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Introduction

Basic works on Gymnopilus taxonomy in Europe

The genus Gymnopilus P. Karst. 1879 (Fungi, Agari-

cales) includes interesting and important saprotrophic,mostly wood-inhabiting fungi occurring all over the world.The history of its study was described by Hesler (1969). Theclassical delimitation of Gymnopilus was summarised byKhner (1980) and Singer (1986). The most important char-acters of Gymnopilus are: presence of lamellae, stipe nevereccentric, presence of a cortinoid to membranaceous veil,taste mostly bitter, spore print rusty brown (ferrugineous-fulvous), spores with verrucose to rugulose ornamentation,no germ pore and mostly dextrinoid wall, presence ofcheilocystidia which are more or less ventricose below andpossess a subcapitate to capitate apex, all hyphae with

clamp connections.In traditional classification, Gymnopilus is placed eitherinto Cortinariaceae (Singer 1986) or, based on its sapro-trophic way of life and presence of styryl-pyrones bis-no-ryangonin and hispidin (e.g. Dangy-Caye et Arpin 1974,Rees et Ye 1999, Rees et Strid 2001), into Strophariaceae(Khner 1980).

The genus has never been monographically elaboratedin the Czech Republic and the same counts for the whole ofEurope. There are several recent studies from some coun-tries or larger areas, which differ in their taxonomic con-cepts and resulting number of accepted taxa, quality ofelaboration (both factual and formal) and number of collec-

tions studied.

1. The classical concept is represented by Khner et Ro-magnesi (1953), Moser (1983), Orton (1993), Breiten-bach et Krnzlin (2000) and Keller et Moser (2001). Inall these works, the delimitation of taxa is similar (al-

though substantial differences occur) and the number ofspecies recognised is about 1015. The results are basedon a limited number of collections studied and problem-atic points are not discussed. A nice summary of theseworks are the descriptions (partly original, partly com-piled) and perfect iconography published by Ludwig(2000, 2001).

2. A detailed monographic study of species occurring inNorway (Hiland 1990) based on a large number of col-lections, detailed analysis of some characters and an ef-fort to respect the original concepts of the species. Sometaxonomic and nomenclatural problems are discussed

and some solutions are proposed (e.g. a proposal to re-place the type species of the genus Gymnopilus G.liquiritiae by G. picreus or a synonymisation of thenames G. sapineus, G. penetrans and G. hybridus).However, the number of taxa treated is rather low (6).

3. A survey of European taxa (Bon et Roux 2002) using aconcept of narrow species and trying to present manytaxa previously omitted or not known from Europe. Theauthors recognise 24 species. Unfortunately, the numberof collections studied is low and some formal errors de-crease the quality of the work (poorly reproduced pho-tographs, poor quality of line drawings, many smallerrors or omissions in the text). However, some newideas are introduced which deserve further research.

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Acta Musei Nationalis Pragae, Series B, Historia Naturalis, 61 (12): 152 issued June 2005Sbornk Nrodnho muzea, Serie B, Prodn vdy, 61 (12): 152

THE GENUS GYMNOPILUS (FUNGI, AGARICALES) IN THE CZECH REPUBLIC WITH

RESPECT TO COLLECTIONS FROM OTHER EUROPEAN COUNTRIES

JAN HOLECNational Museum, Mycological Department, Vclavsk nm. 68, 115 79 Praha 1, Czech Republic, [email protected]

Holec, J. (2005): The genus Gymnopilus (Fungi, Agaricales) in the Czech Republic with respect to collections from other Eu-ropean countries. Acta Mus. Nat. Pragae, Ser. B, Hist. Nat., 61 (12): 152. Praha. ISSN 0036-5343.

The genus Gymnopilus (Fungi, Agaricales, Cortinariaceae) was monographically studied. The study is based on a thoroughobservation of macro- and microcharacters using numerous personal and herbarium collections from the Czech Republic. Se-lected collections from Austria, Bulgaria, Croatia, Finland, Germany, Italy, Poland, Romania, Russia, Slovakia, Sweden, andUkraine were also used for a better understanding of the species concept. In each species, macro- and microcharacters, fruc-tification period, substrates, relation to vegetation and altitude, and distribution are evaluated in detail. Taxonomic remarks onall species are added and contemporary literature on Gymnopilus is summarised. A key for the identification of species gro-wing in the Czech Republic is provided. The following species were recognised in this area: Gymnopilus spectabilis (taxon

with robust fruitbodies), G. bellulus, G. josserandii, G. flavus, G. fulgens, G. decipiens, G. penetrans (= G. hybridus), G. sa-pineus and G. picreus. The following names commonly used in contemporary literature are either not recognised in the CzechRepublic or considered doubtful or hardly interpretable taxa: Gymnopilus stabilis, G. junonius s. str. (taxon with slender fru-itbodies), and G. liquiritiae. Line drawings and photographs of important microcharacters are provided for each species. Co-lour photographs of 6 species are added.

macromycetes, Basidiomycetes, Cortinariaceae, Gymnopilus, taxonomy, nomenclature, ecology, substrates, distribution,macrocharacters, microcharacters, identification key

Received March 8, 2005


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Except for these basic publications, many valuable pa-pers dealing with one or several species have been pub-lished. They are discussed in the chapter Results. Forexample, new European taxa were described by Romagnesi(1976: Gymnopilus corsicus, G. spadiceus; 1979: G.

pseudofulgens) and recently by Moser et al. (2001:

Gymnopilus turficola). A find of Gymnopilus corsicus isdiscussed by Maurice (2001).

My work on Gymnopilus was started after finishing theEuropean monograph of the genus Pholiota Holec (2001a).Some excluded or doubtful taxa of Pholiota proved to beGymnopilus species. Consequently, I tried to elaborate thisgenus in a detailed way, at least in the Czech Republic andsome adjacent regions. In the preparatory phase, severalshorter contributions were published (Holec 2001b, 2002,Holec et al. 2003). This publication is a summary of mystudies on Gymnopilus in the period 20012004.

Aims of this study

1. To check which species are known from the CzechRepublic.

2. To evaluate their delimiting characters.3. To produce thorough descriptions of macro- and mi-

crocharacters based on personal observations and studyof a large number of fresh and herbarium collections.

4. To compare and evaluate different species concepts usedin Europe.

5. To clear up some problems in Gymnopilus taxonomy andnomenclature.

Survey of recent literature on Gymnopilus

Publications on the taxonomy of Gymnopilus in Europehave been cited above. There are also floras or check lists ofGymnopilus species from some countries or regions inEurope. They are cited in the chapter Methods, paragraphData on distribution of Gymnopilus species in Europe.

