0/100 KT591607 Cortinarius impennoides 0.97/75 KT591609 Cortinarius impennoides KX355542 Cortinarius bivelus Biveli 0.99/100 KM085367 Cortinarius bivelus

1.0/100 AJ889977 Cortinarius torvus AY669668 Cortinarius torvus

1.0/100 KC608582 Cortinarius fructuodorus 1.00/99 NR 131827 Cortinarius fructuodorus TYPE 1.0/100 KM273104 Cortinarius niveotraganus Telamonia KM273106 Cortinarius niveotraganus KC608590 Cortinarius agathosmus FJ039571 Cortinarius venustus 1.00/90 GQ159921 Cortinarius venustus

1.0/100 JX407296 Cortinarius oulankaensis 1.0/100 NR 131822 Cortinarius oulankaensis TYPE

NR 120189 Cortinarius bovinus TYPE F u

1.0/100 KU953944 Cortinarius bovinus r 0.57/62 v ola e 1.0/100 NR 131789 Cortinarius neofurvolaesus TYPE KC608593 Cortinarius neofurvolaesus s 1.0/100 DQ139985 Cortinarius sordidemaculatus i 0.97/76 +

DQ139991 Cortinarius sordidemaculatus Bov i 1.00/94 NR 131788 Cortinarius anisatus TYPE

100/98 DQ117930 Cortinarius anisatus n 0.01 NR 131825 Cortinarius alboambitus TYPE i 1.0/100 KC608576 Cortinarius alboambitus Figure 1. The 50% majority rule consensus ITS rDNA phylogram of genus Cortinarius section Hinnulei and related taxa obtained in MrBayes. Nodes supported by >0.95 Bayesian PP or >70% ML BP are shown annotated.

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Table 1. Samples newly sequenced in this study used for the phylogenetic analyses.

GenBank Accession No. Source Identification ITS LSU Holotype SESLIGB 11292 = KATO - Fungi 3455 = Cortinarius Turkey, Trabzon, Maçka, MF696139 conicoumbonatus sp. nov. Sevinç neighborhood SESLIGB 11292 = KATO - Fungi 3455 = Cortinarius Turkey, Trabzon, Maçka, MF696140 conicoumbonatus sp. nov. Sevinç neighborhood SESLIGB 11133 = KATO - Fungi 3455 = Cortinarius Turkey, Trabzon, Maçka, MF696141 conicoumbonatus sp. nov. Sevinç neighborhood

Figure 2. Cortinarius conicoumbonatus: a and b- basidiomata, c and d- basidiospores (scale bars: a and b = 50 mm, c and d = 10 µm). Photos by E Sesli.

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Figure 3. Cortinarius conicoumbonatus: a and b- cross-sections through the , c and d- basidia, basidioles, and marginal cells (scale bars: a = 100 µm, b and c = 20 µm, d = 10 µm). Photos by E Sesli.

