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Turkish Journal of Turk J Bot (2018) 42: 224-232 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1706-64

A new of , H. yadigarii sp. nov. (), with an isolated systematic position within the from the Colchic part of Turkey

1, 2 3 Ertuğrul SESLİ *, Vladimír ANTONÍN , Marco CONTU 1 Department of Biology Education, Faculty of Education, Karadeniz Technical University, Trabzon, Turkey 2 Moravian Museum, Department of Botany, Brno, Czech Republic 3 Via Marmilla, 12 (I-Gioielli 2), I-07026 Olbia (OT), Italy

Received: 27.06.2017 Accepted/Published Online: 20.11.2017 Final Version: 20.03.2018

Abstract: Hygrophorus yadigarii (Hygrophoraceae/) is described as a new species for science based on basidiomata collected from Maçka, Trabzon, Turkey. The new taxon is quite different even from the closest relatives, easily distinguished by the other species because of its grayish to ash-colored, gregarious to subcaespitose, sticky basidioma; a slightly umbonate to depressed pileus; a cylindrical to clavate, grayish ; ellipsoid and smooth ; quite long basidia; clavate, cylindrical or narrowly utriform, apically pyriform or strangulated cheilocystidia; and gelatinous . A description with field and micromorphological illustrations, a phylogenetic tree, a simple key, a comparison chart including similar species, and a short discussion are provided.

Key words: Colchic, Hygrophorus, Maçka, new species, Trabzon

1. Introduction Sesli and Denchev, 2008; Akata and Doğan, 2015; Uzun The genus Hygrophorus belongs to the family et al., 2017), and it is characterized by its tricholomatoid, Hygrophoraceae, proposed by Roze (1876). Singer (1986) collybioid, clitocyboid, or omphalinoid small to large, recognized seven genera in that family: Hygrophorus, thin to fleshy, dry to very glutinous or viscid basidioma; a Neohygrophorus Singer (= ), Hygrotrama whitish, dull-colored, gray, brownish, yellowish, orange, or Singer (= ), Camarophyllus (Fr.) reddish dry to fairly sticky pileus; closely or widely spaced, P.Kumm., (Fr.) P.Kumm., Hygroaster Singer (= adnate to decurrent, typically thick and waxy, generally Hygrocybe, see below), and Omphaliaster D.Lam. whitish, sometimes yellowish or pinkish lamellae; a dry Recent molecular studies have shown that many to glutinous, glabrous or fibrillose, generally pruinose known entities can be classified in Hygrophoraceae such or granulose stipe; an often thick, sometimes reddening as P.M.Jørg.; Redhead, or yellowing whitish content; and long, narrowly clavate Lutzoni, Moncalvo & Vilgalys; Fr.; basidia and smooth, hyaline, nonamyloid basidiospores Singer; H.E.Bigelow & A.H.Sm.; (Singer, 1986; Boertmann, 1995; Young, 2005; Kovalenko, Redhead, Ammirati & Norvell; 2012). Clémençon; Fr.; Corella Vain.; (Donk) According to the monograph by Hesler and Smith Bon; D.A.Reid; Reinsch; (1963), section Hygrophorus is divided into 8 subsections Eonema Redhead, Lücking & Lawrey; Herink; due to the macromorphology of the basidioma: subsect. Kotl. & Pouzar; (Singer) Singer; Chrysodontini Singer (with yellow granules on the Hygroaster Singer; Hygrocybe (Fr.) P.Kumm.; Hygrophorus; basidioma), subsect. Pallidini A.H.Sm. & Hesler (basidioma Redhead, Lutzoni, Moncalvo & Vilgalys; slightly greasy), subsect. Hygrophorus (basidioma distinctly Neohygrocybe Herink; Bresinsky; and slimy), subsect. Erubescentes A.H.Sm. & Hesler (basidioma (Singer) Singer (Boertman, 1995; reddish), subsect. Fulvoincarnati A.H.Sm. & Hesler Lodge et al., 2014). (basidioma more or less orange colored), subsect. Discoidei Hygrophorus Fr. is a large genus with more than 900 (Bataille) Konrad & Maubl. (basidioma yellowish, yellow- records worldwide (Kirk et al., 2008), represented by 30 brown to reddish), subsect. Olivaceoumbrini Bataille species in Turkey (Demirel et al., 2003; Doğan et al., 2007; (basidioma partly slimy and usually shiny), and subsect.