In the past decades, the genus Gymnopilus in Mexico,Central America and the Caribbean has been taxonomicallystudied by Laura Guzmn-Dvalos, sometimes with collab-orators (e.g. Guzmn-Dvalos 1994, 1995, 1996a, 1996b,1997; Guzmn-Dvalos et Guzmn 1986, 1995; Guzmn-Dvalos et Ovrebo 2001). Recently, Guzmn-Dvalos(2003) started with type studies in Gymnopilus, while pub-

lishing the results obtained in 22 species, mostly from Cen-tral and North America.New species and new records from India are described

by Thomas et al. (2003) and older works on Gymnopilusfrom India are cited.

Australian species of Gymnopilus and their relationshipto Northern Hemisphere taxa were studied by Rees (2003),Rees et Lepp (2000), Rees et Strid (2001), Rees et Ye(1999), and Rees et al. (1999, 2002). Red to purple-colouredspecies from Europe and Southern Hemisphere were com-pared (using classical and molecular methods) by Rees et al.(2004). They confirmed the existence of only two separatespecies: Gymnopilus dilepis and G. purpuratus.

The ultrastructure of the spore wall of some Gymnopilusspecies was studied by Clmenon (1974, in the TEM),

Keller (1997, in TEM and SEM) and Rees, Orlovich etMarks (1999, in SEM).

The works on the presence of styryl-pyrones inGymnopilus species were summarised by Hiland (1990).There are also studies on the presence of hallucinogenoussubstances (e.g. psilocybin, psilocin and baeocystin) in

some species of Gymnopilus (e.g. G. purpuratus) publishede.g. by Hatfield et al. (1978), Kreisel et Lindequist (1988),Gartz (1989), Giacomoni (1997) and summarised by Stijve(1995). Gymnopilus spectabilis is considered a hallucino-genic fungus in Japan (Stijve 1995), however, it does notcontain psilocybin but a substance from the group of neuro-toxic oligoisoprenoids (Tanaka et al. 1993). Stijve etKuyper (1988) proved that Gymnopilus spectabilis and G.

fulgens do not contain psilocybin or any related tryptaminederivates. Using the lignin test, Kln (1990) showed that G.sapineus does not contain amanitins or tryptaminederivates.

Mycelial morphology, rhizomorph anatomy and pri-mordium formation of Gymnopilus penetrans was studiedby Clmenon (2002). Cultural studies of 5 species wereperformed by Fausto-Guerra et al. (2002). Both brown-rotand white-rot species were recorded, but all species weremainly cellulose decomposers. Older publications on thecultural characters of Gymnopilus were summarised in thementioned works. Enzyme activity of the mycelium of G.hybridus was studied by Kln et Baudiov (1990: 207).

The only cladistic analysis based on macro- and mi-crocharacters of 6 species from Norway was performed byHiland (1990).

Quite recently, some Gymnopilus species were studied

using DNA techniques, mostly an analysis of ITS sequencesof the nuclear ribosomal gene (for a survey of these works seeGuzmn-Dvalos et al. 2003). Gymnopilus sapineus, G.

penetrans and G. spectabilis (used as an outgroup) were in-cluded by Peintner et al. (2001) in their study of sequestratefungi belonging to Cortinariaceae. Similarly, 5 species ofGymnopilus were used by Thomas et al. (2002) when de-scribingAnamika, a new genus of Cortinariaceae related to

Hebeloma. Moser et al. (2001) studied phylogenetic relation-ships of their new species Gymnopilus turficola by compar-ing its ITS sequences with 7 species of Gymnopilus.Moncalvo et al. (2002, 4 species studied) found a gymnopi-

loid clade within euagarics which contains the gymnopilusclade and Galerina paludosa. Rees et al. (2002) studied thephylogeny of 30 Gymnopilus species from Australia and theNorthern Hemisphere. They found that Gymnopilus is onlymonophyletic if it includes Galerina eucalyptorum andPyrrhoglossum pyrrhum the type species of Pyrrhoglos-sum. Their next study (Rees et al. 2003) proved that the genusGymnopilus is really monophyletic. Guzmn-Dvalos et al.(2003) analysed DNA sequences of 29 Gymnopilus species totest the traditional infrageneric classification of Gymnopilusbased on the presence of membranaceous or arachnoid veil (2groups: Annulatae and Gymnopilus = Cortinatae, see Ro-magnesi 1943, Singer 1986). The genus proved to be mono-phyletic. However, the traditional groups (subgenera) werenot recovered. Five well-supported clades were identified.

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Data on Gymnopilus from the Czech Republic

There is only a small number of detailed contributionson Gymnopilus taxonomy in mycological literature regard-ing the area of the Czech Republic. Svrek (1965) describeda find of the rare species Gymnopilus fulgens. Antonn (inAntonn et kubla 2000) described the new species

Gymnopilus josserandii, a correct name for the invalidlypublished Gymnopilus subsphaerosporus (Joss.) Khner etRomagn. Recently, several works have been published bythe present author (Holec 2001b, 2002, Holec et al. 2003).

Concerning older literature, Velenovsk (19201922:551513, 918) treated 5 species belonging to Gymnopilus(under the generic names Flammula and Pholiota): G. pene-trans, G. hybridus, G. sapineus, G. liquiritiae, and G.spectabilis. No Pholiota and Flammula described by him asnew species belong to Gymnopilus (Holec 1999).

Citations of Gymnopilus species in mycofloristic andecological publications are of limited value for the taxono-

my as the identification cannot be verified in most cases (noherbarium specimens available, no remarks on macro- andmicrocharacters). For this reason, such references are notincluded here. The best source of information on taxonomy,distribution and ecology of individual species are theherbarium specimens kept at important herbaria in theCzech Republic. They were widely used for the preparationof this publication.

Some reliable mycofloristic contributions can be used asa source of data on the distribution and ecology ofGymnopilus species, e.g. Svrek et Kubika (1971: 107,ofnsk prales: G. bellulus, G. picreus, G. sapineus),ern et K (1972: 123, Ranpurk: G. junonius), Antonnet al. (2000: 62, Cahnov, Ranpurk etc.: G. junonius), An-tonn et Vgner (2000: 69, Podyj National Park: G. hy-bridus, G. picreus, G. spectabilis, G. stabilis), Svrek(1990: 86, Krkonoe Mts.: G. sapineus).

