Fungal Biology 123 (2019) 431e447

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Fungal Biology

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Taxonomic update of Clitocybula sensu lato with a new generic classification

* Vladimír Antonín a, , Jan Borovicka b, Jan Holec c, Andrej Piltaver d, Miroslav Kolarík e a Moravian Museum, Department of Botany, Zelný trh 6, CZ-659 37, Brno, Czech Republic b Institute of Geology, Czech Academy of Sciences, Rozvojova 269, CZ-165 00 Prague 6, Czech Republic c National Museum, Mycological Department, Cirkusova 1740, CZ-193 00 Praha 9, Czech Republic d Velika vas 17, SLO-1262 Dol pri Ljubljani, Slovenia e Institute of Microbiology, Czech Academy of Sciences, Vídensk a 1083, CZ-142 20, Praha 4, Czech Republic article info abstract

Article history: The taxonomy and phylogeny of the hydropoid clade (genera Clitocybula s.l., Megacollybia, Leucoinocybe Received 29 August 2018 gen. nov., Hydropus, Trogia, Gerronema, Porotheleum and Lignomphalia gen. nov.) in Europe is studied Received in revised form using morphological and molecular approaches; the first three genera in detail including all known 14 February 2019 European species. Only two European species remain in Clitocybula s.str., Clitocybula lacerata and Clito- Accepted 19 March 2019 cybula familia. The European C. lacerata is a species complex which should be treated as C. lacerata agg. at Available online 24 March 2019 the current state of knowledge. A neotype originating from type area was designated to fix the appli- Corresponding Editor: Nik Money cation of the name. The presence of American species Clitocybula abundans in Europe is insufficiently proved. “Clitocybula dryadicola ˮ belongs to the genus Hydropus, and Clitocybula tilieti has an unclear Keywords: systematic position. The results showed that Megacollybia and Leucoinocybe represent independent Europe genera separated from Clitocybula. The genus Leucoinocybe is validly published with two European Leucoinocybe species, Leucoinocybe lenta and Leucoinocybe taniae. “Clitocybula flavoaurantia” proved to be conspecific Lignomphalia with the latter species. The genus Lignomphalia is published for “Pseudoomphalina lignicola”, a lignicolous nrITS omphalinoid species. The Indian “Clitocybula sulcata” is transferred to Leucoinocybe and “Clitocybula nrLSU atrialba” to Gerronema. The first European records of Megacollybia marginata are published. Phylogeny © 2019 British Mycological Society. Published by Elsevier Ltd. All rights reserved.

1. Introduction restricted it to one species (Clitocybula lacerata e type species) only. Bigelow (1973) characterized this genus as follows (shortened): The genus Clitocybula (Singer) Metrod is a rather small agaricoid “Basidiocarps lignicolous, usually gregarious or caespitose. Pilei genus with c. 15 really existing species (Kirk et al., 2008), and convex to plane or depressed to broadly infundibuliform, margin slightly more than 30 specific names published worldwide (www. incurved at first, surface radiate fibrillose to squamulose at least mycobank.org; accessed June 21, 2018). It was originally near disc. Lamellae usually adnate or decurrent. Stipe relatively described as a subgenus of Fayodia Kühner by Singer (1943), and slender, fibrous or tough or brittle. Spores amyloid, smooth, globose transferred to the generic level by Metrod (1952). Singer (1943) or subglobose or elliptic or ovate, deposit white or pale creamy. characterized it by having amyloid, smooth or asperulate slightly Pileocystidia and caulocystidia present, cheilocystidia present in thick-walled basidiospores without a perispor, rare or absent some species but never pleurocystidia. Pileus: oleiferous hyphae cheilocystidia, absent pleurocystidia, and habitus of small Clitocybe present; clamp connections present; cellular hypoderm absent. species. He only included “Fayodia lacerata ˮ and “Fayodia tilieti ˮ in Hymenophoral trama of parallel or subparallel hyphaeˮ. Singer this subgenus. Metrod (1952) added the character of large, distant, (1979) published the key for the identification of Clitocybula spe- veined, adnate to decurrent lamellae, and a lignicolous habitat. He cies, based on both macroscopical (pileus colour, width and attachment of lamellae, stipe surface) and microscopical (basidio- spore size, presence or absence and a shape of cheilocystidia) * Corresponding author. Fax: þ420 533 435 325. characters, and ecology (growth conifers or broadleaved trees). E-mail addresses: [email protected] (V. Antonín), [email protected] Later, Singer (1962) enlarged the generic characteristics for taxa (J. Borovicka), [email protected] (J. Holec), [email protected] (A. Piltaver), with small clitocyboid, but also omphalioid, collybioid, mycenoid [email protected] (M. Kolarík). https://doi.org/10.1016/j.funbio.2019.03.004 1878-6146/© 2019 British Mycological Society. Published by Elsevier Ltd. All rights reserved. 432 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

Table 1 List of sequences used in the phylogenetical analyses.

Species Specimen voucher Origin ITS (Sequence LSU (Sequence Reference ID) ID)

Baeospora myosura AFTOL-ID 1799, PBM 2748 (CUW) Massachusetts USA DQ484063 DQ457648 Matheny et al. (2006) Clitocybe ulmicola TENN 029208 (HOLOTYPE) USA NR_119887 HQ179668 Matheny et al. (2006) Clitocybula “abundans” PRM 922903 Czech Rep. LT854014 LT854014 this study Clitocybula “abundans” PRM 899161 Slovakia LT854015 LT854015 this study Clitocybula “abundans” PRM 899168 Slovakia LT854016 LT854016 this study Clitocybula aff. lacerata GRSM 77072 USA FJ596916 e Hughes et al. (2009) Clitocybula atrialba AFTOL-ID 1529, PBM 1250 (WTU) Washington USA DQ192179 DQ457659 Matheny et al. (2006) Clitocybula familia PRM 935966 Canada e LT854033 this study Clitocybula familia PRM 921866 Czech Rep. JF730327 JF730320 Antonín et al. (2011) Clitocybula familia BRNM 736053 Slovakia JF730328 JF730323 Antonín et al. (2011) Clitocybula flavoaurantia IB, Contu 18-9-2002 Italy LT854029 LT854058 this study Clitocybula flavoaurantia D Italy HM191743 e Malysheva et al. (2011) Clitocybula flavoaurantia GDOR (TYPE) Italy HM191744 e Malysheva et al. (2011) Clitocybula flavoaurantia LE 262757 Russia HM191745 e Malysheva et al. (2011) Clitocybula lacerata PRM 951559 (NEOTYPE) Slovenia MK713541 e this study Clitocybula lacerata PRM 915404 Czech Rep. LT854030 LT854054 this study Clitocybula lacerata WU 19575 Austria LT854031 LT854053 this study Clitocybula lacerata PRM 933893 Czech Rep. LT854021 LT854021 this study Clitocybula lacerata LE 6639 Russia HM191746 e Malysheva et al. (2011) Clitocybula lacerata LE 262744 Russia HM191747 e Malysheva et al. (2011) Clitocybula lacerata LE 262743 Russia HM191748 e Malysheva et al. (2011) Clitocybula lacerata Bizio 16837 Italy JF908760 e Osmundson et al. (2013) Clitocybula lenta BOZ (EPITYPE) Italy LT854032 LT854032 this study Clitocybula lignicola (Lignomphalia LE 262727 Russia HM191731 e Malysheva et al. (2011) lignicola) Clitocybula lignicola (Lignomphalia LE 6625 Russia HM191732 e Malysheva et al. (2011) lignicola) Clitocybula oculus LIP (P.A. Moreau CAN 13-50) Canada LT854017 LT854017 this study Clitocybula oculus PRM 935965 Canada e LT854034 this study Clitocybula oculus PRM 934963 USA LT854020 LT854020 this study Clitocybula oculus BIOUG 24046-B03 Canada KT695321 e Telfer et al. (2015) Clitocybula oculus AFTOL-ID 1554, PBM 1156 (WTU) New USA DQ192178 DQ151452 Matheny et al. (2006) Hampshire Clitocybula oculus 3512 Canada KM406971 e Berube et al. (unpubl.) Clitocybula oculus BIOUG 24046-E05 Canada KT695404 e Telfer et al. (2015) Clitocybula sp. WU 20008 Austria LT854018 LT854018 this study Clitocybula sp. Halling 4598 NY USA e JF730324 Antonín et al. (2011) Clitocybula sp. CORT 5690 USA - LT854024 this study Clitocybula sp. KA12-0435 South Korea KR673482 e Kim et al. (2015) Clitocybula sp. ASM 10506 (EIU) USA EU623638 e Hughes et al. (2007) Clitocybula sp. TENN 60306/TFB12058 USA EU623637 e Hughes et al. (2007) Clitocybula sulcata CAL 1246 (HOLOTYPE) India KR029720 KR029721 Deepna Latha et al. (2015) Clitocybula taniae BCN-SCM B-4064 Italy LT854028 LT854057 this study Gamundia leucophylla OSC 112535 USA EU669424 EU669424 Gordon et al. (unpubl.) Gerronema nemorale KG 137 South Korea EU883594 e Antonín et al. (2008) Gerronema nemorale KACC 43600 South Korea EU883593 e Antonín et al. (2008) Gerronema sp. TU 112037 Gabon JQ657793 e Tedersoo et al. (2012) Gerronema strombodes TENN 62207 USA FJ596790 e Hughes et al. (2009) Gerronema strombodes DJL05NC72 USA EU623639 e Hughes et al. (2007) Gerronema strombodes TFB 12519/TENN 60718 USA EU623640 e Hughes et al. (2007) Gerronema subclavatum Redhead 5175, DAOM not GSU66434 GSU66434 Lutzoni (1997) indicated Gerronema viridilucens e Brazil e EF514207 Desjardin et al. (2005) Gerronema wildpretii BRNM 788347 Madeira LT854043 LT854045 this study Gerronema xanthophyllum PRM 924657 Czech Rep. LT854023 LT854023 this study Gymnopus contrarius AFTOL-ID 1758, PBM 2711, CUW USA e DQ457670 Matheny et al. (2006) Hemimycena gracilis AFTOLID 1732, PBM 2715, CUW USA DQ490623 DQ457671 Matheny et al. (2006) Hydropus cf. scabripes AFTOL-ID 535, PBM2513 not DQ404389 DQ411536 Matheny et al. (2006) indicated Hydropus fuliginarius DAOM196062 not e AF261368 Moncalvo et al. (2002) indicated Hydropus marginellus AFTOL-ID 1720, PBM 2344 (WTU) Washington not DQ490627 DQ457674 Matheny et al. (2006) indicated Hydropus marginellus OSC 112834 USA EU669314 EU852808 Gordon et al. (unpubl.) Hydropus trichoderma 855 Italy JF908049 e Osmundson et al. (2013) Megacollybia clitocyboidea TENN062231 (HOLOTYPE) Japan NR_119690 e Hughes et al. (2007) Megacollybia clitocyboidea HMJAU4024/TENN62230 China EU623670 e Hughes et al. (2007) Megacollybia fallax MICH 45002 USA EU623714 e Hughes et al. (2007) Megacollybia fallax AN11591 USA EU623716 e Hughes et al. (2007) Megacollybia marginata PRM 860926 Czech Rep. LT854022 LT854022 this study Megacollybia marginata HR 91624 Czech Rep. LT854038 LT854050 this study Megacollybia marginata HR 91607 Czech Rep. LT854039 LT854051 this study V. Antonín et al. / Fungal Biology 123 (2019) 431e447 433

Table 1 (continued )

Species Specimen voucher Origin ITS (Sequence LSU (Sequence Reference ID) ID)