* Correspondence: [email protected] 224 SESLİ et al. / Turk J Bot

Tephroleuci Bataille (not typically viscid). Bresinsky and 2.2. Sequence alignment and phylogenetic analysis Huber (1967), Arnolds (1986), Singer (1986), Bon (1990), BLAST (Altschul et al., 1997) was used to select the most Horak (1990), Candusso (1997), and Kovalenko (2012), closely related 28S rDNA sequences from INSD public among others, proposed other slightly different taxonomic databases. Sequences came mainly from Lutzoni (1997), arrangements, but all recognized sect. Hygrophorus for a Larsson et al. (2004), Matheny et al. (2006), Geml et al. white species with a slimy stem surface. (2012), Lucking et al. (2013), and Lodge et al. (2014), In this paper, a new species belonging to the genus among others. Sequences were first aligned in MEGA 5.0 Hygrophorus collected in Turkey, Hygrophorus yadigarii, software (Tamura et al., 2011) with its Clustal W application sp. nov., is fully described and discussed. and then corrected manually. The final alignment included 352/700 variable sites. Final alignment was subjected 2. Materials and methods to MrModeltest 2.3 (Nylander, 2004) in PAUP* 4.0b10. The basidiomata ofHygrophorus yadigarii were collected Model GTR+I+G was selected and implemented in from the Sevinç neighborhood of Maçka, Trabzon, Turkey, MrBayes 3.1 (Ronquist and Huelsenbeck, 2003), where on 3 November 2016. Color slides of the basidiomata were a Bayesian analysis was performed (data partitioned, taken at the collection site. The pileus of one basidioma two simultaneous runs, six chains, temperature set to was used to obtain a print and the others were dried 0.2, sampling every 100th generation) until convergence for future microscopic studies. Very thin microscopic parameters were met after 3.38M generations, standard sections were obtained from the basidiomata, treated deviation having fallen below 0.01. The first 25% of trees in concentrated ammonia solution and subsequently in were discarded as burn-in. Finally, a full search for the Congo red, and finally investigated under a Zeiss Axio best-scoring maximum likelihood tree was performed Imager A2 trinocular research microscope (Clémençon, in RAxML (Stamatakis, 2006) using the standard search 2009). Microscopic structures such as basidiospores, algorithm (2000 bootstrap replications). The significance basidia, and cystidia were measured with Axio Imager threshold was set above 0.95 for posterior probability (PP) software. The holotype materials are kept at the herbarium and 70% bootstrap proportions (BP). of the Fatih Faculty of Education at Karadeniz Technical University (KATO), Trabzon, Turkey. 3. Results 2.1. DNA extraction, PCR amplification, and DNA 3.1. Molecular analysis sequencing Topology obtained with 28S rDNA failed to resolve Total DNA was extracted from the dry specimens only the most suprageneric relationships among the employing a modified protocol based on Murray and Hygrophoraceae genera included in the analysis, in Thompson (1980). A portion of each sample was blended contrast with the 4-gene analysis by Lodge et al. (2014), with the aid of a micropestle in 600 µL of CTAB buffer although it was enough to support the monophyletic (CTAB 2%, NaCl 1.4 M, 20 mM EDTA pH 8.0, 100 mM status of most of these genera. Hygrocybe and Hygroaster Tris-HCl pH 8.0). The resulting mixture was incubated were not discriminated by 28S rDNA analysis, while no for 15 min at 65 °C. A similar volume of chloroform and isoamyl alcohol (24:1) was added and carefully mixed with significant support for the monophyletic status of the the samples until their emulsion. It was then centrifuged Lichenomphaloideae genera Arrhenia and for 10 min at 13,000 × g, and the DNA in the supernatant was found, and only partial support for Dictyonema and was precipitated with a volume of isopropanol. After a Lichenomphalia was recovered, in agreement with the new centrifugation of 15 min at the same speed, the pellet multigenic analysis of Lodge et al. (2014). The 28S rDNA was washed in cold ethanol 70%, centrifuged again for 2 sequence obtained from the sample KATO - Fungi 3843 min, and dried. It was finally resuspended in 200 µL of (holotype of the new species Hygrophorus yadigarii) was found to be significantly related to the genus Hygrophorus, ddH2O. PCR amplification was performed with primers ITS1F and ITS4 (White et al., 1990; Gardes and Bruns, but no significant similarity with the sections delimited by 1993) for the ITS region, while LR0R and LR5 (Vilgalys Lodge et al. (2014) could be found, suggesting that it might and Hester, 1990; Cubeta et al., 1991) were used to amplify represent a new one. the 28S rDNA region. PCR reactions were performed 3.2. under a program consisting of a hot start at 95 °C for Hygrophorus yadigarii E. Sesli, Antonín & Contu, sp. nov. 5 min, followed by 35 cycles at 94 °C, 54 °C, and 72 °C (Figures 1–5) (45, 30, and 45 s, respectively) and a final 72 °C step for GenBank Acc. No: MF370228 10 min. PCR products were checked on 1% agarose gels, MycoBank No: MB 821862 and positive reactions were sequenced with one or more Diagnosis: Basidiomata tricholomatoid, gregarious to PCR primers. Chromatograms were checked searching for subcaespitose in mixed forests, medium sized, fairly sticky; putative reading errors, and these were corrected. the surface of pileus grayish, slightly yellowish at some