Materials and Methods

Field work and study of macrocharacters

Fresh fruitbodies were collected mainly in the CzechRepublic. The most intensive field work was carried out inthe umava Mountains (= Bohemian Forest), southern Bo-hemia, the esk vcarsko (= Czech Switzerland) National

Park (northern Bohemia) and in the Beskydy Mountains(north-eastern Moravia). Several finds are from Slovakia andAustria. The habitat of all finds was carefully noted, espe-cially forest type, substrate (including tree species and stageof decay), and elevation. Well-developed fruitbodies werephotographed both in the field and in the laboratory.Macrocharacters of all collections were thoroughly observedand recorded. Fruitbodies were dried at 3040 C in a mobileelectric drier. They are deposited in the herbarium of the My-cological Department, National Museum, Prague (PRM).

All fresh fruitbodies were considered for description ofmacrocharacters in this work. The descriptions are comple-mented with data from herbarium documentation (field

notes, photographs, etc.) found in some herbaria (especiallyPRM, BRNM, CB, BRA, W, WU). Consequently, the de-

scriptions are mostly based on rich primary data showingthe variability of Gymnopilus species in Central Europe. Alldeviating or less frequent characters are marked by phraseslike sometimes, in some fruitbodies etc. When freshmaterial was not available, literature data were used andproperly cited. Descriptive terminology is taken from Bas et

al. (1988: 5464). The colour codes are according toKornerup et Wanscher (1981).

Study of microcharacters

The microcharacters were studied on dried fruitbodies(of personal collections and herbarium material). Almostall examinations were made using an Olympus BH-2 mi-croscope. The observations and measurements were madeon material mounted in a 5 % KOH solution. Pigmentationof hyphae of the pileus and stipe cuticle was studied inpure water and their arrangement was observed in radialsections and scalps. Iodine reactions were studied in Melz-ers reagent prepared according to the formula given inMoser (1983), cyanophilous reaction in cotton blue (ac-cording to Kotlaba et Pouzar 1964, Singer 1986) aftershort boiling.

At least five randomly selected cheilocystidia, pleuro-cystidia, and basidia were measured per collection. Thelength of basidia was measured excluding the sterigmata.For spore size measurements, 20 randomly selected maturespores were used per collection (10 spores during visits offoreign herbaria to save time). Immature spores (extremelysmall or having a thin wall and hyaline content) or aberrantspores (1.52 times longer than the normal ones) were notmeasured. Spores were measured without the hilar appendix

(apiculus). All measurements were carried out on prepa-rations from lamellae (not from spore prints) to measureboth personal collections and herbarium specimens in thesame way. The spores were measured directly in prepara-tions at the magnification of 1250x using an eyepiece mi-crometer with a fine scale (basic unit 0.8 m) whichenabled very accurate measurements. The measured valueswere recorded in units of this scale (relative values). Theirconversion to absolute values was calculated after finishingall measurements. Marginal values of spore sizes are givenin brackets and represent at most 10% of all spores mea-sured in each species. These values are not considered for

the purpose of species delimitation and identification.Descriptions of microcharacters are based on a detailedexamination of at least 2 representative collections perspecies. However, all specimens listed in chapter Collectionsstudied were examined, all deviations were recorded and in-corporated into descriptions. Spore size was measured in allcollections studied. Illustrations of microcharacters weredrawn at a magnification of 1250x using a drawing tube.

Standard works used in this work, terminology

Abbreviations of authors names: Brummitt et Powell(1992). Abbreviations of old taxonomic works and data oftheir publication: Taxonomic literature (Stafleu et Cowan

19761988). Journal abbreviations: Botanico-Periodicum-Huntianum (Lawrence et al. 1968). Standard taxonomic,

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nomenclatural, and bibliographic abbreviations: BotanicalLatin (Stearn 1986). Acronyms of herbaria: Index Herbari-orum (Holmgren et al. 1990). Terms for description ofmacro- and microcharacters: Flora Agaricina Neerlandica,vol. 1 (Bas et al. 1988: 5464). The term suprahilar disc in-stead of plage is used here for the smooth area in the region

of the suprahilar depression in accordance with Pegler etYoung (1971: 21) and Rees, Orlovich et Marks (1999).

Data on distribution of Gymnopilus species in Europe

For all species, the following publications were consulted:Dennis et al. (1960: Great Britain), Kreisel et al. (1987: east-ern part of Germany), Hiland (1990: Norway), Krieglsteiner(1991: Germany), Ryman (1992: Norway, Sweden, Finland,Denmark), Orton (1993: Great Britain), Kuyper (1995: TheNetherlands), Breitenbach et Krnzlin (2000: Switzerland),Ludwig (2001: mostly Germany, but also Sweden), Keller etMoser (2001: Austria), Bon et Roux (2002: especially France,rarely Italy, Belgium, Slovakia), kubla (2003: Slovakia), En-derle (2004: the vicinity of Ulm, Germany). These works arenot cited in paragraphs on the distribution of each species. Ex-cept for these collective publications, papers dealing with in-dividual species were used and cited.

Iconography

In the paragraph Selected illustrations, iconographieswere cited as follows:Bon et Roux: Bon M. et Roux P. (2002): Le genre

Gymnopilus P. Karst. en Europe. In: Fungi non de-lineati, vol. 17: 152, Alassio.

Breitenbach et Krnzlin: Breitenbach J. et Krnzlin F.

(2000): Pilze der Schweiz. Vol. 5. 340 p., Luzern.Bresadola: Bresadola J. (19271960): Iconographia myco-

logica. Vol. 128. Milano.Cetto: Cetto B. (19701993): I funghi dal vero. Vol. 17.

3042 figs., Trento.Dhncke: Dhncke R. M. (1993): 1200 Pilze in Farbfotos.

1179 p., Aarau.Fries: Fries E. (18671884): Icones selectae Hymenomyce-

tum nondum delineatorum, vol. 12. Stockholm.Hagara et al.: Hagara L., Antonn V. et Baier J. (1999): Hou-

by. [Fungi]. 416 p. Praha (in Czech).Lange: Lange J. E. (19351940): Flora agaricina danica.

Vol. 15. Copenhagen.Ludwig: Ludwig E. (2000): Pilzkompendium. Band 1. Ab-bildungen. 192 p., Eching.

Moser et Jlich: Moser M. et Jlich W. (19852002): Farb-atlas der Basidiomyceten. Lieferung 120. Stuttgart.

Phillips: Phillips R. (1981): Mushrooms and other fungi ofGreat Britain & Europe. 288 p., London.

Ryman et Holmsen: Ryman S. et Holmsen I. (1992):Pilze. 718 p., Braunschweig (German edition ofSvampar en flthandbok).