Megacollybia marginata HR 90203 Czech Rep. LT854047 e this study Megacollybia marginata PRM 859785 Czech Rep. LT854042 LT854046 this study Megacollybia marginata TFB7206/TENN58493 Czech Rep. EU623689 e Hughes et al. (2007) Megacollybia marginata LE 202274 Russia EU623688 e Hughes et al. (2007) Megacollybia marginata TENN 060752 (HOLOTYPE) Russia NR_119691 e Hughes et al. (2007) Megacollybia platyphylla PRM 924408 Czech Rep. LT854019 LT854019 this study Megacollybia platyphylla BRNM 737654 Czech Rep. LT854036 LT854048 this study Megacollybia platyphylla BRNM 766972 Czech Rep. LT854037 LT854049 this study Megacollybia platyphylla HR 103494 Czech Rep. e LT854040 this study Megacollybia platyphylla PRM 920937 Czech Rep. e LT854041 this study Megacollybia platyphylla BRNM 788384 Czech Rep. LT854044 e this study Megacollybia platyphylla TFB11569/TENN59541 Finland EU623691 e Hughes et al. (2007) Megacollybia platyphylla C45901 Denmark e AM946457 Saar et al. (2009) Megacollybia platyphylla GLM 45963 Germany e AY207239 Walther et al. (2005) Megacollybia platyphylla LE212071 Russia EU623699 e Hughes et al. (2007) Megacollybia rodmanii BRNM 781115 USA e LT854025 this study Megacollybia rodmanii AFTOL-ID 534, PBM2431 not e AY702016 Matheny et al. (2006) indicated Megacollybia rodmanii TENN 059430 (HOLOTYPE) USA NR_119695 e Hughes et al. (2007) Megacollybia rodmanii CIF2004-71/FCME25109 USA EU623782 e Hughes et al. (2007) Megacollybia rodmanii PRM 861168 USA LT854027 LT854056 this study Megacollybia rodmanii PRM 861169 USA e LT854059 this study Megacollybia rodmanii DAOM195782 not e AF261366 Moncalvo et al. (2002) indicated Megacollybia rodmanii AFTOL-ID 560, TENN59432 not DQ249275 AY635778 Matheny et al. (2006) indicated Megacollybia sp. BRNM 747514 South Korea LT854026 LT854055 this study Megacollybia subfurfuracea TENN059558 (HOLOTYPE) USA NR_119694 e Hughes et al. (2007) Megacollybia texensis DPL7405/TENN62058 USA EU623726 e Hughes et al. (2007) Mycena aurantiidisca AFTOL-ID 1685, PBM 1282 (WTU) Washington not DQ470811 DQ470811 e indicated Mycena olida KA131261 South Korea KR673718 e Kim et al. (2015) Myxomphalia maura DAOM187839 not e AF261378 Moncalvo et al. (2002) indicated Pleurotopsis longinqua RV95/473 not e AF042604 Moncalvo et al. (2000) indicated Porotheleum fimbriatum AFTOL-ID 1725, CBS 788.86 not DQ490626 AF261370 Matheny et al. (2006) indicated Trogia infundibuliformis KUN HKAS56709 China JQ031776 JQ031781 Yang et al. (2012) Trogia infundibuliformis KUN HKAS63661 China JQ031775 JQ031780 Yang et al. (2012) Trogia venenata KUN HKAS54710 China JQ031772 JQ031778 Yang et al. (2012) Trogia venenata KUN HKAS56679 China JQ031773 JQ031779 Yang et al. (2012) Trogia venenata TC228 China KT968080 e Mi et al. (2016) Trogia venenata MHHNU 8750 not KX268227 e Chen et l. (unpubl.) indicated Trogia venenata KUN HKAS56421 China JQ031771 JQ031777 Yang et al. (2012)

The sequences generated within this study are in bold.

and pleurotoid habitus, absent to numerous cheilocystidia, some- seems too be heterogeneous. Collections of some extra-European times present pleurocystidia, absent dermatocystidia (or with species available to us are also discussed. cystidioid pileipellis terminal cells at pileus centre), the presence of clamp connections, pileus mostly radially fibrillose to even radially 2. Materials and methods rimose, thin-to thick-fleshy context, irregular-ramose, sometimes seemingly cellular subhymenium, regular hymenophoral (some- 2.1. Morphology times subparallel to interwoven) trama, intracellular pigment, and lignicolous habitat. Except for the last characterization by Singer The original diagnoses and later published descriptions of taxa (1986), all mentioned descriptions included only species with are cited for macroscopic descriptions; remarks or descriptions amyloid basidiospores. The recent concept of Clitocybula agrees resp. on C. lacerata agg., Clitocybula familia, Megacollybia marginata, more or less with the last mentioned one. Megacollybia platyphylla and some other taxa are based on our Later, species of several other genera, e.g. Hydropus Kühner ex freshly collected material. Colour abbreviations follow Kornerup Singer, Megacollybia Kotl. & Pouzar, and Leuco-Inocybe Singer and Wanscher (1983). Microscopic features are described from (invalid name, see below) were also transferred into Clitocybula dried material mounted in 5 % KOH, Melzer's reagent, and Congo (e.g. Malençon and Bertault, 1975; Horak, 1987; Ludwig, 2001). Red, using an Olympus BX 50 and BX 43 light microscopes (Tokyo, Moncalvo et al. (2002) and Matheny et al. (2006) included all Japan) with a magnification of 1000. Authors of fungal names are these genera (except of Leuco-Inocybe) into/hydropoid clade. Aim of cited according to the Authors of Fungal Names page (http://www. this paper is the clarification of phylogeny and taxonomy of Euro- indexfungorum.org/AuthorsOfFungalNames.htm). Herbarium ab- pean representatives of this group which, in its broad concept, breviations follow Thiers (2018). Exclamation mark (!) after 434 V. Antonín et al. / Fungal Biology 123 (2019) 431e447 V. Antonín et al. / Fungal Biology 123 (2019) 431e447 435 herbarium number of type material means that the collection was and TIM2þI þ G for LSU. The resulting tree generated in MB agreed revised by the authors. Cited articles of the nomenclatural code with the ML analysis in the more supported nodes and thus only refer to its newest edition (Shenzhen Code, Turland et al., 2018). MB tree is further presented (Fig. 1). All our sequences can be For basidiospores, the factors E (quotient of length and width in assigned to the hydropoid clade sensu Matheny et al. (2006). Eight any one spore) and Q (mean of E values) are used. SEM micro- main well supported clades can be recognised within our dataset. photograph of basidiospores of Clitocybula tilieti was performed The basal one contains “Clitocybula lignicola ˮ and Mycena olida (the using the Tescan Mira 3 LMU electron microscope. taxonomic position of which will be analysed another time). Next clades are formed by species of Clitocybula including “Clitocybula ˮ 2.2. Molecular genetic analyses lenta , the type species of Leucoinocybe. Main part of Clitocybula species, incl. the type species C. lacerata are aggregated in the Total DNA was extracted from vouchers using the NucleoSpin separate clade (Clitocybula s.s. clade) which is sister (although in Plant II DNA extraction kit (MachereyeNagel) following the man- the ML analysis only) to clade of Hydropus marginellus and Hydro- ufacturer's protocol. The nuclear ITS rDNA region (ITS1-5.8S- ITS2) pus fuliginarius. The next large clade shows strongly supported in- and the large subunit (LSU) rDNA sequences were generated using ternal separation into the sub-clades formed by Trogia species the primers pairs ITS1F-ITS4B (ITS4) and NL1-NL4 correspondingly. (Trogia sub-clade), Gerronema spp., Gerronema wildpretii and Cli- Thermal cycling parameters were as in Borovicka et al. (2011). The tocybula atrialba (Gerronema sub-clade) and Megacollybia spp. PCR products were sequenced in Macrogen, Europe. The dataset (Megacollybia sub-clade). This whole clade is clustered with the ¼ was compiled using our data, sequences deposited during the minimal statistical support (posterior probability 0.77, bootstrap ¼ previous studies of Clitocybula, Megacollybia, Gerronema and support 65) with the clade formed by Clitocybe ulmicola and fi Hydropus and the most similar sequences (>91 % sequence simi- Porotheleum mbriatum. The detailed topology inside of the above fi larity in LSU rDNA) selected from GenBank using BlastN similarity de ned clades and sub-clades is described below. search tool (Table 1). The matrix was supplemented with the spe- cies of Gymnopus (Pers.) Roussel, Pleurotopsis (Henn.) Earle, Hemi- 3.2. Taxonomy mycena Singer and Baeospora Singer identified as sister lineages to the hydropoid clade (Matheny et al., 2006). The members of the Our phylogenetic studies showed, that the studied group (Cli- fi fayodioid clade (Gamundia and Myxomphalina) identi ed by tocybula s.l.) is distinctly polyphyletic (Fig. 1). Only two European Moncalvo et al. (2002) as standing outside the whole Clitocybe/ taxa (C. lacerata, C. familia) belong to the monophyletic Clitocybula. Collybia group were used as outgroup. The ITS and LSU sequences The discovered genotype variability of C. lacerata suggests an ex- originating from the single voucher or strain were concatenated (49 istence of cryptic taxa. Studies of more collections and genes, sequence pairs) and this matrix was supplemented with other ITS together with morphological characters, are necessary to under- (42 sequences) and LSU sequences (12 sequences). Thus, the se- stand their delimitation, as well as to verify if the American species quences originating from different vouchers or strains were not Clitocybula abundans really occurs in Europe. On the other hand, the concatenated. Sequence editing was done in Bioedit 7.09 (Hall, genus Megacollybia, included in Clitocybula by some authors (e.g. 1999) and sequence alignment in MAFFT 6 (Katoh et al., 2009) Ludwig, 2001; Matheny et al., 2006), represents a separate mono- using the MAFFT online server (https://mafft.cbrc.jp/alignment/ phyletic genus. This is in agreement with conclusions by Hughes fi server/). To lter both gaps and variable regions, we used Gblocks et al. (2007). The separate position of Megacollybia is supported version 0.91b (Talavera and Castresana, 2007) with less stringent also by the presence of inamyloid basidiospores (on the contrary to fi selection allowing smaller nal blocks and gap positions within the amyloid ones in Clitocybula). fi nal blocks. Bayesian searches (MB) were conducted with MrBayes According to our phylogram, “C. lenta ˮ, together with “Clito- 3.0 (Ronquist and Huelsenbeck, 2003) and 80 million replicates cybula taniae ˮ (and its synonym “Clitocybula flavoaurantia ˮ, the estimated together with burn-in value in Tracer v1.5 (Rambaut and conspecificity of which is discussed in this paper) belong to the Drummond, 2007). Maximum likelihood (ML) searches were con- separate group, originally described by Singer (1943) invalidly as ducted in IQ-TREE multicore version 1.4.1. in IQ-TREE (Nguyen et al., “Leuco-Inocybe ad interim ˮ. Therefore, the new genus Leucoinocybe “ ” 2015) including standard automated model selection ( -m TEST ) is validly described here. Its separate position is also supported by þ as well as FreeRate ( R) model, where the site rates and pro- the presence of different cystidia types than in Clitocybula. portions are inferred independently from the data. Branch supports Having asperulate basidiospores and lacking all types of cystidia, were assessed by the standard nonparametric bootstrap approxi- the position of C. tilieti within studied groups is rather strange. “ ” mation ( -b 1000 ). The substitution models were estimated in According to these micromorphological characters, it surely does jModeltest 0.1.1 (Posada, 2008). Two data partitions (one for each not belong to Clitocybula. However, its systematic placement re- subunit) were recognized in the rDNA concatenated dataset. mains unclear until its re-collection and sequencing. Although the sequencing of the studied specimens of “Clito- 3. Results and discussion cybula dryadicolaˮ was not successful, we are of the opinion that this species belongs to the genus Hydropus with the correct name 3.1. Molecular phylogeny Hydropus dryadicola (Kühner) E. Horak. The main reasons for this classification (see also Horak, 1987) are non-amyloid basidiospores The original concatenated dataset consisted of 104 taxa and and indistinct cheilocystidia (marginal cells). 1743 positions. The Gblocks curated dataset consisted of 1260 po- The only species of the newly described genus Lignomphalia sitions and 738 patterns (550 informative sites, 591 constant sites). (Lignomphalia lignicola, see below) forms a distinct phylogenetic The best substitution models were estimated as: HKY þ I þ G for ITS clade. It is rather similar to the omphalinoid species of the genus