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Figure 1. Hygrophorus yadigarii: a, b, and c- basidiomata (scale bars: 30 mm). Photos by E Sesli. places (probably slime layer leaving yellowish stains in are raised up in maturity; margin incurved for a long age), margin often cleft; center usually slightly umbonate; time; the surface grayish, ash-colored, slightly yellowish at stipe grayish and lamellae slightly bluish white. some places (probably slime layer leaving yellowish stains Holotype: Turkey, Trabzon, Maçka, Sevinç in age); slightly fibrillose to floccose. Lamellae adnate; neighborhood, 40°50′51.06″N, 39°37′41.07″E, 751 m alt., becoming subdistant or subdecurrent in age; whitish to 03.11.2016, leg. E. Sesli (KATO - Fungi 3843). creamy, slightly bluish; broad; thick; waxy; L = 35–45, I = Etymology: The specific epithet honors the father of 1–3. Stipe 40–70 mm long and 5–10 mm wide; cylindrical the first author (Yadigar Sesli), who died in 2016. to clavate; grayish; longitudinally fibrous; somewhat larger Pileus 20–60 mm wide; hemispherical when young, toward the base; sometimes flattened; base coated with later convex to plane; slightly depressed in the center; white ; sometimes twisted or compressed and somewhat irregular, sometimes cleft and undulating; grooved. Context slightly bluish white and soft. Odor and umbo indistinct or broadly hill-shaped; surface very taste not distinctive. slimy-lubricous when moist, viscid when dry; the edges

226 SESLİ et al. / Turk J Bot C yp h e

AY586715 Suillus luteus e llost 1.00/99 KF443243 Cyphellostereum imperfectum KF443244 Cyphellostereum phyllogenum Di c r