331 SESLİ and LIIMATAINEN / Turk J Bot injured or eaten by insects; longitudinally whitish fibrillose Photographica (C. hinnuleus (II) in the phylogenetic tree, on a dark reddish brown background; covered with a Figure 1) from Finland, France, Italy, and Sweden; and whitish woolly veil from the middle towards the base C. hinnuleoarmillatus from Finland, France, and Sweden and forming membranous annular zone. Pileus context (Brandrud et al., 1989; Bidaud et al., 1997; Niskanen et al., thin; grayish to beige, biscuit- to fawn-colored. Odor 2006; Li et al., 2016). Cortinarius badioflavidusdiffers from unpleasant. Basidiospores (6.7–)7.3–8.6(–9.1) × (4.2– C. conicoumbonatus by the overall red brown coloration, )4.6–5.8(–6.8) µm (n = 75 and Q = 1.3–1.7), on average green corn-like odor of lamellae and larger (8.1–10.5 × 7.9 × 5.2 µm; broadly ellipsoid to obovoid; moderately to 5.8–6.5 µm), broadly ellipsoid to broadly amygdaloid, strongly verrucose; always with a distinct apiculus; light to very coarsely verrucose basidiospores (Li et al., 2016). The dark ocher-yellow; moderately dextrinoid; spore deposit known distribution and ecology of C. badioflavidus also cinnamon-brown. Basidia clavate; (25–)30–35(–45) × are different. European C. hinnuleoarmillatus has orange (6.5–)8–9(–10.5) µm (n = 15); 2-4-spored; sterigmata red , which makes it a distinct species from triangular, up to 5 µm. Basidioles 20–25(–30) × 5.5–7.2(– other Hinnulei species (Niskanen et al., 2006, 2008a). 7.5) µm. Cystidia absent. Marginal cells (14–)9–23(–26) Another European species, C. hinnuleus s.s. Cortinarius × (4.5–)7–9.5(–10) µm, various, cylindrical to clavate and Flora Photographica, has strongly verrucose, subglobose thin-walled (n = 15). Pileipellis duplex: Epicutis made up to obovoid-subglobose basidiospores (Niskanen et al., of more or less cylindrical hyphae, 4–10 µm across, hyaline 2008a). The ITS sequence of C. conicoumbonatus is distinct to yellow. Hypocutis consists of cylindrical to enlarged from other members of the section Hinnulei and forms a (more cellular in shape) hyphae that are 15–30 µm wide. monophyletic clade. It has a sister group relationship with Clamp connections present in all tissues. C. hinnuleus s.s. M. Moser (as C. hinnuleus (I) in Figure Ecology and distribution: Gregarious in mixed 1) and differs from it by 1.5%. Thus, based on ITS region, spruce-beech forests in soil and debris, autumn. Known C. conicoumbonatus is a clear and distinct species and only from mixed spruce-beech forests of the East Black Sea suitable for barcode identification. Region of Turkey. Acknowledgments 4. Discussion The first author is thankful to the Karadeniz Technical Cortinarius conicoumbonatus belongs to subgen. Telamonia, University management for financial support (BAP: FAT- sect. Hinnulei. It can be distinguished from other closely 2017-7044). Our most sincere thanks are due to André related species by its basidiospores, which are broadly Bidaud (Dauphiné-Savoie Federation for Mycology and ellipsoid to obovoid, moderately to strongly verrucose, Botany, France) for his helpful comments. We are grateful and 7.3–8.6 × 4.6–5.8 µm in size. Morphologically similar to Dr Pablo Alvarado (Oviedo, Spain) and Tuba Oyur species (Table 2) are C. badioflavidus from Western Kurt (Triogen, İstanbul) for assistance with the molecular North America; C. hinnuleus s.s. Cortinarius Flora studies.

Table 2. Comparison of the diagnostic characteristics of Cortinarius conicoumbonatus with similar species (Brandrud et al., 1989; Bidaud et al., 1997; Niskanen et al., 2006; Li et al., 2016).

Species Pileus Stipe Basidiospores Basidia Habitat

Rounded conic, plano-convex, Cylindrical, yellowish cream, Broadly ellipsoid to umbonate, pale yellowish to light ochraceous, buff, reddish broadly amygdaloid, very Clavate, 4-spored, Mixed forests of oak, Cortinarius white-silky or red-brown, 20–60 to yellowish brown, coarsely verrucose, 8.1–10.5 × 29–31 × 8.7–9.2 µm fir, pine, willow badioflavidus mm 43–88 mm 5.8–6.5 µm

Hemispherical, convex to Cylindrical to clavate, curved, Cortinarius Broadly ellipsoid to obovoid, umbonate, gray-beige, light gray-beige, light ocher to Clavate, 2–4-spored, Gregarious in spruce- conicoumbonatus moderately to strongly v ocher to beige brown, beige-brown, 50–100 × 5–10 30–35 × 8–9 µm beech mixed forests errucose, 7.3–8.6 × 4.6–5.8 µm 25–50 mm mm

Conical to campanulate, convex Cylindrical, brown, ocher- to Cortinarius Subglobose to obovoid, strongly Gregarious in to plane, red-brown to ocher- red brown, light brown to Clavate, 4-spored, hinnuleus verrucose, hardwood forests and brown, or ocher-yellow, 20–70 gray-brown, 10–120 × 2–15 23–34 × 7.5–10 µm 7–8.5 × 5–6.6 µm dwarf-shrubs mm mm

Cylindrical to clavate, orange- Solitary or caespitose Cortinarius Conical to convex, orange to Obovoid to oblong ellipsoid, Clavate, 4-spored, red on a whitish background, in deciduous forests of hinnuleoarmillatus ochraceous brown, 30–50 mm verrucose, 8.8–10.1 × 5.4–6.3 µm 30–35 × 8–10 µm 60–100 × 5–10 mm oak, nut, and birch