Abbreviations used in the text

(only those which are not included in the standard works

listed above)

CR: Czech Republic, E: length/width ratio of the spores,ICBN: International Code of Botanical Nomenclature(Greuter et al. 2000), L: number of lamellae reaching up tothe stipe, l: number of lamellulae between two lamellae,SEM: scanning electron microscope, Q: mean value of E(= length/width ratio of the spores) for all spores studied.

Abbreviations used in line drawings

B: basidia, BD: basidioles, CH: cheilocystidia, P: pleu-rocystidia, PC: pileocystidia, S: spores, TC: terminal cellsof hyphae on pileus cuticle, TCS: terminal cells of hyphaefrom stipe cuticle.

Results

Taxonomy, ecology and distribution of the genus

Gymnopilus in the Czech Republic

Gymnopilus P. Karst., Bidrag Knnedom FinlandsNatur Folk 32: xxi, 1879.

= Fulvidula Romagn., Rev. Mycol. 1: 209, 1936 (invalid name without Latin description, see Donk 1962: 101).

Type: Gymnopilus liquiritiae (Pers.) P. Karst. The historyof typification was described by Donk (1962: 117118). Hi-land (1990) wrote that this choice was mechanical, the name G.liquiritiae is hard to interpret and has been variously interpret-ed during the years. He proposed a new lectotype, Gymnopilus

picreus (Pers.: Fr.) P. Karst. I fully agree with his arguments. Itis also possible that G. liquiritiae is a synonym of G. picreus a sanctioned name which must be preferred. For a detailed dis-

cussion see under G. liquiritiae in the chapter Comments onsome taxa not reported from the Czech Republic.

Key to the species of the genus Gymnopilus based on

taxa known from the Czech Republic*

1. Fruitbodies with a distinct membranaceous annulus, robust,medium-sized to large . . . Gymnopilus spectabilis, p. 5

1.* Fruitbodies without membranaceous annulus, small tomedium-sized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Fruitbodies with traces of purple to violet colour . . . 32.* Fruitbodies without purple or violet colour (at most with

orange or red tinges) . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Spores large [8.09.5(11) 6.06.8(7.2)m

], pileustomentose-fibrillose to fibrillose-scaly . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . Gymnopilus igniculus, p. 8

3.* Spores smaller (68.5 46 m, average sporeswithout extremely large ones), pileus with more distinct,mostly erect scales covering the whole surface . . . . . .. . . . . . . . . . . . Gymnopilus purpuratus, G. luteifolius,. . . . . . . . . . . . . . . . . . . . . . . G. peliolepis, G. dilepis

(taxa not known from the Czech Republic,for differences, see discussion under G. igniculus)

4. Spores small, most of them measuring up to6.0 4.5m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4.* Spores on average larger than 6) 4 m . . . . . . . . . 7

5. Growing on decaying wood . . . . . . . . . . . . . . . . . . . . 6

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5.* Growing in grassy places in tufts of grasses, typically amongDactylis glomerata, spores mostly 5.06.0 3.54.5m

. . . . . . . . . . . . . . . . . . . . . . Gymnopilus flavus, p. 156. Pileus bright coloured (rusty orange to rusty brown with

orange tinge), surface slightly lustrous, lamellae bright(deep yellow, then yellow-brown to yellow-rusty),

spores prolonged (ellipsoid to amygdaliform-ellipsoid:length/width ratio mostly 1.41.9) with a distinctsuprahilar depression, mostly 4.56.0 3.03.5 m,cheilocystidia narrowly lageniform with more or lessprominent globose head which is not sharply dividedfrom the neck . . . . . . . . . . Gymnopilus bellulus, p. 11

6.* Pileus dull coloured (ochre-brown, dark brown to rustybrown), surface fibrillose-tomentose, lamellae dark(dark brown at maturity), spores subglobose but alsobroadly ellipsoid to broadly obovoid in side view(length/width ratio mostly 1.11.4), without suprahilardepression, mostly 4.56.0 3.54.5 m, cheilocys-

tidia very distinctive (tibiiform with a narrowly lageni-form or cylindrical basal part, long narrow neck anddistinct globose head, often with slightly thickened andrusty brown wall) . . . . . Gymnopilus josserandii, p. 13

7. Growing on peat, soil or burnt substrates (ash, charcoaletc.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7.* Growing on wood, often strongly decayed or hidden insoil, but never on burnt wood . . . . . . . . . . . . . . . . . 10

8. Spores large, mostly 811 57 m, stipe with darkred-brown lower part, habitat: peat-bogs or sand-duneheaths on peaty soil, growing on peat or peaty soil or di-rectly among Sphagnum and other mosses or amonglichens. . . . . . . . . . . . . . . . Gymnopilus fulgens, p. 17

8.* Spores smaller, mostly 69(10) 45 m, stipe with-out dark red-brown lower part, habitat and substrateother . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Pileus dirty yellow-brown, rusty brown to greyish brownwith fibrillose-tomentose to tomentose-scaly surface,spores longer: 79(10) 45 m, taste completelymild . . . . . . . . . . . . . . . . Gymnopilus decipiens, p. 18

9.* Pileus bright coloured: orange red-brown, surfacesmooth, at most finely fibrillose-scaly, spores shorter:(6)6.57.5(8.5) (3.5)4.04.8(5.5) m, taste bit-terish . . . . . . . . . . . . . . . . . . Gymnopilus odini, p. 38

(not documented from the Czech Republic;

other species growing on soil or burnt substratesbut hitherto not known from Central Europe:

Gymnopilus pseudofulgens, G. humicola;for differences see discussion under G. decipiens)

10. Spores large, mostly 8.510.5 5.56.5 m, coarselyornamented, with suprahilar disc, without suprahilar de-pression, fruitbodies small, rarely medium-sized (pileus555 mm, mostly up to 40 mm), pileus at centre typi-cally orange-brown, red-brown to reddish rusty brown(but also yellow-rusty to rusty brown when dry), lamel-lae vividly deep yellow for a long time, stipe typicallydark rusty brown to umber-brown with a red or violettinge . . . . . . . . . . . . . . . . . Gymnopilus picreus, p. 35

10.* Spores smaller, mostly 79 45 m, moderately or-namented, without suprahilar disc, with more or less dis-

tinct suprahilar depression, fruitbodies mostly medium-sized, pileus and stipe without red tinges and not verydark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Hyphae of upper layer of pileus cuticle narrow,310(12) m in diam., cylindrical, pileus surface prin-cipally smooth but rusty ochre to rusty brown fibrillose-

striped to appressed fibrillose-scaly, young pilei coveredwith white to greyish-white velum, pileus context paleyellow, young lamellae pale yellow, stipe covered withwhite tomentose-fibrillose velum remnants . . . . . . . . .. . . . . . . . . . . . . . . . . . . Gymnopilus penetrans, p. 19. . . . . . . . . . . . . (= G. sapineus sensu Hiland 1990)