KF443254 Dictyonema spongiosum e

1.00/100 u ty o

KF443253 Dictyonema sericeum m KF443247 Dictyonema irpicinum n

EU825957 Dictyonema glabratum e 1.00/93 KF443264 Cora strigosa m 1.00/77 KF443262 Cora reticulifera a 1.00/98 KF443261 Cora pavonia Cora 1.00/100 EU825969 Cora minor 1.00/93 U66428 Arrhenia auriscalpium U66429 EU118604 Arrhenia retirugis 0.98/90 U66448 Arrhenia obscurata DQ071732 Arrhenia auriscalpia Arrhenia 1.00/100 1.00/90 KP965784 U66453 Arrhenia sphagnicola 0.97/82 1.00/99 U66449 Arrhenia philonotis U66442 U66455 Arrhenia velutipes JQ065875 Lichenomphalia hudsonia 1.00/99 GU811053 Lichenomphalia alpina Lichenomphalia GU811047 Lichenomphalia sp. AF261445 Lichenomphalia umbellifera MF370228 SESLI 3843 Hygrophorus yadigarii sp. nov. AF042562 Hygrophorus sordidus 1.00/83 AY586663 Hygrophorus KF291210 Hygrophorus poetarum AY586660 0.99/72 KF291214 Hygrophorus pustulatus DQ457678 AY586661 1.00/100 KF291208 Hygrophorus lucorum 1.00/97 KF381557 Hygrophorus 1.00/94 HM026551 Hygrophorus fuligineus AY635769 Hygrophorus flavodiscus 1.00/100 AY586662 Hygrophorus olivaceoalbus AF430280 Hygrophorus leucophaeus AF430279 KF381556 Hygrophorus discoxanthus KF381555 1.00/100 KF291050 Chrysomphalina grossula 1.00/82 EU652372 Chrysomphalina grossula Chrysomphalina DQ457656 Chrysomphalina chrysophylla KF381546 1.00/97 1.00/69 AF261446 Gliophorus laetus Gliophorus KF381545 Gliophorus graminicolor 1.00/98 KF291111 Humidicutis sp. DQ457672 Humidicutis auratocepha 1.00/90 KF291145 Humidicutis marginata Humidicutis 1.00/94 AF042580 Humidicutis marginata KF291239 Porpolomopsis lewelliniae 1.00/100 KF291243 Porpolomopsis calyptriformis Porpolomopsis 1.00/100 KF291234 Neohygrocybe ovina 0.99/70 KF291137 Neohygrocybe subovina Neohygrocybe KF291226 Neohygrocybe ingrata KF291073 Chromosera citrinopallida 1.00/93 DQ457655 Chromosera 1.00/100 EF561625 Hygroaster nodulisporus 1.00/100 KF291171 Hygrocybe andersonii EF551314 Hygroaster albellus KF291134 1.00/87 KT020862 Hygrocybe firma KF381554 Hygrocybe substrangulata 1.00/99 AF261452 0.99/72 KF291109 Hygrocybe ceracea 1.00/100 KF291189 Hygrocybe parvula Hygrocybe KF291159 1.00/100 KF291155 Hygrocybe melleofusca Hygroaster 1.00/83 KF291183 Hygrocybe helobia 1.00/93 KF291190 Hygrocybe pseudoconica EU522785 1.00/100 AF261451 Hygrocybe spadicea KF306330 Hygrocybe konradii AM946460 0.05 KF291193 Hygrocybe purpureofolia

Figure 2. The 50% majority rule consensus 28S rDNA phylogram of the Lichenomphalioideae, Hygrophoroideae, and Hygrocyboideae lineages of the Hygrophoraceae obtained in MrBayes from 25,350 sampled trees. Nodes supported by 0.95 Bayesian PP or 70% ML BP are shown annotated.

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Figure 3. Hygrophorus yadigarii: a, b, and c- cross-sections through the (scale bars: 10 µm). Photos by E Sesli.

Basidiospores (6.5–)7.5–8.5(–9.5) × (4.3–)4.5– a gelatinous matter. Subpellis consists of elongate hyphae 5.5(–6.3) µm (n = 54 and Q = 1.5–1.7), on average 8.1 8.5–20 µm wide. Clamp connections present in all tissues. × 5.2 µm (basidiospores rarely very large, e.g., 16 × 9 Ecology and distribution: Gregarious to subcaespitose µm from 2-spored basidia); ellipsoid; hyaline; smooth; in hornbeam- dominated forest, on soil, in debris, always with drops; with a distinct apiculus; medium to under hornbeam. Autumn. Known only from the Colchic thin-walled; greenish in water and pale yellow in Congo part of Turkey. red. Spore deposit white. Basidia slenderly clavate; 55–70(–72) × (7.5–)8–9(–9.5) µm (n = 15); 2–4-spored; 4. Discussion some with siderophilous granules; sterigmata long Hygrophorus yadigarii can be distinguished from other (5–7.5 µm). Basidioles 45–60(–62) × 5.5–7.5(–7.7) µm. closely related species because of its tricholomatoid, Cheilocystidia present; 30–60(–70) × (3.9–)4.5–7.5(–9) grayish to ash-colored, gregarious to subcaespitose, µm; variable; consisting of clavate, cylindrical or narrowly fairly sticky basidioma; a slightly umbonate to depressed, utriform, apically pyriform, strangulated, sinuous clavate; somewhat irregular, 20–60 mm wide pileus; a cylindrical to slightly thick- to thin-walled; hyaline elements (n = 15). clavate, grayish, longitudinally fibrillose, 40–70 mm stipe; Pleurocystidia absent. Pileipellis made up of irregular to ellipsoid, hyaline, smooth, 8.1 × 5.2 µm basidiospores; parallel hyphae; (2.8–)4–8(–12.3) µm across; occasionally slenderly clavate and very long (55–70 µm) basidia; branched; some hyphal ends ascending and embedded in clavate, cylindrical or narrowly utriform, apically pyriform