Fig. 1. Phylogenetic placement of the studied fungi based on the concatenated nrITS and nrLSU sequences. The topology represents 50 % majority rule consensus tree from analysis in MrBayes. Bayesian PP 0.70 followed by Maximum likelihood BS 60 supports are indicated. Maximum likelihood tree was computed by IQ-TREE software. The Myxomphalia maura and Gamundia leucophylla were used as outgroups. Strongly supported branches (1.00/95) are doubled in width and the sequenced obtained during this study are printed in bold. 436 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

Leucoinocybe. It differs from them especially by the stipe colour and lignicolous way of life. Clitocybula (Singer) Metrod, Revue de Mycologie 17: 74, 1952. Fayodia subgen. Clitocybula Singer, Ann. Mycol. 41: 63, 1943. Original description (Singer 1943): “Sporis amyloideis, sub- globosis, levibus, vel leniter subasperulatis, membrana sub- crassiuscula instructis, sed perisporio visibili destitutis. Hymenio cystidiis destituto, cheilocystidiis rarissimis vel nullis. Habitu ad Clitocybes minores accedit. ˮ. Generic description by Metrod (1952): “Port clitocyboïde-colly- bioïde; pied furfurace finement; lamelles larges, espacees, veinees, adnees puis decurrentes, blanches a grisatres; spores rondes, amyloïdes; pas de cystides. Sur le bois. Une espece. ˮ. Type species: C. lacerata (Scop.) Metrod (see Donk, 1962). Clitocybula s.s. clade is further separated in five sub-clades (Fig. 1). Two clades with uncertain taxonomic identity are repre- sented by the Clitocybula sp. ‒ collections, the first one by Halling Fig. 2. Clitocybula lacerata (PRM 951559, neotype). Basidiomata in situ. Slovenia, Upper 4598 (Antonín et al., 2011) and CORT 5690, the second one by TENN 60306 and GRSM 77072 (C. aff. lacerata)(Hughes et al., 2007). Other Carniola region, near the town of Skofja Loka, close to the village of Vincarje, 20 Sep. 2018, leg. B. Arzensek & A. Piltaver. Photo A. Piltaver. two well defined sub-clades contains collections of Clitocybula oculus and C. familia respectively. The remaining sub-clade contains specimens identified as C. abundans and C. lacerata and can be Smell faint but distinctively aromatic, resembling the odour of fresh further subdivided to three distinct lineages. spruce or fir resin in liquid state. Taste mild but distinctively Clitocybula lacerata (Scop.) Metrod, Rev. Mycol. 17: 74, 1952. adstringent. (Figs. 2e5). Microscopic description (neotype, Fig. 3‒5): Basidiospores e e m ¼ e ¼ Basionym: Agaricus laceratus Scop., Fl. Carniolica, Ed. 2, Vol. 2: 5.0 6.0 4.0 5.0 m, E 1.10 1.33, Q 1.17, subglobose to 439, 1772. broadly obovoid, with small but distinct hilar appendix, smooth, e e m Collybia lacerata (Scop.) Gillet, Hymenomyc etes: 310, 1876 (this distinctly amyloid. Basidia 4(1)-spored, 28 32 7 8 m, clavate. e e m e e m combination is based on Gillet's reference to Fries who refers to Basidiolae 25 27 5 6 m. Cheilocystidia rare, 30 32 8 9 m, e Scopoli); Collybia platyphylla subsp. lacerata (Scop.) Konrad & clavate, with 1 2 slight constrictions. Gloeoplerous hyphae pre- e e m Maubl., Icones Selectae Fungorum, Vol. 3, Fasc. 7: pl. 204, 1932; sent, some of them protruding from lamellae edge, 10 25( 30) m Fayodia lacerata (Scop.) Singer, Annales Mycologicae 43: 63, 1944. e broad, cylindrical or attenuated towards the apex which is obtuse. Combinations into the genera Hydropus and Baeospora cited in In- Pleurocystidia not observed. Lamellar trama regular, of cylindrical e m dex Fungorum are invalid as basionyms are not indicated and ref- hyphae 4 24 m broad, cells long, hyaline. Pileipellis a cutis of e m erences omitted. Clitocybula lacerata var. odorata Bon, Bull. densely arranged parallel hyphae 3 13 m broad, of long cylin- Trimestriel Fed. Mycol. Dauphine-Savoie 31 (121), 17, 1991, is not drical cells, with pale encrusting pigmentation. Pileocystidia scat- e e m included in this study. tered, mostly in nests, 15 20( 25) m broad, clavate to almost Holotype: Not existing. No herbarium material is known to exist sphaeropedunculate, some of them slightly constricted in the broad of Scopoli's collections although it is evident from several sources part, with slightly thickened wall, hyaline. Stipitipellis of the same that these collections did exist. Probably they were burned by two characters like pileipellis. Caulocystidia frequent, isolated or in consequent fires in his house in Idria (Voss, 1885: 16). Holotype nests, mostly curved or perpendicularly upturned, clavate but e e m area: Carniola, historical region that comprised parts of present- irregular in shape, with upper part 15 20( 27) m broad, with day Slovenia (roughly its western half). slightly thickened wall, hyaline. Clamp connections present in all Neotype: Slovenia, Upper Carniola region (Gorenjska region) in tissues. NW Slovenia, near the town of Skofja Loka, close to the village of Ecology and Distribution: Growing in clusters on dead wood of Vincarje: at the end of a narrow, small, humid valley, at the right conifers, mostly P. abies and Abies alba (our data), but also Pinus side of a small creek, 46.1601269N, 14.2916650E, alt. ± 410 m, deep sylvestris (Krieglsteiner, 2001), preferably in natural forests. Ac- highly grown forest (predominantly Picea, Fagus, Abies, Acer), Picea cording to literature, this species is distributed in Europe (see abies: on rotten trunk and stump, 20 Sep. 2018, B. Arzensek & A. above) and North America (e.g. Bigelow, 1973), especially at higher Piltaver (PRM 951559, neotypus hic designatus; MycoBank MBT altitudes. € € 386133). Specimens examined: Austria: Oberosterreich:Vocklabruch, 10 & Selected descriptions and iconography: Breitenbach and Kranzlin€ Sep. 1999, K. Helm (WU, 20008); Ibid., 10 Sep. 1999, K. Helm H. (1991); Ludwig (2000, 2001); Krieglsteiner (2001); Roux (2006); Dietsch (WU, 19575). Czech Republic: Bohemian Forest, Boubínský Eyssartier and Roux (2011). prales National Nature Reserve, alt. 970 m, 4 Sep. 2013, J. Holec JH “ ” Macroscopic description (neotype, Fig. 2): Growing in big groups 165/2013 (PRM 922903, as C. abundans ); Ibid., 13 Sep. 2013, close together or in smal clusters of 2e3 specimens. Pileus M. Kríz (PRM 933893); Bohemian Forest, Zatonska hora Nature & 15e40 mm, thin-fleshed, translucent, dry, at first hemisphaerical, Reserve, alt. 940 m, 5 Aug. 2014, J. Holec M. Kríz JH 63/2014 (PRM then convex with depressed centre, surface innately radially 924505). Novohradske hory Mts., Zofínský prales National Nature fibrillose and radially cracked at margin when old, beige grey to Reserve, alt. 800 m, 28 Sep. 2008, J. Holec JH 173/2008 (PRM pale grey brown (Methuen 6B2, 7B2 or paler). Gills medium spaced, 915404). Slovakia: Kremnicke vrchy Mts., Badínsky prales Nature adnexed to adnate, white. Stem 30e50 (e80) 3e7 mm, hollow at Reserve, alt. 720 m, 28 Sep. 2009, J. Holec JH 139/2009 (PRM 899161, “ ” maturity, cylindrical, slightly bulbous at the base, which is finely as C. abundans ). Veporske vrchy Mts., Dobrocský prales National whitish fibrillose, at first white, becoming beige grey or grey-brown Nature Reserve, alt. 970 m, 29 Sep. 2011, J. Holec JH 143/2011 (PRM downwards. Decayed specimens are turning deeper grey brown. 899420); Ibid., 29 Sep. 2009, J. Holec JH 148/2009 (PRM 899168, as V. Antonín et al. / Fungal Biology 123 (2019) 431e447 437

Fig. 3. Clitocybula lacerata (PRM 951559, neotype). Microcharacters (AeE: stained with Congo Red, F: Melzer's reagent). (A) lamella edge with protruding apices of gloeoplerous hyphae (arrows); (B) lamella edge with protruding gloeoplerous hyphae (arrows), squashed mount; (C) lamella edge with protruding gloeoplerous hyphae and hymenium cells, basidium marked with arrow, squashed mount; (D) lamella edge with cheilocystidium; (E) lamella edge with two chelocystidia (arrows) and gloeoplerous hyphae in squashed lamellar trama; (F) basidiospores. (AeE): mounted in 5 % KOH with Congo Red; (F): mounted in Melzer's reagent. Bars ¼ 20 mm for AeB, 10 mm for CeF. Photo J. Holec.

“C. abundans”). Slovenia: Upper Carniola region, near the town of almost all collections except for PRM 899420 and WU 20008. They Skofja Loka, close to the village of Vincarje, 20 Sep. 2018, B. Arzensek are either scattered or very rare, hyaline, variable in size and shape & A. Piltaver (PRM 951559, neotype; for details see neotype (Figs. 3D, F and 4D, F), cylindrical to clavate (rarely almost sphaer- designation). opedunculate), ofter flexuous, measuring 20e50 8e20 mm, some Notes: According to literature (Bigelow, 1973; Moser, 1983; of them slightly thick-walled on apex. They are often oriented Breitenbach and Kranzlin,€ 1991; Holec, 2012), C. lacerata is char- parallel to the lamellae edge. acteristic by its growth in clusters, depressed mature pilei with Moreover, we observed another “cystidiod” elements. They radially cracked margin, distinctly radially fibrillose pileus surface protrude several mm to almost 100 mm above the lamellae edge, are and spores slightly larger [4.5e7.0 (‒8.0) 4e6 mm] in comparison cylindrical, often flexuose, with obtuse apex, sometimes slightly with C. abundans and C. familia. In the cited literature, the almost thick-walled, with 6e10 mm broad upper part and sometimes total absence of cheilocystidia in C. lacerata is used as the key dis- swollen basal part (in hymenium level) reaching the width of up to tinguishing character against the cheilocystidia-possessing Amer- 20 mm(Fig. 3AeC and 4B, E). They represent terminal cells or apical ican species C. abundans. As shown below, the real situation is more parts of 15e24 mm wide hyphae of the lamellar trama which are complicated. distinct also by their “lipid-like ˮ content, either homogeneous or in Our study confirmed that European material of C. lacerata is very the form of droplets, and almost total absence of septa. We inter- variable morphologically. Depending on weather conditions and pret them as gloeoplerous hyphae. In some mounts they were ar- age, its pileus and stipe is rather dark (grey-brown, especially when ranged in loose bundles hidden in the lamelar trama and ending in young and fresh) or pale (whitish with darker centre when mature the hymenium layer or above it (Fig. 4A). Such gloeoplerous hyphae and dry). Its habit changes considerably during development, from were observed scarcely to frequently in all studied collections from rather thick young basidiomata with convex pileus to slim old ones the Czech Republic and Slovakia. We are convinced “cheilocystidia” having depressed pileus with radially cracked margin and long that up to 200 mm long reported and depicted e.g. by Breitenbach stipe. Microscopically, the material studied by us is also variable. and Kranzlin€ (1991: no. 181) in C. abundans are in fact apical parts of The most important finding is that cheilocystidia are present in gloeoplerous hyphae, not the true cheilocystidia (which, as shown 438 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