11.* Hyphae of upper layer of pileus cuticle broad,(4)620 m in diam. (mostly 816 m), coarsely in-crusted, with narrowly clavate, clavate to pyriform ter-minal cells, pileus surface fibrillose-tomentose,tomentose to tomentose-scaly, without velum, pileuscontext mostly deep yellow, lamellae typically deep yel-low when young, stipe yellow fibrillose-tomentose . . .. . . . . . . . . . . . . . . . . . . . Gymnopilus sapineus, p. 24

sensu Khner et Romagnesi (1953), Ludwig(2000, 2001), Breitenbach et Krnzlin (2000) etc.;

non G. sapineus sensu Fries (1821), Hiland (1990).

Gymnopilus spectabilis (Weinm.: Fr.) A. H. Smith(Text-fig. 1; Pl. 3, figs 23; Pl. 4, figs 12; Pl. 9)

B a s . : Agaricus spectabilis Weinm., published in Fries, Elench.fung. 1: 28, 1828.

Agaricus spectabilis Weinm.: Fr., Elench. fung. 1: 28, 1828. Gymnopilus spectabilis (Weinm.: Fr.) A. H. Smith, Mush-rooms and their natural habitats: 471, 1949. Pholiota spectabilis (Weinm.: Fr.) P. Kumm., Fhr. Pilzk.:84, 1871. Fulvidula spectabilis (Weinm.: Fr.) Romagn., Bull. Soc.Mycol. France 48: 89, 1943 (1942), invalid combinationas the generic name Fulvidula is published invalidly.

= Agaricus quercicola Lasch, Linnaea 4: 545.= Pholiota aurantiaca Thesleff, Bidrag Knnedom Finlands Natur

Folk 79(1): 34, 1920 (for type study see Holec 2001a: 181).= Pholiota gigantea R. Naveau, Natuurwetenschappelijk Tijd-

schrift 5: 77, 1923 (for type study see Holec 2001a: 188).= ?Pholiota grandis Rea, British Basidiomycetae: 118, 1922 (for

taxonomic discussion see Holec 2001a: 189).= ?Agaricus (Pholiota) villosus Fr., Elench. fung. 1: 28, 1828.

?Pholiota villosa (Fr.) Sacc., Syll. fung. 5: 752, 1887 (fortaxonomic discussion see Holec 2001a: 206207).

Se l ec te d il lu st ra t io ns : Lange: fig. 108B (as Pho-liota spectabilis). Phillips: p. 144 (as G. junonius). Ry-man et Holmsen: p. 482 (as G. junonius). Dhncke: p.699. Moser et Jlich: III Gymnopilus 1, top figure. Bre-itenbach et Krnzlin: fig. 144 (as G. junonius). Ludwig:fig. 31.4. A, B, C (as G. junonius). Hagara et al.: p. 291,fig. 425. Bolets de Catalunya: vol. 3, fig. 119. Cetto: fig.919, fig. 2669.

C h a r a c t e r i s t i c s i n b r i e f : Fruitbodies singly or inclusters, large to very large, fleshy; pileus 50150(200) mm,dry, bright yellow, ochre-yellow, orange-yellow, ochre-rustyto rusty orange, surface distinctly innately tomentose-fibril-lose to tomentose-scaly, the covering being yellow, yellow-

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rusty to rusty brown; stipe with cylindrical upper part butwith fusiformly inflated to thickly bulbous lower part andfusiformly attenuated base, with prominent membranaceousannulus when young; taste distinctly bitter, spores large,mostly 8.510.5 5.56.8 m, ellipsoid, ovoid-ellipsoid toamygdaliform-ellipsoid, medium to coarsely verrucose toverrucose-rugulose. Growing as a saprophyte or parasite onwood of deciduous trees, rarely of conifers, in areas with awarm or temperate climate, never in mountains, common.

D e s c r i p t i o n : Fruitbodies growing singly or in fasci-cles. Pileus 30150(200) mm, fleshy, at first subglobose,then convex or broadly obtusely conical, sometimes plano-convex when old, margin slightly inflexed, without umbo,surface dry, mat, not translucently striate, not hygro-phanous, ground colour bright yellow, ochre-yellow (5B8),orange-yellow, ochre-rusty to rusty orange (6C8) at centre(depending on age and weather), towards the margin brightyellow (4A78) when fresh, the entire surface distinctly in-nately tomentose-fibrillose to tomentose-scaly, scalesdensely arranged, mostly appressed but slightly upraisedwhen old, this covering being yellow, yellow-rusty to rustybrown, pileus margin sometimes with overlapping, tomen-tose-membranaceous, bright yellow to yellow-ochre veil rem-nants. Lamellae crowded, L = 5080, l = 17, 412(15) mm

high, segmentiform or slightly ventricose, near the stipeemarginate and decurrent with a small tooth, rarely slightlydecurrent, at first pale yellow to sordid yellow, at maturityrusty yellow to yellow-brown with a rusty tinge, rustybrown spotted, edge even or finely irregularly serrulate, paleyellow. Stipe very variable in shape and size (depending on

sort of stipe insertion, age and size of the whole fruitbody orcluster), 50150(200) 1030(40) mm, upper part cylin-drical, in lower part either fusiformly inflated with conicalyattenuated or even rooting base or thick bulbous (up to 50mm, in extreme cases even up to 100 mm), mostly withfusiformly attenuated base, annulus (veil remnant) promi-nent, about 515 mm broad, upright, membranaceous,bright yellow to rusty yellow, partly missing at maturity andremaining only as a disrupted fibrillose-tomentose annularzone on stipe surface, stipe pale (lemon) yellow and smoothabove the annulus, below the annulus yellow, yellow-ochreto ochre, surface ochre rusty to rusty fibrillose-grooved,

spotted to marbled. Context thick, fleshy, in young fruit-bodies yellowish-white to pale yellow in pileus, but yellow-ochre below pileus cuticle and above the lamellae, in stipepale yellow to deep lemon yellow, in base brownish, in old-er fruitbodies darker, bright yellow to yellow-rusty, with5 % KOH immediately orange to orange-brown. Taste im-mediately distinctly bitter. Smell indistinct or aromaticallyfruity on section.