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Figure 4. Hygrophorus yadigarii: a- basidiospores, b and c- pileipellis (scale bars: a = 10 µm, b and c = 20 µm). Photos by E Sesli. or strangulated cheilocystidia; and a gelatinous pileipellis lamellae; larger (8.5–12 × 5.5–7.5 µm), ellipsoid to oblong (Table). ovoid basidiospores (Bon, 1990; Kovalenko, 2012). Morphologically close species are H. tephroleucus The other morphologically similar species from (Pers.) Fr., H. mesotephrus Berk. & Broome, and H. , H. pustulatus, has a smaller (15–35 mm), slightly pustulatus (Pers.) Fr. from Europe and H. fuligineus Frost, viscid, scaly, gray brown pileus; subdecurrent lamellae; H. occidentalis A.H.Sm. & Hesler, and H. virgatulus Peck smaller (50–65 × 7–9 µm) basidia; and lacks cheilocystidia from North America (Hesler and Smith, 1963; Bon, 1990; (Bon, 1990; Breitenbach and Kränzlin, 1991). Kovalenko, 2012). Hygrophorus fuligineus has a larger (40–120 mm), Hygrophorus tephroleucus differs from H. yadigarii by blackish, clove brown to brownish olive, grayish brown or the gray brown to dark brown pileus, decurrent lamellae, olive-gray pileus; a longer stipe (40–100 mm); and lacks a whitish stipe, larger (9–11 × 5–6 µm) basidiospores, cheilocystidia. Hygrophorus occidentalis is different from and smaller basidia (40–60 × 6–8 µm). Hygrophorus our new species by its slightly larger (20–80 mm) and mesotephrus differs by a smaller (20–30 mm), bell-shaped variably colored (hairs brown to fuscous, yellowish or or convex, pale grayish brown pileus; decurrent and white smoky, white to pale cinereous) pileus; smaller (6–8 × 3.5–

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Figure 5. Hygrophorus yadigarii: a, b, and c- cheilocystidia, d- basidia, basidioles, and cheilocystidia (scale bars: 10 µm). Photos by E Sesli.

5 µm) basidiospores; and the absence of cheilocystidia. 11 × 4.5–6.2 μm) basidiospores; and grows in coniferous Another close species from North America, H. virgatulus, forests (Hesler and Smith, 1963; Bon, 1990; Kovalenko, has a smaller (25–50 mm), whitish-brownish pileus; 2012). Another molecularly close species, H. pustulatus, narrower (3.5–5 µm) basidiospores; and smaller basidia has a smaller (20–45 mm), convex, at times papillate, ashy (32–54 × 5–7 µm) (Hesler and Smith, 1963). to darker brownish, fibrillose pileus; somewhat decurrent, Genetically, 28S rDNA data suggest that H. yadigarii bluntly adnate, close to subdistant, pure white lamellae; a is different from all records in public databases. BLAST dark-gray punctate stipe; and lacks cheilocystidia (Hesler top matches are Hygrophorus hyacinthinus, H. pustulatus, and Smith, 1963; Bon, 1990). Hygrophorus agathosmus and H. agathosmus (95% similarity), although ITS data differs from H. yadigarii with its larger (40–80 mm), ashy are almost 20% different from all other Hygrophorus ITS gray, glutinous to viscid, glabrous pileus; white to sordid sequences available. grayish lamellae; a whitish to pale ashy, solid, dry or moist, Hygrophorus hyacinthinus has a larger (20–100 mm), evenly fibrillose pruinose stipe; larger (8–10.5 × 4.5–5.5 grayish to dirty white pileus; a taller (50–100 mm), dry, μm) basidiospores; smaller basidia (48–65 × 6–8 μm); smooth, naked, dirty white or light gray stipe; larger (7.5– and lack of cheilocystidia (Hesler and Smith, 1963; Bon,

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Table. Comparison of the diagnostic characteristics of Hygrophorus yadigarii with similar species (Hesler and Smith, 1963; Bon, 1990; Breitenbach and Kränzlin, 1991; Boertman, 1995; Kovalenko, 2012).