above, are smaller, hyaline and mostly clavate). Holec (2012) used the same misinterpretation and identified collections with such “cheilocystidia ˮ as C. abundans. In fact, all collections from the Czech Republic and Slovakia preliminary identified as C. lacerata or C. abundans possess more or less protruding but usually very distinct gloeoplerous hyphae. Additionally, they have less distinct, smaller, hyaline, mostly clavate cheilocystidia, which are rare to scattered. It means that the distinguishing of C. lacerata and C. abundans based on absence/presence of cheilocystidia is inap- plicable. As shown in discussion on C. abundans, there is not even certainty that this species really occurs in Europe. Spores are also variable in our material. Their size range is 4.5e5.5 4e5 mm in some collections (e.g. PRM 922903) but 5.5e7.2 5e6 mm on others (e.g. PRM 915404). In these “utter- most” collections the values do not overlap. In all studied collections the stipe surface is covered by nests of cylindrical-clavate to clavate caulocystidia measuring 22e65 8e30 mm. They are macroscopically apparent as pruinose to floccose stipe covering. Pileus cuticle contains numerous pileo- cystidia resembling the caulocystidia. The large morphological variability of studied material is demonstrated molecularly, too. The European and Asian collections preliminary identified as C. lacerata or C. abundans are positioned in one lineage, which is, however, separated into three well- supported branches (Fig. 1). There is also a sister groups consist- ing of two unidentified North American collections (TENN 60306, GRSM 77072) not studied by us. The variability inside the branches is very low whereas the individual branches differ considerably (with ITS similarity only 94e95.6 %). The most distant group in terms of genetic distance is formed by two Russian collections (Siberia, Altaj Republic: LE 262744; Primorsky Territory: LE 6639; Malysheva et al., 2011) and one from Austria (WU, 19575), all of Fig. 4. Clitocybula lacerata agg. (PRM 915404: AeD; PRM 922903 preliminary identified as “C. abundans”:EeF). (A). Schematic view of undulate lamella edge showing arrange- them not seen by us in fresh state. Our collections from the Czech ment of gloeoplerous hyphae (not protruding in this case) in lamellar trama; (B) detail of Republic, Slovakia and Slovenia are distributed in two clades con- lamella edge with basidiolae and protruding gloeoplerous hypha; (C) basidium; (D) taining also material from Italy (Bizio 16837), Austria (WU, 20008) cheilocystidium; (E) detail of lamella edge with basidiolae, basidia and long protruding and Russia (Siberia, Altaj Republic: LE 262743). The Slovenian gloeoplerous hypha; (F) cheilocystidia. Scale bars ¼ 10 mm. Del. J. Holec. collection, originating from the type area of C. lacerata, has identical ITS and LSU sequences like the Czech (PRM 922903), Austrian (WU, 20008) and Siberian (LE 262743) ones. Unfortunately, we were not able to recognize any pattern of correlation of morphological characters or geographical distribution compatible with the three groups (plus the American one) discovered molecularly. However, the discovered genotype variability shows an existence of cryptic taxa. Study of more collections and genes is necessary to under- stand their delimitation. To do at least the first step to fix the taxonomic and nomenclatural situation within the C. lacerata group, we designated the collection from Slovenia as neotype (in situation when no original materials exist, see typification obove). Generally, we consider the European C. lacerata a species com- plex which should be treated as C. lacerata agg. at present. At the current state of knowledge (when no collection of the American C. abundans is sequenced) it is unclear if this complex contains the true C. abundans (for details see discussion on C. abundans). Clitocybula abundans (Peck) Singer, Sydowia 15: 53, 1962 [1961]. (Fig. 6). Basionym: Agaricus (Collybia) abundans Peck, Annual Rep. New York State Mus. Nat. Hist. 29: 38, 1878 [1876]. Collybia abundans (Peck) Sacc., Syll. Fung. 5: 241, 1887; Fayodia abundans (Peck) Singer, Lloydia 5: 126, 1942; Clitocybula abundans (Peck) Singer, Sydowia 8: 110, 1954, invalid combination (basionym not indicated). Holotype: Not designated; two collections (syntypes) from decaying trunks in woods are briefly mentioned by Peck in the Fig. 5. Clitocybula lacerata (PRM 951559, neotype). SEM microphotograph of basidio- protologue: USA, New York State, Sandlake, Aug. (NYS); Ibid., Greig, spores. Photo L. Ilkovics. V. Antonín et al. / Fungal Biology 123 (2019) 431e447 439

bad state of preservation (shrinked hymenial cells). Cheilocystidia can also have a cylindrical-fusiform shape. Ecology and Distribution: Growing caespitose on dead wood of mostly conifers, but also broadleaved trees, proved with certainty only from the USA (see Remarks). Specimens examined e collections by C.H. Peck: USA: New York State: Greig, Sep., C.H. Peck (NYS!, mycology type database no. 27, syntype); New York State: Caroga, Jul., C.H. Peck (NYS); New York State: Berne, Aug., C.H. Peck (NYS); New York State: Piseco, Sept., C.H. Peck, NYS. All specimens have no day and year indication. We do not have data enough to judge if the non-type collections represent original material used for the description of Agaricus abundans Peck or later collected specimens. Other studied specimens: USA: Idaho: Cascade Lake, on well decayed wood, 18 Jun. 1983, during USA coll. foray, det. O.K. Miller OKM 20432 (NY 01933720). Montana: Yellow Bay, on a mossy conifer log, 11 Jul. 1983, L. Bailey, det. O.K. Miller OKM 20510 (NY 01933721); Montana: Noisy Creek, on strand fir log, 22 Jun. 1981, Fig. 6. Clitocybula abundans (NYS, mycology type database no. 27, syntype). (A) chei- O.K. & H.H. Miller OKM 19309 (NY 01933722). New York: Bergen locystidia; (B) basidiospores. Scale bars ¼ 10 mm. Del. J. Holec. Swamp, hemlock? stump, 5 Oct. 1971, H.S. Vishniac (NY V,127;57-7). Maine: Penobscot Co., Orono, on buried wood, 4 Aug. 1979, R.E. Sep. (NYS!). A lectotype can be selected from them after clearing up Halling (NY Halling no. 2924). the species delimitation. Notes:AsC. abundans was described from North America, we Original description by Peck (1878): Pileus thin, convex or primarily revised the American material to understand its charac- fi expanded, subumbilicate, innate- brillose, whitish inclining to ters and variability. The studied Peck's collections are very similar fi fuscous, often a little darker and more densely brillose on the disk, morphologically and the same fact relates to the more modern the thin margin easily splitting; lamellae narrow, close, adnate, material from Yellow Bay, Noisy Creek, and Bergen Swamp. The sometimes veiny, white; stem equal, smooth, hollow, easily split- collections from Cascade Lake and Orono are somewhat different as ting, often curved, colored like the pileus, pruinose at the top. Plant 0 0 0 0 no cheilocystidia were observed and their spores are slightly larger, gregarious or subcaespitose, 1 -2 high, pileus 1 -1.5 broad, stem e ‒ e ‒ m 00 measuring 5.6 7.2 (4.0 )4.4 5.6( 6.4) m, with prevailing 1 thick. This fungus is not frequent, but when it does occur it is broadly ellipsoid to ellipsoid shape. It could indicate their deviating usually in great abundance. When drying the margin rolls inward or even non-conspecific character. In other collections studied, the and the color becomes darker. spores measure 4.8e5.6(‒6.4) 4.0e4.8(‒5.6) mm and are mostly Selected macroscopic descriptions: Peck (1878), Bigelow (1973). subglobose to broadly ellipsoid. Macromorphologically, the studied Microscopic description: (part of the Greig syntype: 2 pileus American material is represented by rather small and pale basi- fragments, 5 5 mm each, 15 mm of stipe; material hard to study, diomata (pileus up to 40 mm broad, whitish with darker centre). “ fi ” “ ” overdried, most cells scleri ed and shrinked with indistinct Unfortunately, our attempts to isolate DNA from the modern col- details, mostly not possible to divide them and observe individual lections studied by us failed (we did not try it from Peck's collec- e ‒ e ‒ m cells). Basidiospores 4.8 5.6( 6.4) 4.0 4.8( 5.6) m (in Melzer), tions), probably due to their age and massive use of fumigation subglobose to broadly ellipsoid, rarely globose, with small but chemicals for their preventive conservation. Moreover, no se- distinct hilar appendix, smooth, distinctly amyloid (Fig. 6B). Basidia quences of C. abundans are available in GenBank. All these facts m 4-spored, about 20 5 m, narrowly clavate. Cheilocystidia scat- disable to fix its phylogenetic position and compare it with other tered, not much protruding above the lamellae edge, some of them sequenced Clitocybula species. parallel to the edge and only slightly upturned, As far as we know, C. abundans was introduced to European e e m 26 44 5.5 8.0 m, narrowly clavate, thin-walled, rarely there literature by Moser (1983), based probably on the description and fl are bundles of exuose hyphae protruding high above the edge but key by Bigelow (1973). Bigelow used the pubescent stipe and not differentiated to the form of regularly shaped cystidia (Fig. 6A). presence of cheilocystidia in C. abundans as the distinguishing Pleurocystidia not observed. Lamellar trama regular, of cylindrical characters against C. lacerata which according to him should have e m hyphae 3 12 m broad, with yellow membranal pigment and en- glabrous stipe and no cheilocystidia. Moser emphasized also the crustations, yellow in Melzer's reagent, not dextrinoid. Pileus whitish, at centre often darker to grey-brown, smaller pileus (up to cuticle observed on scalp surface (sections not made to save the 40 mm) of C. abundans and slightly smaller spores type material), a cutis of parallel or slightly interwoven cylindrical (4.4e6.5 3.5e5.5 mm). Nice pictures of a supposed C. abundans e m hyphae 3 10 m broad. Pileocystidia rare, narrowly clavate to were published by Breitenbach and Kranzlin€ (1991: no. 181) and clavate, resembling cheilocystidia in shape but larger, about Moser (Farbatlas: Clitocybula 1, lower picture). m 65 15 m, thin-walled. Stipe cuticle (scalp) a cutis of parallel The cited data, pictures and some “cheilocystidia-having” col- e m cylindrical hyphae 2.5 5.0 m broad, covered by encrustations, at lections from Central Europe inspired Holec (2012) to include the e m places with nests of upturned to interwoven hyphae 2.5 5.0 m species to a key in Funga Nordica. However, during our study the broad. Caulocystidia scattered, lageniform to clavate, irregular in putative cheilocystidia proved to be a matter of misinterpretation fl e e m shape, often exuose, about 30 35 7 10 m. Clamp connections and inconspicuous but clear cheilocystidia were found in present. C. lacerata, too (for details see remarks on that species). There is also The non-type collections by Peck (Caroga, Berne, Piseco; see a discrepancy between the pale American C. abundans and its dark- below) are in the same (i.e. not optimal) state of preservation like coloured European interpretation (Breitenbach and Kranzlin,€ 1991: the syntype. They agree with the Greig syntype in almost all no. 181; Moser, Farbatlas: Clitocybula 1, lower picture; Holec, 2012). characters. Some of their spores are broadly obovoid. In Berne Generally, for all these reasons it is not clear if the European collection, no cheilocystidia were observed but probably due to the collections identified as C. abundans are really conspecific with the 440 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