Spores (8.0)8.510.5(11.2) (5.2)5.56.8(7.2)m,E = 1.351.80, Q = 1.50, ellipsoid to amygdaliform-ellip-soid in side view, with suprahilar depression; ellipsoid,ovoid-ellipsoid to amygdaliform-ellipsoid in face view,rusty yellow in KOH, hilar appendix small but visible, or-

namentation very variable in appearance and degree of de-velopment, in fully developed spores medium to coarse,verrucose to verrucose-rugulose, up to 1 m high, in somespores poorly developed or fully absent, with small andsometimes poorly developed suprahilar disc, spore proto-plasm colouring vinaceous reddish brown in Melzersreagent (dextrinoid) but wall remaining bright yellow to yel-low-rusty. Basidia 2631 68 m, 4(2)-spored, narrowlyclavate or cylindrical with 12 slight constrictions. Basidio-lae 619 56 m, narrowly clavate. Cheilocystidia abun-dant at edge or mixed with basidiolae, 2137 59 m,variable in shape, mostly lageniform to narrowly lageniform

but also narrowly to moderately fusiform, utriform, clavate,mostly with a cylindrical or subcapitate to capitate upperpart, head 36(8) m, thin-walled, hyaline, sometimeswith a granular or homogeneous yellow content. Pleurocys-tidia absent. Lamellar trama regular, of parallel hyphae312 m broad, cells cylindrical, hyaline. Velum (from an-nulus) composed of parallel to slightly interwoven hyphae512 m broad, cells cylindrical, mostly hyaline, some ofthem with yellow-rusty granular content, wall only slightlyyellow-rusty incrusted. Pileus cuticle a cutis of densely ar-ranged hyphae 410 m broad, the whole layer yellow, cellsyellow-rusty incrusted, cylindrical, this layer covered with athin layer of ascending and interwoven velum hyphae (seeabove), in a scalp visible as a loose net or cords of velumhyphae forming the fibrillose to scaly pileus covering, cells

6

Text-fig. 1. Gymnopilus spectabilis: 1 Kneves (PRM892723); 2 dike between Nadje and Vra fish-ponds, JH

482/02 (PRM). For explanations see Material and Methods.Scale bar = 10 m.

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yellow-rusty incrusted, with slightly clavate terminal cells;pileocystidia absent. Stipe cuticle a cutis, yellow-rustycoloured, cells cylindrical, 26 m broad, terminal elementscapitate or clavate, at places covered with nests of ascend-ing velum hyphae (see above); true caulocystidia resem-bling cheilocystidia sometimes present but rare. Clamp

connections abundant in all tissues.F r u c t i fi c a t i o n : August October (CR).E c o l o g y : In the Czech Republic, G. spectabilis is

mostly found as a saprophyte on dead wood of broadleavedtrees, especially on stumps, fallen trunks and wood in soil(roots etc.). Less frequently it grows as a parasite on roots oron the base of living trees. Most finds are from Quercus spp.(Q. roburif the tree species was noted). Other known hostsfrom the CR:Malus domestica, Pyrus communis, Acer pseu-doplatanus, Populus alba, Salix caprea, Carpinus betulus,

Tilia, Betula, Ulmus, Corylus. Finds from conifers (Pinus)are known from Southern Bohemia (M. Beran, personal com-munication, collections in CB). I personally studied at leasttwo collections from conifers (Pinus: WU 7308, Pinus nigra:Fungi Exsiccati Suecici no. 2718). It well corresponds withthe data from North America (Hesler 1969) where the speciesgrows on wood of conifers and deciduous trees. Hiland(1990) mentions finds on Fagus from Norway. In southernEurope, the species is known from e.g.Erica arborea orEu-calyptus (Bon et Roux 2002). In the CR, G. spectabilis growsboth in natural forests as well as in man-influenced or man-made habitats (especially in parks, gardens, on fish-ponddikes, in avenues etc., along forest roads or forest paths,mostly on old trunks or stumps). It is mostly found in low-lands and occurs also in the hills up to altitude of about 500

m. It has never been found in mountains.D i s t r i b u t i o n : Gymnopilus spectabilis is rather com-

mon in suitable habitats of the Czech Republic (see Ecolo-gy). It is distributed in areas with a warm or temperateclimate (called thermophyticum and mesophyticum inthe CR, see Hejn et Slavk 1988). The species is commonin most European countries except for mountainous areas(which is well seen e.g. in the map by Krieglsteiner 1991:742). In Scandinavia, it only grows in southern regions (inthe temperate and hemiboreal zone).

D i s c u s s i o n : Gymnopilus spectabilis is well recog-nisable by its robust fruitbodies mostly growing in clusters,

tomentose-fibrillose to tomentose-scaly pileus covering,presence of a prominent annulus (when young), bitter tasteand large spores with coarse verrucose to verrucose-rugu-lose ornamentation. The fruitbodies are very variable insize, shape and colour which depends on their age, sort ofstipe insertion and weather conditions. Extremely largefruitbodies with a swollen stipe base (up to 100 mm) arefound in some cases.

In the literature, small and slender forms are mentioned,mostly under name Gymnopilus junonius (Fr.: Fr.) P.D. Or-ton. However, in the last four decades of the 20th century,the name G. junonius has been used both for slender and ro-bust forms (the forms were considered conspecific by mostauthors) instead of the name G. spectabilis. Both G. juno-nius and G. spectabilis are sanctioned names, but G. juno-

nius is older. If the two names really represent the samespecies, then G. junonius must be used as the correct name.However, even some recent authors (Bon et Roux 2002, Ro-bich 1989) report finds of a slender fungus named G. juno-nius or G. spectabilis var. junonius. I personally do notknow such a fungus (neither from nature nor from herbaria),

but until its relation to robust forms (G. spectabilis) ischecked, I prefer to use the traditional name G. spectabilisfor the robust forms which I know from the Czech Repub-lic. Another reason is the fact that some important parts ofFries description of G. junonius (Fries 1821: 244, 1874:223) are in contradiction to the collections I know from theCR. Fries writes that Agaricus junonius is small (minor:pileus up to 5 cm), has glabrous pileus surface, cylindricalstipe (stipite aequali) and grows solitarily. All these char-acters disagree with the robust fungus widely known as G.spectabilis and, in my opinion, the name G. junonius in itsoriginal sense is not applicable for it.