Character H. fuligineus H. mesotephrus H. occidentalis H. pustulatus H. tephroleucus H. virgatulus H. yadigarii

Blackish brownish, Hair brown, fuscous, Pallid-cinereous, Whitish with a Pale drab, cinereous, Ashy with a darker Grayish, grayish brown or yellowish, smoky, dark ashy, gray, brownish buff, avellaneous brownish disc ash-colored olive-gray white pale ashy gray disc Pileus 40–120 mm 15–40 mm 20–100 mm 20–45 mm 10–30 mm 25–50 mm 20–60 mm

Concolorous White White Whitish White White Grayish with the pileus Stipe 40–100 mm 40–70 mm 20–70 mm 60–90 mm 40–60 mm 60–100 mm 40–70 mm

Spores 7–9 × 4.5–5.5 µm 8–11 × 5–6 µm 6–8 × 3.5–5 µm 7–9 × 4–5 µm 8–10 × 4–5 µm 7–9 × 3.5–5 µm 7.5–8.5 × 4.5–5.5 µm

4-spored, 4-spored, 45–60 × 4-spored, 2- and 4-spored 4-spored, 4-spored, 2- and 4-spored, Basidia 38–58 × 5–9 µm 8–10 µm 29–43 × 6–8 µm 46–61 × 6–8 µm 40–58 × 6–8 µm 32–54 × 5–7 µm 55–70 × 8–9 µm

In coniferous and Under fir and r In debris under Habitat Under In mixed oak- Under In open woods mixed woods edwood hornbeam

1990; Breitenbach and Kränzlin, 1991). According to some 3* Stipe hair brown, fuscous, yellowish, smoky or white, studies, Hygrophorus agathosmus (Fr.) Fr. and Hygrophorus 20–70 mm...... H. occidentalis hyacinthinus Quél. are the same taxon, but this needs to be 4 Basidiospores up to 10 µm...... 5 confirmed with further studies (Kirk et al., 2008). Phylogenetic analysis of 28S rDNA did not recover a 4* Basidiospores smaller...... 6 significant relationship between H. yadigarii and any of the 5 Basidia 40–58 × 6–8 µm, under conifers...... other Hygrophorus sections as defined by Lodge et al. (2014), ...... H. tephroleucus maybe because of the monogenic approach employed here, 5* Basidia 45–60 × 8–10 µm, under oak...... H. mesotephrus or else because H. yadigarii actually represents a different, independent section within Hygrophorus. To test this 6 Basidia 2- and 4-spored...... 7 hypothesis, a thorough multigenic review of Hygrophorus 6* Basidia 4-spored, in open woods...... H. virgatulus should be conducted. 7 Under fir and redwood...... H. pustulatus 4.1. A simple key to the Hygrophorus species close to H. yadigarii 7* Under hornbeam...... H. yadigarii We prepared a simple key below including the close relatives of Hygrophorus yadigarii to date. The key is in Acknowledgments accordance with those of Hesler and Smith (1963), Bon The first author is thankful to the Karadeniz Technical (1990), Breitenbach and Kränzlin (1991), Boertman University management for financial support (BAP: (1995), and Kovalenko (2012). 11300). Our most sincere thanks are due to Dr Deborah Jean Lodge (USA) and Dr David Boertmann (Denmark) for their very useful comments. We are grateful to Dr Pablo 1 Pileus up to 100–120 mm...... 2 Alvarado for assistance with the phylogenetic tree. The 1* Pileus smaller...... 4 studies of the second author were enabled by the support 2 Under beech, stipe whitish to pale ash...H. agathosmus provided to the Moravian Museum by the Ministry of Culture of the Czech Republic as part of its long-term 2* Under other trees...... 3 conceptual development program for research institutions 3 Stipe white, 40–100 mm...... H. fuligineus (DKRVO, Ref. MK000094862).

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