American ones. Moreover, we cannot compare them molecularly as substrate. It is known from Austria, Czech Republic, France, there are no available sequences of the true American C. abundans. Slovakia, Sweden and Switzerland in Europe, and from North At moment, we have to consider the presence of C. abundans in America (Antonín et al., 2011). Europe insufficiently proved. Specimens examined: Czech Republic: Novohradske hory Mts., Clitocybula familia (Peck) Singer, Sydowia 8 (1e6): 110, 1954. Zofínský prales Nature Reserve, alt. c. 750e800 m, 30 Sep. 2008, Basionym: Agaricus familia Peck, Annual Rep. New York State V. Antonín & V. Kabat VA 08.251 (BRNM 736054); Ibid., J. Borovicka 0 00 0 00 Mus. Nat. Hist. 23: 79, 1873. (PRM 921866); Ibid., alt. 780 m, 48º40 03 N, 14º42 36 E, 5 Oct. 2009, J. Novotný (CB 16204). Vsetínske vrchy Mts., SE slopes of Cab Collybia familia (Peck) Sacc., Syll. Fung. 5: 241, 1887; Gymnopus e familia (Peck) Murrill, N. Amer. Fl. 9 (5): 365, 1916; Baeospora familia hill, Kutaný Nature Reserve, alt. c. 650 750 m, 11 Oct. 2008, D. Dvorak 645/08 (BRNU 618420). Canada: Ontario: Long Point, L. (Peck) Singer, Rev. Mycol. 3: 193, 1938. Collybia familia var. compressa Romagn., Collect. Bot. 7 (2), 58: Temagami, 7 Sep. 1936, R.F. Cain (NY). Quebec: St. Adolphe, 2 Sep. 1959, R.F. Cain (NY). Manitoba: Saint-Thuribe, Saint Leon, 13. Sep. 1090, 1968; Clitocybula familia var. compressa (Romagn.) H.E. Bige- low, Mycologia 65 (5): 1102, 1973. 2015, R. Lebeuf HRL 2153 (PRM 935966) France: Nay, B. Pyrenees, 2e4 Nov. 1965, J. Beller (PC!, holotype of Collybia familia var. com- Type specimens: USA, Adirondack Mts., North Elba, Aug., Peck (NYS, not studied). France, Nay, B.-Pyrenees, 2e4 Nov. 1965, J. Beller pressa). Slovak Republic: Kremnicke vrchy Mts., Badín, Badínsky prales Nature Reserve, alt. c. 750 m, 28 Sep. 2009, V. Antonín & J. (PC!, holotype of Collybia familia var. compressa). º 0 º 0 Selected macroscopic descriptions and iconography: Romagnesi Lederer VA 09.296 (BRNM 736053); Ibid., 48 41.321 N, 19 03.293 E, J. Holec JH 142/2009 (PRM 899163). Veporske vrchy Mts., Dobroc, (1968); Lennox (1979); Ludwig (2000, 2001); Groger€ (2006); e Antonín et al. (2011). Dobrocský prales Nature Reserve, alt. c. 800 900 m, 29 Sep. 2009, V. Antonín VA 09.310 (BRNM 736052); Ibid., 48º40.8270 N, Microscopic description: Basidiospores 3.5e5.3 (‒ º 0 5.5) 3.5e5.0 mm, average 4.4 4.0 mm, E ¼ 1.00e1.43, 19 40.504 E, alt. 920 m, 28 Sep. 2009, J. Holec JH 154/2009 (PRM Q ¼ 1.09e1.21, globose, subglobose to broadly ellipsoid, thin- 899174). USA: Idaho: Idaho Co., Papoose Creek, 7 Devil Mts., 3 Sep. walled, amyloid, smooth. Basidia 18e28 5.0e7.5 mm, 4-spored, 1954, H.E. Bigelow 1940 (NY). Maine: Franklin Co., Flagstaff Lake narrowly clavate. Basidioles up to 28 3e7 mm, narrowly clavate to Road, Carrabassett, 24 Sept. 1971, H.E. Bigelow 16757 (NY). Michi- cylindrical. Lamellae edge fertile. Marginal cells sometimes scat- gan: Luce Co., Tahquamenon, 12 Sep. 1969, W. Habermehl, Bigelow tered, sometimes ± frequent, (16‒)21e34 (‒40) 3e7 mm, cylin- 16061 (NY). drical, narrowly clavate, regular, irregular, some with irregular, Notes: Clitocybula familia is a distinct, molecularly well- sometimes branched narrow rostrum, thin-walled. Trama hyphae supported species, differing from other European Clitocybula spe- cylindrical to fusoid, thin-to slightly thick-walled, smooth, non- cies by its growth in larger and denser clusters, having a more or dextrinoid, clamped, up to 25 (‒35) mm wide. Pileipellis a cutis of less mycenoid, usually not centrally depressed (or only sometimes fi radially arranged, ± cylindrical, 3e10 mm wide hyphae, with when old), smooth or sometimes very slightly innately brillose fi adpressed to suberect, cylindrical or fusoid, less frequently sub- pileus (never distinctly radially brillose), and by small basidio- cylindrical, obtuse, slightly thick-walled, smooth or minutely spores. According to Lennox (1979) the basidiomata of the North- incrusted, subhyaline or pale greyish, up to 55 mm wide terminal American collections have a yellowish tan to honey coloured e cells. Stipitipellis a cutis of cylindrical, smooth, non-dextrinoid, ± pileus and closer lamellae (48 56 reaching the stipe), however, slightly thick-walled, up to 6.0 mm wide cells. Caulocystidia specimens studied by us are microscopically identical with Euro- pean ones. For detailed comments, see Antonín et al. (2011). numerous, (22‒)32e55 4e12(‒16) mm, cylindrical, fusoid, clavate, regular, (sub)rostrate or slightly irregular, thin-walled. Clitocybula tilieti (Singer) Singer, Sydowia 15 (1961): 53, 1962 Ecology and Distribution: In Central Europe, C. familia was mostly (Figs. 7 and 8). collected on decaying fallen trunks of Abies alba, rarely of P. abies. Basionym: Fayodia tilieti Singer, Ann. Mycol. 41: 63, 1943. Most American collections lack detailed information on substrates; Holotype: Russia, Mordovsky zapovednik [Mordovsky Nature in one case (Quebec, Cain s.n.), a decayed Acer is mentioned as the Reserve], 1937, Kuznetsov (LE 17627!).

Fig. 7. Clitocybula tilieti (LE 17627, holotype). (A) basidiospores; (B) marginal cells; (C) pileipellis; (D) stipitipellis. Scale bar ¼ 20 mm. Del. V. Antonín. V. Antonín et al. / Fungal Biology 123 (2019) 431e447 441

Republic of Mordovia, on the forested right bank of the Moksha River, a tributary of the Oka River. The sequencing of the holotype was unsuccessful. Because of having asperulate basidiospores, a unique character within all studied genera, and lacking all types of cystidia, it surely does not belong to Clitocybula, and the right generic position of this taxon is unclear. The final decision on its generic placement would be possible only after its re-collection and successful sequencing. Leucoinocybe Singer ex Antonín, Borovicka, Holec & Kolarík, gen. nov. MycoBank MB 825128. Leuco-Inocybe Singer, Ann. Mycol. 41: 144, 1943 (invalid name: described provisionally). Description: Basidiospores moderately large, smooth, amyloid; cheilocystidia present, but sometimes scattered; pleurocystidia absent or present but scattered, similar to cheilocystidia; pileipellis an (ixo)cutis with the presence or absence of pileocystidia, if pileocystidia absent, then pileocystidioid terminal cells present; caulocystidia present, distinct; clamp connections present in all studied tissues. Type species: Mycena lenta Maire, Bull. Soc. Mycol. France 44: 40, 1928 (Leucoinocybe lenta (Maire) Antonín, Borovicka, Holec & Kolarík; for a valid combination see below). Etymology: Macroscopically similar to species of the genus Ino- cybe but having white lamellae. Fig. 8. Clitocybula tilieti (LE 17627, holotype). SEM microphotograph of one basidio- Notes: The original description by Singer (1943) was invalid spore. Photo L. Ilkovics. because the genus was described provisionally (ad interim). Singer delimited the group as follows: “Leuco-Inocybe gen. nov. ad int. Pileo epicute haud diverticulata, hypodermate haud celluloso, sed Macroscopic description: Singer (1943). pilis liberis instructo. Lamellis subliberis, latiusculis. Sporis ellip- ‒ Microscopic description (holotype revision): Basidiospores 4.7 soidalibus vel amygdaliformibus, amyloideis, mediocriter volumi- e ‒ m m ¼ e 6.2 3.7 4.5( 5.0) m, average 5.4 4.3 m, E 1.17 1.38, nosis. Trama lamellarum regulari, haud amyloidea. Habitus ¼ Q 1.26, (broadly) ellipsoid to subglobose, sometimes subovoid or marasmioidearum vel inocybis. Typus gen. L. lenta (Mre.) (Singer, subamygdaloid, thin-walled, asperulate, amyloid. Basidia 1943)ˮ. The invalidity was mentioned by Donk (1962) and Horak e e m 24 33 6.5 7.5 m, 4-spored, clavate, clampless. Basidioles (1968), too. The genus is validly described here. 11e35 3.5e7.5 mm, clavate, subcylindrical or subfusoid, clamp- less. Cheilocystidia absent. Marginal cells 14e17 4.5e6.5 mm, clavate, subfusoid, thin-walled, clampless. Pleurocystidia absent. Trama hyphae cylindrical, thin-to slightly thick-walled, non-dex- trinoid, clampless, 3e8 mm wide. Pileipellis a cutis composed of ± cylindrical, ± thin-walled, smooth, non-dextrinoid, clampless, 3.5e7.0 mm wide hyphae; subpileipellis hyphae incrusted, grey- black in KOH; terminal cells adpressed to erect, cylindrical or narrowly clavate, obtuse, thin-walled, up to 7 mm wide. Stipitipellis a cutis made up of cylindrical, parallel, ± slightly thick-walled, non- dextrinoid, clampless, 3e6 mm wide hyphae; terminal cells adpressed to erect, cylindrical, narrowly fusoid, narrowly clavate, obtuse, subcapitate or slightly attenuated, slightly thick-walled.o.€ Ecology and Distribution:OnTilia twigs in fallen leaves and detritus layer in a Tilia stand; in August and September. It is known only from the type locality in Russia till now. Specimen examined. Russia: Mordovsky zapovednik [Mordovsky Nature Reserve], 1937, Kuznetsov, det. R. Singer (LE 17627!; holotype). Notes: Clitocybula tilieti is characterized by a up to 40 mm broad (when dry), grey-brown to black-brown pileus, decurrent, narrow and white lamellae, a 40 5e7 mm, smooth, glabrous, stipe con- colourous with pileus, 4.7e6.2 3.7e4.5(‒5.0) mm, asperulate, amyloid basidospores, the absence of cheilocystidia, pleurocystidia and true caulocystidia, and clampless hyphae. The location of the type specimen was considered unclear because of the misinter- pretation of the type locality name: Mordavia (Singer, 1943), Moldavia (Mueller and Wu, 1997). However, it has been kept in the LE herbarium where Singer worked in those years. Mordovsky Fig. 9. Leucoinocybe lenta (BOZ, epitype): (A) basidiospores; (B) cheilocystidia; (C) Nature Reserve is located in European Russia on the territory of the pileocystidia; (D) caulocystidia. Scale bar ¼ 20 mm. Del. V. Antonín. 442 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