The nameAgaricus spectabilis was created by JohannesAnton Weinmann, probably in a letter sent to Fries togetherwith dried fruitbodies from the vicinity of Petropolin (= St.Petersburg) and published by Fries (1828) in Elenchus fun-gorum (where it is simultaneously sanctioned). This can bededuced from Fries abbreviations in litt. (= in correspon-dence, which means a description obtained from Weinmannused for part of his broader description of A. spectabilis)and v. s. (= vidi siccum: dried fruitbodies obtained fromWeinmann). Fries really ascribed the name to Weinmann,which is clear from Index alphabeticus (Fries 1832: 42)where it is cited asA. spectabilis Weinm.

According to Dennis et al. (1960: 70), the name was

published in Weinmann (1824). At the moment, I have nopossibility to check this as the book is not available fromany Czech library.

In later works, Fries (1838: 166, 1874: 221) cited thename as A. spectabilis Fr., Elench. fung. 1: 28, 1828 (nonWeinm.) which means that he taxonomically and nomen-claturally excluded Weinmann from his concept of A.spectabilis. He probably insisted that the Weinmanns partof the original description does not correspond to his ownpart based on fresh fruitbodies seen by him (see abbrevia-tion v. v. = vidi vivum in Elenchus). However, this has noinfluence on the fact that the basionym A. spectabilis was

created by Weinmann.Recently, the name Gymnopilus pampeanus (Speg.)Singer commonly used for collections from Southern Hemi-sphere was added to the synonymy of G. spectabilis by Reeset Strid (2001).

C o l l e c t i o n s s t u d i e d :Austria Burgenland, Oberpullendorf, Pinus, on stump, 17 Aug

1988, leg. W. Klofac (WU 7308). Niedersterreich, Maissau,Grnhof, 23 Sep 1998, leg. ? (illegible) (WU 18398). Niedersterreich, W of Pulkau, Pulkautal, on stump (Carpinusforest), 21 Oct 1979, leg. G. and M. A. Fischer (WU 0340). Niedersterreich, Wien, Galitzinberg, on stump near the

ground (Fagus

?,Quercus

?), 23 Sep 1984, leg. Reisinger (WU3556). Niedersterreich, Wien, Lainzer Tiergarten, Fagus, onstump, 18 July 1982, leg. W. Zhrer (WU 2345).

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Bulgaria Burgas, between Lovno chane and Slanev Brig, onsoil on forest path, 20 Aug 1982, leg. F. Kotlaba (PRM830098). Ropotamo, Arkutino reserve, Quercus, at base andon roots, 11 June 1978, leg. J. Kuthan (PRM 824635).

Czech Republic Bohemia, esk Brod, Tilia, on trunk, Sep 1936,leg. J. Skora (PRM 28583). esk vcarsko National Park,Mezn Louka near Hensko, Koz hbety, Quercus robur, roots

at base of a living trunk, 28 Sep 2002, leg. J. Holec, JH 305/02(PRM 896824). Horn Cerekev, 2 km NE (direction of NovRychnov), Tilia, 14 Oct 1979, leg. F. ejnost (CB 2221). Hradec Krlov, 5 km SE of the town, Fagus sylvatica, 22 Sep1985, leg. J. Valter (CB 6345). Praha, Seminsk zahradapublic garden, Pyrus, 30 Oct 1961, leg. Chromcov (PRM616093). Praha-Stromovka, Quercus, on strongly decayedtrunk, 27 Aug 1950, leg. Z. Pouzar (PRM 729213). Rakovnk,Kneves, on wood in soil, 25 Sep 1998, leg. V. Bazika (PRM892723). Southern Bohemia, 1 km NW of Rakovice, Tilia, atbase, leg. M. Koch (CB 5386). Southern Bohemia, 1 km Wof Chnov, Populus, on stump, 6 Oct 1985, leg. J. Valter (CB6346). Southern Bohemia, Frahel near Tebo, dike betweenNadje and Vra fish-ponds, Quercus, on stump at soil level, 11Oct 2002, leg. J. Holec, JH 482/02 (PRM). Hlubok nad Vl-tavou, park in front of Ohrada castle, Quercus, on stump, 11Oct 2002, leg. J. Holec, JH 479/02 (PRM). Hlubok nad Vl-tavou, bank of Munick rybnk fish-pond, Populus alba, onstump, 11 Oct 2002, leg. J. Holec, JH 480/02 (PRM 900671). esk Budjovice, embankment by the Vltava river, Tilia, onstump, 25 Sep 1984, leg. M. Janouek (CB 3921). SouthernBohemia, Klec, dike of Dobr Vle fish-pond,Alnus glutinosa,on stump, 16 Sep 1986, leg. T. Papouek (CB 4623). South-ern Bohemia, Klec, dike of the Lska fish-pond, Corylus avel-lana, at base, 9 Oct 1980, leg. J. Va (CB 2499). SouthernBohemia, Klec, dike of the Nadje fish-pond, Quercus, onstump, 21 Aug 1983, leg. J. Novotn (CB 3692). Southern

Bohemia, Maleice, on dike of a fish-pond, in grass underQuercus, 15 Oct 1989, leg. Z. Vrzk (CB 5671). SouthernBohemia, Tebo,Betula, at base, 29 Aug 1958, leg. R. Vesel(PRM 519210). Southern Bohemia, Turovec, near Lun fish-pond, Pinus sylvestris, on stump, 16 Sep 1988, leg. M. Beran(CB 5929). Vesel nad Lunic, Boilec, near Hlin fish pond,Quercus, on stump, Sep 1979, leg. Chalupsk (PRM 889214). Brno-idenice, Malus domestica, old stump, 28 Sep 1964,leg. K. Koncerov (BRNM 312592). Moravia, Beclav,Lanhot, Quercus, 12 Sep 1967, leg. M. Svrek (PRM 629482). Beclav, Lanhot, Cahnov floodplain forest, Quercus robur,at base of a dead trunk, 10 Oct 1965, leg. F. Kotlaba and J.Lazebnek (PRM 617465). Ditto, Ulmus, on fallen trunk, 6Oct 1967, leg. J. Lazebnek and A. Vgner (BRNM 312591).