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References

Akata I, Kabaktepe S, Akgül H (2015). Cortinarius caperatus (Pers.) Høiland K, Holst-Jensen A (2000). Cortinarius phylogeny and Fr., a new record for Turkish mycobiota. Kastamonu University possible taxonomic implications of ITS rDNA sequences. Journal of Forestry Faculty 15: 86-89. Mycologia 92: 694-710. Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ (1997). Basic Kaya A, Uzun Y, Karacan İH (2009). Macrofungi of Göksun local alignment search tool. J Mol Biol 215: 403-410. (Kahramanmaraş) District. Turk J Bot 33: 131-139. Ammirati JF (1989). Dermocybe, subgenus Dermocybe, section Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of Sanguineae in Northern California. Mycotaxon 34: 21-36. the Fungi. 10th ed. Wallingford, UK: CABI. Ammirati JF, Hughes KW, Liimatainen K, Niskanen T, Matheny PB Kytövuori I, Niskanen T, Liimatainen K, Lindstrom HK (2005). (2013). Cortinarius hesleri from eastern North America and Cortinarius sordidemaculatus and two new related species, related species from Europe and Western North America. C. anisatus and C. neofurvolaesus, in Fennoscandia Botany 91: 91-98. (, Agaricales). Karstenia 45: 33-50. Bidaud A, Moënne-Loccoz P, Reumaux P (1997). Atlas des Li GJ, Hyde KD, Zhao RL, Hongsanan S, Abdel-Aziz FA, Abdel- Cortinaires, Les Cortinaires Hinnuloides. La Roche-sur-Foron, Wahab MA, Alvarado P, Alves-Silva G, Ammirati JF, France: Federation Mycologique Dauphine-Savoie (in French). Ariyawansa HA et al. (2016). Fungal diversity notes 253-366: taxonomic and phylogenetic contributions to fungal taxa. Bidaud A, Moënne-Loccoz P, Reumaux P, Carteret X (2006). Atlas Fungal Divers 78: 1-237. des Cortinaires 16. La Roche-sur-Foron, France: Federation Mycologique Dauphine-Savoie (in French). Liimatainen K, Niskanen T, Dima B, Kytövuori I, Ammirati JF, Frøslev TG (2014). The largest type study of Agaricales species to date: Brandrud TE, Bendiksen E, Dima B (2015). Some new and little Bringing identification and nomenclature of Phlegmacium known telamonioid Cortinarius species from Norway. Agarica (Cortinarius) into the DNA era. Persoonia 33: 98-140. 36: 11-42. Moser MM (2002). Studies in the North American Cortinarii 7. Brandrud TE, Lindström H, Marklund H, Melot J, Muskos S New and interesting species of Cortinarius subgen. Telamonia (1989). Cortinarius Flora Photographica I. Matfors, Sweden: (Agaricales, Basidiomycotina) from the Rocky Mountains. Cortinarius HB. Feddes Repertorium 113: 48-62. Clémençon H (2009). Methods for Working with Macrofungi: Moser MM, McKnight KH, Ammirati JF (1995). Studies on North Laboratory Cultivation and Preparation of Larger Fungi for American Cortinarii 1. New and interesting taxa from the Light Microscopy. Berchtesgaden, Germany: Berchtesgadener greater Yellowstone area. Mycotaxon 55: 301-346. Anzeiger. Murray MG, Thompson WF (1980). Rapid isolation of high Cubeta MA, Echandi E, Abernethy T, Vilgalys R (1991). molecular weight plant DNA. Nucleic Acids Res 8: 4321-4325. Characterization of anastomosis groups of binucleate Rhizoctonia species using restriction analysis of an amplified Niskanen T, Kytövuori I (2012). Key F: subgenus Telamonia sects ribosomal RNA gene. Phytopathology 81: 1395-1400. Bovini, Illumini, Saturnini, Sciophylli, Subbalaustini and Sordescentes. In: Knudsen H, Vesterholt J, editors. Funga Demirel K, Erdem Ö, Uzun Y, Kaya A (2010). Macrofungi of Hatila Nordica, 2nd revised edition. Agaricoid, Boletoid, Clavarioid, Valley National Park (Artvin, Turkey). Turk J Bot 34: 457-465. Cyphelloid and Gastroid Genera. Copenhagen, Denmark: Dima B, Liimatainen K, Niskanen T, Kytovuori I, Bojantchev D Nordsvamp Press, pp. 847-856. (2014). Two new species of Cortinarius, subgenus Telamonia, Niskanen T, Kytövuori I, Bendiksen EK, Brandrud TE, Frøslev TG, sections Colymbadini and Uracei, from Europe. Mycol Prog 13: Hoiland K, Jeppesen TS, Liimatainen K, Lindström H (2008a). 867-879. Cortinarius (Pers.) Gray in: Knudsen H, Vesterholt J, editors. Gardes M, Bruns TD (1993). ITS primers with enhanced specificity Funga Nordica. Copenhagen, Denmark: Nordsvamp, pp. 661- for Basidiomycetes. Application to the identification of 778. mycorrhizae and rusts. Mol Ecol 2: 113-118. Niskanen T, Liimatainen K, Ammirati JF (2013). Five new Telamonia Garnica S, Weiss M, Oberwinkler F (2005). A framework for species (Cortinarius, Agaricales) from Western North America. a phylogenetic classification in the genusCortinarius Botany 91: 478-485. (Basidiomycota, Agaricales) derived from morphological and Niskanen T, Liimatainen K, Kytövuori I (2006). Taxonomy, molecular data. Can J Bot 83: 1457-1477. ecology and distribution of Cortinarius rubrovioleipes Harrower E, Ammirati JF, Cappuccino AA, Ceska O, Kranabetter and C. hinnuleoarmillatus (Basidiomycota, Agaricales) in JM, Kroeger P, Lim S, Taylor T, Berbee ML (2011). Cortinarius Fennoscandia. Karstenia 46: 1-12. species diversity in British Columbia and molecular Niskanen T, Liimatainen K, Kytövuori I, Lindström H (2008b). phylogenetic comparison with European specimen sequences. Subgen. Telamonia in: Knudsen H, Vesterholt J, editors. Funga Botany 89: 799-810. Nordica. Copenhagen, Denmark: Nordsvamp, pp. 721-778.