Leucoinocybe lenta (Maire) Antonín, Borovicka, Holec & Kolarík, Holotype: Spain: Girona: S'Arenella, el Port de la Selva, 18 Nov. comb. nov. (Fig. 9). 1997, L. Vila & X. Llimona (BCN-SCM B-4064!). MycoBank MB 825130. Macroscopic descriptions and iconography: Vila (2002,as Basionym: Mycena lenta Maire, Bull. Soc. Mycol. France 44: 40, C. taniae); Contu (2003,asP. flavoaurantia); Malysheva et al. (2011, 1928. as C. flavoaurantia). Collybia lenta (Maire) Maire, Fungi Catalaunici: 68, 1933; Clito- Microscopic description (holotype revision): Basidiospores cybula lenta (Maire) Malençon & Bertault, Fl. Champ. Super. Maroc 7e9 5.5e7.0 mm, average 8.1 6.2 mm, E ¼ 1.14e1.55 (‒1.64), 2: 395, 1975; Leuco-Inocybe lenta (Maire) Singer, Ann. Mycol. 41: Q ¼ 1.32, (broadly) ellipsoid to subglobose, thin-walled, smooth, 144, 1943 (invalid, The Code Art. 35.1). amyloid. Basidia 30e37 9.0e11 mm, 4-spored, clavate, clamped. Lectotype: Algeria:Foret^ de la Reghaïa, a terre (sur brindilles Basidioles up to 35 6e11 mm, clavate or subcylindrical, clam- enterrees), 18 Nov. 1923, R. Maire 8285 (MPU!, lectotypus hic des- ped. Cheilocystidia distinct, (28‒)44e64 7e12 mm, fusoid, ignatus; MycoBank MBT 385412). The specimen is not well- (sub)lageniform, mostly (sub)acutely rostrate, thin-walled, preserved (similarly like other collections by Maire in MPU, but clamped. Pleurocystidia absent. Trama hyphae cylindrical to this one is the best one of the original material). To support it by a subinflated, thin-walled, non-dextrinoid, clamped, 5e25 mm better material, we decided to designate an epitype, a more recent wide. Pileipellis a cutis composed of ± cylindrical, thin-walled, collection from the Mediterranean area, microscopied and non-dextrinoid, clamped, 3.5e8.0 (‒10) mmwidehyphae;ter- sequenced by us. minal cells adpressed to suberect, cylindrical, fusoid, clavate, Epitype: Italy: Sardinia: Cala Gonone, on wood of Quercus ilex, 31 obtuse, thin-walled, up to 10 mm wide. Stipitipellis a cutis made Oct. 1985, C. Rossi (BOZ!, epitypus hic designatus; MycoBank MBT up of cylindrical, parallel, ± thin-walled, non-dextrinoid, clam- 381559; GenBank: LT854032 and LT854060). ped, 3e7 mm wide hyphae. Caulocystidia mostly in groups, Macroscopic descriptions and iconography: Maire (1928); 19e51 5e11 mm, fusoid, (sub)lageniform, clavate, often (sub) Malençon and Bertault (1975); Ludwig (2000, 2001); Groger€ acutely rostrate, thin-walled. (2006); Eyssartier and Roux (2011). Ecology and Distribution: In fascicles, apparently terricolous, but Microscopic description: Basidiospores (5.5‒)6.0e7.7 (‒ growing on underground roots probably of living grasses. It is 8.0) 4.0e5.2 (‒5.5) mm, average 7.0 4.7 mm, E ¼ 1.33e1.67, known from several localities in the European Mediterranean Q ¼ 1.47e1.48, (broadly) ellipsoid, sometimes with a slight supra- region. hilar depression, thin-walled, smooth, amyloid. Basidia Specimens examined: Italy: Sardinia: Diga del Liscia, Lato Luras, 23e35 8.5e9.5 mm, 4-, rarely 2-spored, clavate, sometimes sub- Podda Carana (SS), 20 Sep. 2002, M. Contu (GDOR!, holotype of capitate, clamped. Basidioles 12e38 3e10 mm, clavate, sometimes P. flavoaurantia); Sardinia: Stazzo Montesu, Tempio Pausania, 13 subcapitate, subcylindrical, subfusoid, clamped. Cheilocystidia Sep. 2002, 17 Sep. 2002, 7 Oct. 2002 & 15 Oct. 2002, M. Contu scattered, 31e60 9e19(‒22) mm, clavate, fusoid, sometimes (GDOR, as P. flavoaurantia). Spain: Girona: S'Arenella, el Port de la subcapitate, sometimes irregular, thin-walled, clamped. Pleuro- Selva, 18 Nov. 1997, L. Vila & X. Llimona (BCN-SCM B-4064!, cystidia absent. Trama hyphae cylindrical to subinflated, thin- holotype). walled, voluminous, non-dextrinoid, clamped, 5e30 mm wide. Notes: Leucoinocybe taniae is characterized by a small, orange or Pileipellis a cutis composed of ± cylindrical, thin-walled, non- brown-red pileus, decurrent, cream or pale beige lamellae, a root- dextrinoid, clamped, 2e8 mm wide hyphae. Pileocystidia ing, white-pruinose stipe paler than pileus, 7e9 5.5e7.0 mm, 32e140 6e11 mm, subulate, fusoid, lageniform, rarely (sub)cy- (broadly) ellipsoid to subglobose, amyloid basidiospores, and lindrical, slightly to distinctly thick-walled (walls up to 1.5 mm distinct (28‒)44e64 7e12 mm, fusoid, (sub)lageniform, mostly thick), obtuse to subacute, rarely branched or irregular. Stipitipellis (sub)acutely rostrate cheilocystidia. a cutis made up of cylindrical, parallel, ± thin-to slightly thick- “Clitocybula flavoaurantiaˮ morphologically differs only in walled, non-dextrinoid, clamped, 3e7 mm wide hyphae. Caulocys- slightly smaller basidiospores (their size averages vary between tidia 50e150 6e16 mm, subulate, lageniform, fusoid, thick-walled 7.1 and 7.5 mminlengthand4.9e6.0 mminwidthinstudied (walls up to 1 mm thick), sometimes with thin-walled base, sub- specimens) and slightly broader caulocystidia (varying between acute to obtuse, clamped; cylindrical or (narrowly) clavate, mostly 8and23mm). Phylogenetically, ITS rDNA sequences derived irregular, thin-walled, 2.5e5.0 mm wide terminal cells also present. from holotypes of C. taniae (LT854057) and C. flavoaurantia Ecology and Distribution: On soil (twigs under the soil surface) (HM191744) differed in six unambiguous substitutions (0.94 % and on wood of Q. ilex. It is widely distributed in the Mediterranean from 641 bp) and three additional sites were heterozygotic, region in Europe and North Africa. containing shared as well as unique nucleotide. These phylo- Specimens examined: Algeria:Foret^ de la Reghaïa, a terre (sur genetical differences as well as the morphological ones are too brindilles enterrees), 18 Nov. 1923, R. Maire 8285 (MPU!, lectotype). small for a differentiation of both species. Therefore, we Italy: Sardinia: Cala Gonone, on wood of Q. ilex, 31 Oct. 1985, C. Rossi consider the two taxa conspecific. (BOZ!, epitype). Other taxa belonging to Leucoinocybe. Raj & Manimohan (in Notes: Leucoinocybe lenta is characterized by having rather Deepna Latha et al., 2015) described a new species Clitocybula sul- small, amyloid basidiospores, only scattered, clavate, fusoid chei- cata from India, which phylogenetically undoubtedly belongs to the locystidia, and subulate, fusoid, lageniform, rarely (sub)cylindrical, genus Leucoinocybe (Fig. 1). This fact is also supported by its macro- and slightly to distinctly thick-walled pileo- and caulocystidia. and micromorphological characters. The formal new combination Leucoinocybe taniae (Vila) Antonín, Borovicka, Holec & Kolarík, is proposed below. comb. nov. Leucoinocybe sulcata (K.N.A. Raj & Manim.) Antonín, Borovicka, MycoBank MB 825131. Holec & Kolarík, comb. nov. Basionym: Clitocybula taniae Vila, Rev. Catalana Micol. 24: 283, Mycobank MB 825132. 2002. Basionym: Clitocybula sulcata K.N.A. Raj & Manim., Phytotaxa Pseudoomphalina flavoaurantia Contu, Micol. Veg. Medit. 18 (1): 208 (1): 65, 2015. 65, 2003; Clitocybula flavoaurantia (Contu) E.F. Malysheva, O.V. Holotype: India: Kerala State: Kozhikode District, Atholi, 6 Aug. Morozova & Contu, Sydowia 63 (1): 98, 2010. 2011, K. N. Anil Raj, AR791 (CAL 1246). V. Antonín et al. / Fungal Biology 123 (2019) 431e447 443

Lignomphalia Antonín, Borovicka, Holec & Kolarík, gen. nov. trama lamellarum maturitate cum hyphis valde inflatis, tandem MycoBank MB 825129. crasse tunicatis; rhizomorphis dimiticis e hyphis generativis et Diagnosis: It differs from omphalinoid species of the genus sceleticis compositis. Leucoinocybe by its separate phylogenetic position, ochraceous, Type species: Collybia platyphylla (Pers.: Fr.) P. Kumm. yellowish or brightly orange stipe, and especially by its lignicolous Genus characteristics: Basidiomata collybioid or clitocyboid, way of life. pileus surface fibrillose, delicately radially streaked, or, in age, Type species: Pseudoomphalina lignicola Lj.N. Vassiljeva (≡ rimose, lamellae adnexed to adnate, usually with a tooth merging Lignomphalia lignicola (Lj.N. Vassiljeva) Antonín, Borovicka, Holec & into the longitudinal ridges of the stipe, usually broad, basidio- Kolarík (for valid combination see below)). spores inamyloid, Cheilocystidia present, typically broadly clavate Etymology: Omphalia growing on wood (lignum). (see Hughes et al. (2007)). Description: Pileus clitocyboid or omphalioid, usually distinctly Megacollybia platyphylla (Pers.: Fr.) Kotl. & Pouzar, Ceska infundibuliform with deeply depressed centre, glabrous or slightly Mykol. 26 (4): 220, 1972. pruinose to minutely scaly at centre, ochraceous, umbrinous, clay Basionym: Agaricus platyphyllus Pers., Observ. Mycol. 1: 47, 1796. coloured, orange, or lightly red-brown; lamellae deeply decurrent, Collybia platyphylla (Pers.: Fr.) P. Kumm., Führer Pilzk.: 117, 1871; cream coloured or yellowish; stipe cylindrical or broadened to- Gymnopus platyphyllus (Pers.: Fr.) Murrill, N. Amer. Fl. 9 (5): 367, wards base, pruinose or minutely pubescent, yellowish at apex and 1916; Hydropus platyphyllus (Pers.: Fr.) Kühner, Bull. Mens.Soc. Linn. darker at base, ochraceous, yellowish or brightly orange; spore Lyon 49: 895, 1980; Oudemansiella platyphylla (Pers.: Fr.) M.M. print white; basidiospores thin-walled, amyloid; cheilocystidia Moser, Kl. Kryptogamenfl. e Die Rohrlinge€ und Blatterpilze€ (Agar- present; pleurocystidia absent; pileipellis a cutis made up of radi- icales) IIb/2: 156, 1983; Tricholomopsis platyphylla (Pers.: Fr.) Singer, ally arranged hyaline hyphae; pileo- and caulocystidia present; Schweiz. Z. Pilzk. 17: 56, 1939; Clitocybula platyphylla (Pers.: Fr.) clamp connections present. Malençon & Bertault, Fl. Champ. Super. Maroc 2: 398, 1975 (invalid Ecology: On trunks or decayed wood of coniferous, more rarely combination, basionym not indicated, The Code Art. 41.5); Clito- deciduous trees. cybula platyphylla (Pers.: Fr.) E. Ludw., Pilzkompendium 1: 58, 2001. Notes: At the present state of knowledge, the genus Lignom- Agaricus grammocephalus Bull., Herb. France 13: t. 594, 1793; phalia contains only the type species. Collybia grammocephala (Bull.) Quel., Fl. Mycol. France Pays Limi- Lignomphalia lignicola (Lj.N. Vassiljeva) Antonín, Borovicka, trophes: 228, 1888. Holec & Kolarík, comb. nov. Type specimen: Holotype probably not existing. Neotype not MycoBank MB 825144. designated for the time being, probably due a large infraspecific Basionym: Pseudoomphalina lignicola Lj.N. Vassiljeva, Agarikovye variability (presence of three genetic types, for discussion see Shlyapochnye Griby (por. Agaricales) Primorskogo Kraya: 153, 1973. Hughes et al., 2007). Clitocybula lignicola (Lj.N. Vassiljeva) E.F. Malysheva & O.V. Selected macroscopic descriptions and iconography: Ryman and € Morozova, Sydowia 63 (1): 91, 2011. Holmåsen (1984); Breitenbach and Kranzlin (1991); Boekhout Type specimens: Holotype: Russia, Russian Far East: Primorye (1999); Ludwig (2000, 2001); Hughes et al. (2007); Eyssartier and Territory, Ussurijsky District, Suputinsky (¼ Ussurijsky) Reserve, Roux (2011); Holec (2012). valley of Suputinka River, mixed forest, on a decaying wood of Microscopic description: Basidiospores (7.0‒)7.5e10(‒ coniferous tree, 16 Sep. 1961 (lost, see Malysheva et al., 2011). e 11) 6e8(‒9) mm, average 8.6 6.7 mm, E ¼ 1.13e1.38(‒1.46), Lectotype: Russia: Sakhalin Island: vicinities of Novo-Aleksandrovsk, Q ¼ 1.27e1.28, subglobose, broadly ellipsoid, subovoid, thin-walled, flood plain forest with Alnus hirsuta, 20 Sep. 1960, Lj. N. Vassiljeva smooth, non-dextrinoid, inamyloid. Basidia 40e45 8e12 mm, 4- (VLA M-1462, designated by Malysheva et al., 2011: 91, not studied spored, clavate, clamped. Basidioles 15e45 2.5e10.0 mm, by us). Malysheva et al. (2011) erroneously used the term neotype clavate, cylindrical, clamped. Cheilocystidia numerous, for this collection which represents lectotype in fact as it was 35e82 7e25 mm, clavate, fusoid, sometimes subcapitate and selected from the original material collected by Vassiljeva and kept pedicellate, rarely irregular or with projection(s), thin-to slightly in VLA (see The Code Arts. 9.3, 9.4, 9.8, 9.11e9.13). This error in use thick-walled, clamped. Pleurocystidia absent. Trama hyphae cylin- of a term denoting type is corrected here in accordance with The drical to subinflated, thin-walled, non-dextrinoid, clamped, Code, Art. 9.10. 3e20 mm wide. Pileipellis a subhymeniderm at centre when young, Selected descriptions: Malysheva et al. (2011), Shiryaeva (2016). ± a cutis otherwise; terminal cells 40‒82 9‒16 (‒22) mm, clavate, Notes: This species is morphologically similar to some species of narrowly clavate, (sub)fusoid, sometimes with apical projection(s), Leucoinocybe. However, it differs from them by its lignicolous way thin-to slightly thick-walled, clamped, colourless to brown in KOH. of life. Phylogenetically, it has an isolated position outside of Cli- Stipitipellis made up of cylindrical, parallel, ± slightly thick-walled, tocybula and Leucoinocybe species (Fig. 1). non-dextrinoid, clamped, 2e7 mm wide hyphae. Caulocystidia It was described from the Russian Far East and it is widely single or in groups, adpresed to (sub)erect, 30e75 6e15 (‒21) mm, distributed in Siberia (Malysheva et al., 2011). In 2016, however, clavate, narrowly clavate, fusoid, subcylindrical, subcapitate and collections from the European part of Ural and the Russian East pedicellate, sometimes irregular, thin-to slightly thick-walled, European Plain were published by Shiryaeva (2016). clamped. Ecology and Distribution: Solitary or gregarious, on or along & Megacollybia Kotl. Pouzar, Ceska Mykol. 26 (4): 220, 1972. decaying wood of especially broadleaved trees (most common on Original description: Carposomata pileata, collybioidea, Fagus, Betula and Quercus, Holec, 2012), less frequently conifers, camose succosa, lamellata, stipite centrali basi rhizomorphis usually connected by mycelial cords with wood. This species is crassiusculis ramosisque proviso; lamellis crassis, latis, dente widely distributed in Europe, Scandinavia, western and central adnatis; cute pilei grisea vel griseo-brunnea; pulvere sporarum Russia (Hughes et al., 2007). albo; sporis ovoideis vel late ellipsoideis, pariete glabra, tenui, Specimens examined: Czech Republic: Labske pískovce Land- inamyloidea; carne pilei atque stipitis e hyphis monomiticis, scape Protected Area, Hrensko, SuchaBela valley, decaying stem of nodoso-septatis; cute pileorum juvenilium centro pilei hyme- Picea, 27 Jun. 2002, V. Antonín 02.39 (BRNM 670807). Bíla, Salajka niformi, postea e cellulis distincte separatis, clavatis constituta; National Nature Reserve, on base of a dead stem of Picea, 29 Jul. 444 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

2011, V. Antonín & D. Janda VA 11.68 (BRNM 737297). Drahonín, Trenckova rokle valley, near the decaying coniferous stem, 30 Jul 2009, S. Kubesova (BRNM 724528). Bedrichov, Hersica Nature Monument, stump of Picea, 7 Sep. 2002, Z. Bieberova (BRNM 724818). Brno-Bystrc, Horak uv zleb valley, on wood remnants of a broadleaved tree, 24 May 2012, V. Antonín & H. Sevcíkova (BRNM 737654). Vranovice nad Svratkou, Plackuv les Nature Reserve, fallen mossy Salix stem, 26 Jul. 2000, A. Vagner (BRNM 664912). Bíle Karpaty Landscape Protected Area, Straní, VelkaJavo rina National Nature Reserve, decaying wood of a broadleaved tree, 24 Aug. 2003, A. Vagner (BRNM 704823). Lednice, Pastvisko National Nature Reserve, on soil under Fraxinus, 3 Oct. 2001, Z. Bieberova (BRNM € 728280). Hungary: Mecsek Mts., Pecs, Obanya near Mecseknad asd, Kis-Tuft, on fallen branch of Fagus, 3 Jun. 1987, V. Antonín (BRNM 462110). Slovakia:Kacín, a hornbeam stand with birch, 16 Sep. 1975, K. Kríz (BRNM 305724). Notes: Megacollybia platyphylla is characterized by having robust basidiocarps, a distinctly radially fibrillose to scaly, beige-brown to grey-brown pileus, distant and very broad lamellae, distinct white mycelial cords at stipe base, inamyloid basidiospores, and distinct clavate or fusoid cheilocystidia. In our phylogenetic analysis, this species formed a relativelly well supported clade (0.91/98), sister to M. marginata. Similarly like C. lacerata, it represents an aggregate species (Hughes et al., 2007) deserving further taxonomic study. Megacollybia marginata R.H. Petersen, Morozova & J.L. Mata, Rep. Tottori Mycol. Instit. 45: 21, 2007 (Figs. 10 and 11). Holotype: Russia: Primorsky Reg.: Khasansky Dist., Kedrovaya Pad Nature Reserve, 20 Aug. 2005, O. Morozova (as M. platyphylla), Fig. 11. Megacollybia marginata (TENN 60752, holotype). (A) basidiospores; (B) chei- TFB 11869 (TENN 60752!). locystidia; (C) pleurocystidia; (D) caulocystidia. Scale bar ¼ 20 mm. Del. V. Antonín. Macroscopic descriptions: Hughes et al. (2007); Kim et al. (2014). Description of own collections: Pileus 35e150 mm broad, low- convex, applanate to slightly funnel-shaped, with slightly rarely subglobose, thin-walled, smooth, inamyloid. Basidia 33‒ depressed centre and involute to straight margin, with very small 52 8.5e11.5 mm, 4-spored, clavate, clamped. Basidioles 15‒ obtuse papilla within central depression, except for centre radially 40 3e12 mm, clavate, less frequently subfusoid or subcylindrical, fibrillose or fibrillose-squamulose, grey-brown (6E‒F5, 7E‒F4), clamped. Cheilocystidia 30‒65 8‒20 mm, clavate, broadly clavate, slightly paler (6D4) among fibrils. Lamellae distant, L ¼ c. 40, l ¼ 2‒ spathulate, often pedicellate, thin-walled, clamped, colourless or 4(‒5), broadly adnate with small tooth, broad (up to 15 mm), with pale brown contents in KOH. Pleurocystidia scattered some- intervenose ad base when old, very pale cream with dark edge times absent, similar to cheilocystidia but predominantly clavate (pileus colour). Stipe 60‒90 5‒10 mm, cylindrical or laterally and smaller (34‒45 9‒13 mm). Trama hyphae cylindrical, ± thin- compressed with groove, not broadened at base, finely pale grey- walled, colourless, 3‒20 mm wide. Pileipellis a cutis with transition brown fibrillose-squamulose at apex, finely longitudinally fibril- to subtrichoderm (centre), of cylindrical, ± thin-walled, terminal lose, white or whitish, with white or whitish rhizoids at base. cells 18‒72 6‒14 mm, clavate, fusoid, subutriform, sometimes Context white, grey-brown under pileipellis, hollow in stipe, (long) pedicellate, thin-to slightly thick-walled, sometimes irreg- without any distinct smell. ular, clamped, with black-brown contents. Stipitipellis a cutis of Basidiospores (7.0‒)7.5e9.5 (5.0‒)5.5e7.0 (‒7.5) mm, average cylindrical, thick-walled (±1 mm), smooth, clamped, 2‒5 mm wide 8.18 6.12 mm, E ¼ (1.16‒)1.30e1.60, Q ¼ 1.36, broadly ellipsoid, hyphae. Caulocystidia 25‒72(‒110) 5‒18(‒29) mm, clavate, broadly clavate, fusoid, capitate, often pedicellate, sometimes rostrate, slightly thick-walled, clamped. Ecology and Distribution: On or near very rotten wood of Alnus? (holotype), on a strongly decayed wood of P. abies, Fagus sylvatica (Czech Republic) and a broadleaved tree and wood remnants in a mixed forest (Republic of Korea). It was described from eastern Russia, collected also in East Asia, newly recorded from Europe (see below). Specimens examined: Czech Republic:StríbrnaSkalice, Vlkancice ‒ StaraH ura, Baba Hill, 24 Jul. 2012, J. Borovicka (PRM 860926). Nedvedice near Sobeslav, V Starem forest, 26 Aug. 2010, F. Kotlaba (PRM 859785, as M. platyphylla). Panska Habrova, 30 Jun. 2012, T. Tejklova & H. Tejklova (HR 90203, as M. platyphylla). Trckov, Trckov National Nature Reserve, 13 Sep. 2011, T. Tejklova (HR 91607, as M. platyphylla); Ibid., 4 Sep. 2012, Z. Egertova & T. Tejklova (HR 91624, as M. platyphylla). Republic of Korea: Hongcheon Prov.: Dong-myeon, 17 Aug. 2013, V. Antonín & R. Ryoo VA 13.176 (BRNM 801861). Kangwon Prov.: Fig. 10. Megacollybia marginata (PRM 860926). Basidiomata. Photo J. Borovicka. Pyeongchang County, Dae-hwa-myeon, Mt. Kariwang, 21 Aug. V. Antonín et al. / Fungal Biology 123 (2019) 431e447 445