333 SESLİ and LIIMATAINEN / Turk J Bot

Nylander JAA (2004). MrModeltest v2. Program Distributed by Smith AH (1939). Studies in the Genus Cortinarius I. Contributions the Author. Uppsala, Sweden: Evolutionary Biology Centre, from the University of Michigan Herbarium. Ann Arbor, MI, Uppsala University. USA: University of Michigan. Peintner U, Moser MM, Thomas KA, Manimohan P (2003). First Stamatakis A (2006). RAxML-VI-HPC: Maximum likelihood-based records of ectomycorrhizal Cortinarius species (Agaricales, phylogenetic analyses with thousands of taxa and mixed Basidiomycetes) from tropical India and their phylogenetic models. Bioinformatics 22: 2688-2690. position based on rDNA ITS sequences. Mycol Res 107: 485- Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S 494. (2011). MEGA5: Molecular evolutionary genetics analysis Ronquist F, Huelsenbeck JP (2003). MrBayes 3: Bayesian phylogenetic using maximum likelihood, evolutionary distance, and inference under mixed models. Bioinformatics 19: 1572-1574. maximum parsimony methods. Mol Biol Evol 28: 2731-2739. Sesli E (2006). New records of Tricholomataceae and Cortinarius Uzun Y, Akata I, Acar İ, Akçay ME (2013). Three new records for (Pers.) Gray from Turkey. Turk J Bot 30: 59-62. Turkish Cortinarius from Bingöl province. Biological Diversity and Conservation 3: 160-163. Sesli E, Contu M, Vila J, Moreau PA, Battistin E (2015). Taxonomic studies on some agaricoid and boletoid fungi of Turkey. Turk Vilgalys R, Hester M (1990). Rapid genetic identification and J Bot 39: 134-146. mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J Bacteriol 172: 4238-4246. Sesli E, Denchev CM (2008). Checklists of the myxomycetes, larger ascomycetes and larger basidiomycetes in Turkey. Mycotaxon White TJ, Bruns T, Lee S, Taylor JW (1990). Amplification and direct 106: 65-67. sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ, editors. PCR Sesli E, Moreau PA (2015). Taxonomic studies on some new fungal Protocols: A Guide to Methods and Applications. New York, records from Trabzon, Turkey. Turk J Bot 39: 857-866. NY, USA: Academic Press, pp. 315-322.

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