2012, V. Antonín, K.-H. Ka & R. Ryoo VA 12.135 (BRNM 747514). R.W. Kerrigan (PRM 861168); ibid., 15 Jul. 2012, J. Borovicka (BRNM Taean Peninsula: Deoksung, Sudeoksa Monastery, 8 Jul. 2014, 781115). Connecticut: New Haven, West Rock Ridge State Park, near V. Antonín & R. Ryoo VA 14.51 (BRNM 801862). Russia: Primorsk Lake Wintergreen (south edge), 15 Jul. 2012, J. Borovicka & E.C. County: Hasansky, town of Primorsk, 20 Aug. 2005, O. Morozova Vellinga (PRM 861169). (TENN 60752!, holotype). Notes: We included this species in our study as we had a good Notes: Megacollybia marginata is macroscopically very close to material at hand for comparison with European collections. the common European species M. platyphylla. It macroscopically M. rodmanii is macroscopically very close to M. platyphylla. It differs differs only by a constantly dark coloured lamellar edge due to the by less fleshy basidiomata, mostly globose or subglobose basidio- darker contents of cheilocystidia (however, one of infraspecific spores, larger cheilocystidia, more voluminous trama hyphae and forms of M. platyphylla marked as “A” by Hughes et al., 2007 may pileipellis in a form of a cutis also in centre. Neverthelles, its sometimes have a very delicately marginate to dark purple lamella phylogenetic position is outside of the clade containing edge; for a discussion see there), and by distinct DNA sequences. M. platyphylla, M. marginata, M. clitocyboidea and M. falax (Fig. 1). It Hughes et al. (2007) considered it apparently endemic to far forms a well supported lineage together with other American eastern Russia. The collections from the Czech Republic represent species Megacollybia texensis and Megacollybia subfurfuracea. the first published records not only for this country, but also for the whole Europe. The records from the Republic of Korea have already been published (Kim et al., 2014). Other consequent taxonomical results and comments Hughes et al. (2007) mentioned the presence of one specimen from the Czech Republic and one from Caucasus (Russia, Teberda) As shown in our phylogram (Fig. 1), the genera Trogia, Gerro- with sequences fitting phylogenetically to M. marginata (Fig. 1), but nema and Hydropus form separate clades. However, our study was lacking marginate gills, which suggest that the value of this char- not focused on species of these groups. Below we discuss only their acter should be further studied. representatives which were formerly placed to Clitocybula s. lato. Phylogenetically, M. marginata specimens were clustered Hydropus dryadicola (Kühner) E. Horak, Sydowia 39: 117, 1987. together, but with a minor statistical support due to the relatively Basionym: Clitocybula dryadicola Kühner, Sydowia 36: 165, 1983. high intragroup variability and limited variability from Holotype: Switzerland: Grisons: partie haute du Val Schombrina, M. platyphylla. This suggests that more variable alternative genetic 29 Aug. 1966, R. Kühner (G 59278!). markers should be used for future taxonomic studies. Selected macroscopic descriptions: Kühner (1983); Horak (1987); Senn-Irlet (1987). Extra-European Megacollybia species studied Microscopic description: Basidiospores of the holotype: (8.0‒) 8.5e11 (‒12) (5.0‒)5.5e7.5 (‒8.0) mm, average 9.7 6.5 mm, Megacollybia rodmanii R.H. Petersen, K.W. Hughes & E.B. E ¼ 1.30e1.80, Q ¼ 1.50; BERN collection: (8.5‒)9.0e11.5 (5.5‒) Lickey, Rep. Tottori Mycol. Instit. 45: 42, 2007. 6.5e8.0 (‒8.5) mm, average 9.8 7.3 mm, E ¼ 1.12e1.64, Q ¼ 1.35; LIP Holotype: USA, Tennessee, Sevier Co., vic. Gatlinburg, GSMNP, collection: 8.5e11.5 (‒12) 6.5e8.0 (‒9.0) mm, average Greenbrier, Porter's Creek Rd., 5 May 2002, R.H. Petersen & K.W. 9.6 7.4 mm, E ¼ 1.18e1.71, Q ¼ 1.31], (broadly) ellipsoid to Hughes, TFB 11485 (TENN 59430!). ellipsoid-fusoid, rarely broadly subamygdaloid, thin-walled, Macroscopic description and iconography: Hughes et al. (2007). smooth, non-amyloid. Basidia 26e45 (7.5‒)9.5e13 mm, 1-, 2-, 3- Microscopic description:Basidiospores7e9(‒11) (5.5‒)6.5e8.0 and 4-spored (2-spored usually dominating), clavate, sometimes (‒8.5) mm, average 8.0 6.9 mm, E ¼ 1.0e1.32, Q ¼ 1.15, globose, with median constriction, clampless or with very rare clamp con- subglobose to broadly ellipsoid, thin-walled, smooth, inamyloid. nections. Basidioles 1535 5e11 mm, clavate, (sub)fusoid or (sub) Basidia 31‒50 8e12 mm, 4-spored, clavate or fusoid. Basidioles cylindrical. Cheilocystidia indistinct (marginal cells), (18‒) (10‒)20‒40 3e12 mm, clavate, subfusoid or cylindrical. Cheilocys- 21e40 (3‒)5e10(‒13) mm, clavate, fusoid, subcylindrical, mostly tidia 33‒64 (7‒)10‒19 (‒24) mm, clavate, broadly clavate, spathu- irregular, sometimes subcoralloid or rostrate, thin-walled. Pleuro- late, subfusoid, often pedicellate, sometimes 2-celled, irregular to cystidia absent. Trama hyphae cylindrical to subinflated, thin- or branched, thin-walled, clamped, colourless; on addition, 29‒ slightly thick-walled, non-dextrinoid, clamped or clampless, 40 3.5e8.5 mm, subfusoid to cylidrical, branched cells are present 4e25 mm wide. Pileipellis a cutis composed of ± cylindrical or in the type specimen. Pleurocystidia absent. Trama hyphae cylin- subinflated, thin-walled, non-dextrinoid, clamped or clampless, drical, thin- and slightly thick-walled, colourless, non-dextrinoid, 3‒ 3e9 mm wide hyphae; terminal cells 15e30 4e8 mm, adpressed to 40 mm wide. Pileipellis a cutis made up of cylindrical, radially ar- erect, cylindrical, fusoid, (narrowly) clavate, obtuse, thin-walled. ranged, ± thin-walled, 3e8.0 mm wide hyphae; terminal cells 25‒ Stipitipellis a cutis made up of cylindrical, parallel, ± thin-walled, 80 (4.5‒)8e15 (‒25) mm, clavate, fusoid, subutriform, sometimes non-dextrinoid, clamped or clampless, 3e6 mm wide hyphae. (long) pedicellate, thin-to slightly thick-walled, sometimes irregular, Caulocystidia mostly in groups, 14e35 4.0e8.5 mm, fusoid, cy- with black-brown contents. Stipitipellis a cutis of cylindrical, thick- lindrical, clavate, thin-walled. walled (±1 mm), smooth, clamped, 2e6 mm wide hyphae. Caulocys- Ecology and Distribution: In small groups in alpine zone, on soil tidia 29e85 (‒105) (4‒)6‒14 mm, clavate, fusoid or cylindrical and on basic (calcareous) substrate in Dryas octopetala, Salix retusa, capitate, often pedicellate, thin-to slightly thick-walled. All tissues S. helvetica and S. herbacea stands, and on raw humus in the Saxi- clamped, but clamps not at all septa and sometimes loop-like. fragi-Caricetum frigidae association. It is known only from France Ecology and Distribution: On or close to very rotten wood; when and Switzerland till now (Kühner, 1983; Horak, 1987, studied on forest debris, then probably attached to buried wood in hard- specimens). wood forests. It is known from Eastern North and Central America Studied specimens: France: F. Savoie: Bessans, L'Ecot, Plan de (Costa Rica to Newfoundland; Hughes et al., 2007). Evettes, 13 Aug. 1995, P.-A. Moreau 95081308 (LIP). Switzerland: Studied specimens: USA: Tennessee: Sevier County, Gatlinburg, 5 Grisons: partie haute du Val Schombrina, 29 Aug. 1966, R. Kühner (G May 2002, R.H. Petersen & K.W. Hughes (TENN 59430!, holotype); 59278, holotype). Wallis: St. Luc, Les Faches, 2342 m alt., 30 Aug. Connecticut: New Haven, West Rock Ridge State Park, 15 Jul. 2012, 1985, B. Senn-Irlet (BERN). 446 V. Antonín et al. / Fungal Biology 123 (2019) 431e447

Acknowledgements

The authors wish to thank curators of the herbaria BCN, BERN, BRNM, CB, G, GDOR, HR, LE, LIP, NY, NYS, PRM, S and Francesco Bellù (Bolzano, Italy) for a kind sending us the type and other herbarium specimens. The studies of the first author were enabled by the support provided to the Moravian Museum by the Ministry of Culture of the Czech Republic as part of its long-term conceptual development program for research institutions (DKRVO, ref. MK000094862). The work of the second author was supported by the Long-term Development Project RVO67985831. The work of the third author was financially supported by the Ministry of Culture of the Czech Republic (DKRVO 2018/08, National Museum, 00023272). The collecting trips of the first author to the Republic of Korea were supported by project FP 0801-2010-01 of the National Institute of Forest Science, Seoul; this author also thanks Kang- Fig. 12. Gerronema wildpretii (BRNM 788347). Basidiomata. Photo J. Borovicka. Hyeon Ka and Rhim Ryoo (National Institute of Forest Science, Seoul) for collaborating on field excursions. The authors thank Pierre-Arthur Moreau (Lille, France) for valuable comments to the fi Notes: This species is characterized by having a small, 4e16 mm typi cation of Mycena (Leucoinocybe) lenta and Veronique Bour- ^ fi broad, grey-brown or sepia brown, then pale brown-grey or gade and Caroline Loup (Herbier MPU, Pole Patrimoine scienti que, brownish yellowish (when drying-out) pileus, decurrent lamellae, a Universite Montpellier 2, France) for the access to Maire and Mal- 6e21 1e2 mm, yellowish grey, sometimes brownish ochraceous ençon's collections. Collections of North American Clitocybula stipe, (broadly) ellipsoid to ellipsoid-fusoid, non-amyloid basidio- samples provided by Renee Lebeuf and Eric Smith are also greatly spores, and indistinct cheilocystidia (marginal cells). Kühner (1983) appreciated. The authors also thank Luis Alberto Parra Sanchez mentioned the presence of 2-spored and clampless, and 4-spored (Aranda de Duero, Spain) for very valuable nomenclatorical com- and clamped basidiocarps. However, according our studies, 1-, 2-, ments to the genus Leucoinocybe and Ladislav Ilkovics (Masaryk 3-, and 4-spored basidia may be present in one basidiocarp, and the University, Medical Faculty, Department of Histology and Embry- presence/absence of clamps is very irregular. Kühner (1983) also ology, Brno, Czech Republic) for providing the SEM microphoto- mentioned the size of basidiospores as 5.7e7 4.7e5.5 mm (4- graph of C. tilieti. spored form), and 7e10 5.7e7 mm (2-spored form). Basidio- spores of both collections studied by us have the same length but References differ by their broadness (see above). Although the sequencing of the studied specimens was not Antonín, V., Ryoo, R., Shin, H.-D., 2008. Gerronema nemorale (Basidiomycota, successful, we are convinced similarly like Horak (1987) that this Agaricomycetes): anatomic-morphological, cultivational, enzymatic and mo- lecular characteristics and its first records in the Republic of Korea. Czech taxon belongs to the genus Hydropus. The main characters sup- e fi Mycol. 60, 197 212. porting this classi cation are the non-amyloid basidiospores and Antonín, V., Beran, M., Borovicka, J., Dvorak, D., Holec, J., 2011. Clitocybula familia indistinct cheilocystidia (marginal cells). Therefore, the correct (Fungi, Agaricales) e taxonomy, distribution, ecology and first records in the name of this species really is H. dryadicola (Kühner) E. Horak. Czech Republic and Slovakia. Czech Mycol. 63, 1e11. Arora, D., 1986. Mushrooms Demystified: A Comprehensive Guide to the Fleshy Gerronema atrialba (Murrill) Borovicka & Kolarík, comb. nov. Fungi, second ed. Ten Speed Press, Berkeley. Bigelow, E.H., 1973. The genus Clitocybula. 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