Moravia, Lanhot, Mysliveck palouk, Quercus, on stump,22 Oct 1998, leg. V. Antonn (BRNM 642470). Beclav,Lanhot, Ranpurk virgin (floodplain) forest, Quercus robur,on decayed stump, 14 Aug 1979, leg. L. Kubikov (PRM821870). Moravia, Heroltice, coniferous tree, on stump, 22Sep 1962, leg. K. Koncerov (BRNM 312393). Moravia, Le-tovice,Malus domestica, on injured trunk, 12 Sep 1971, leg. V.Beneov (BRNM 301809). Moravia, Vlkov, Ondruky,coniferous forest (Picea, Pinus), 17 Sep 1974, leg. H. Hurtov(BRNM 289660). Znojmo, Boice, Hoja, Carpinus betulus,on decayed stump, 29 Aug 1971, leg. Z. Pouzar and F. Kotlaba(PRM 796550). Nov Bydov, Kobylice, on stump of abroadleaved tree, 6 Oct 1989, leg. L. Drahokoupil (PRM873975). Ostrava-Tebovice, Turkov nature monument,

Quercus, on soil among roots, 1 Oct 2002, leg. H. Deckerov(PRM 900663). Ostrava, Polanka, Pemyovsk mokad na-

ture reserve, Salix caprea, on roots of a fallen trunk, 4 Oct2002, leg. J. Holec, JH 336/02 (PRM 900664). Czech Silesia,Javornk, Ra dol valley,Malus?, on trunk, 12 Sep 1945, leg.V. Pospil (BRNM 312569).

Romania Cluj, Hoia-Cluj, Quercus, 28 Oct 1956, leg. G. Silaghi(PRM 533794).

Sweden Gteborg, Hisingen, Rya skog (Fungi Exsiccati Suecici

no. 2718), Pinus nigra, on stump, 9 Sep 1952, leg. F. Karlvall(PRM 576613). Gteborg, Slottsskogen (Fungi ExsiccatiSuecici no. 2022), on stumps of Quercus and Corylus, Oct1951, leg. F. Karlvall (PRM 729196).

Gymnopilus igniculus

Deneyer, P.-A. Moreau et Wuilbaut(Text-fig. 2)

Gymnopilus igniculus Deneyer, P.-A. Moreau et Wuilbaut, Doc.Mycol., vol. 32, no. 125: 11, 2002 (the species was at firstpublished without Latin description and type designation inBon et Roux 2002: p. 4, p. 1516)

Se l ec te d il l us tr at i on s: Bon et Roux (2002: pl. 1-B), Wuilbaut (2002: p. 31), Holec et al. (2003: 165166).

Ch ar ac t er is ti cs i n br ie f (based on finds from theCzech Republic): Fruitbodies small to medium-sized, pileus1540 mm, pileus covering tomentose-fibrillose when youngand fibrillose-squamulose to distinctly scaly at maturity, pur-plish to vinaceous or reddish brown on yellow ground, veilwhitish membranaceous to fibrillose, ring lacking, stipe dis-tinctly longitudinally purplish brown fibrillose on dirty whiteor slightly violaceous ground, context yellowish with reddishviolaceous tinge, smell fungoid, spores relatively large,8.09.5(11) 6.06.8(7.2) m, broadly ellipsoid, with

rough verrucose to verrucose-rugulose ornamentation, shapeof cheilocystidia variable, pleurocystidia absent. Growing ondecaying wood on burning coal mine dumps.

D e s c r i p t i o n (based on fruitbodies from the CzechRepublic which were thoroughly described by Holec et al.2003): Basidiocarps single or in small groups, never cespi-tose. Pileus (7)1540 mm broad, broadly conical to con-vex, involute at margin when young, then convex withapplanate centre and inflexed margin, almost applanate withslightly inflexed margin in the end, entirely distinctly to-mentose when young, then except for centre adpressed ra-dially fibrillose, almost pyramidal fibrillose-squamulose atcentre and radially adpressed fibrillose towards marginwhen old, vesture purplish or vinaceous brown (911E67,10F7), with paler margin (9E7) when young, then vinaceousonly at centre and paler, purplish ochraceous brown(89D78) towards margin, when old reddish brown(9DE7) at centre and paler (89D6) towards margin,ground yellowish to pale dirty yellow; margin sometimesdecorated with almost membranaceous velar remnants.Lamellae rather distant, L = 1824, l = 23, broadly adnateor emarginate and shortly decurrent with tooth, horizon-tal, light yellow (3A4) to orange-yellow (45A5), with con-colorous, irregularly serrulate, finely pubescent edge. Stipe2080 26 mm, cylindrical, sometimes slightly clavate

(up to 7 mm) or attenuated towards base, often curved, dis-tinctly longitudinally purplish brown (10D56, 1011E67)

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fibrillose or fibrillose-squamulose on dirty white or slightlyviolaceous ground, whitish or with violaceous tinge, lessdistinctly fibrillose (paler than other parts of stipe) andsometimes striate (decurrent lamellar tooth) at apex; withdirty whitish basal mycelium. Velum membranaceous, dirtywhitish or pale dirty brownish, at margin yellowish when

young, its remnants sometimes distinct at pileus margin andnear stipe apex (when young) and only as indistinct fibrilson stipe surface (when old). Context hollow in stipe, whitishto yellowish, in stipe apex and above lamellae more dis-tinctly yellow, pale violaceous under pileipellis, slightly vi-olaceous (reddish) in stipe middle; with fungoid orindistinct smell and bitterish taste.

Spores 8.09.5(11) 6.06.8(7.2) m , E =1.251 .50, Q = 1.39, broadly ellipsoid, with small but dis-tinct suprahilar depression visible in side view, rusty yellowin KOH with darker, rusty brown wall which is slightlythick-walled, surface densely covered with low but large (up

to 1.5m

broad) and irregular verrucose to rugulose-verru-cose ornamentation, hilar appendix tiny, spore protoplasmdistinctly dextrinoid (staining vinaceous reddish brown) inMelzers reagent with the wall remaining rusty yellow. Ba-sidia 2428 89 m, mostly 4-spored, rarely 2- or 1-spored, clavate to subutriform, sometimes with a slightmedial constriction, clamped. Basidioles 1030 39 m,cylindrical when young, then distinctly clavate, some ofthem filled with a homogeneous yellow-rusty pigment (in5 % KOH), clamped. Cheilocystidia 2435 610 m, ar-ranged in nests on edge or mixed with basidioles, nar-rowly clavate to fusiform-lageniform when young, thentypically lageniform to subutriform with 34 m broad

neck and more or less distinct globose head 58 m in di-ameter, thin-walled, hyaline, clamped. Pleurocystidia notobserved. Lamellar trama regular to subregular, consistingof parallel hyphae 320 m broad, narrower hyphae locatednear the subhymenium, cells cylindrical, slightly fusiformto narrowly ellipsoid, with hyaline content and yellowishwall, non-dextrinoid, clamped, subhymenium of densely ar-ranged hyphae. Pileus cuticle a cutis, 6070 m thick,2layered, upper layer thin, dark reddish to violet brown inKOH, consisting of densely and radially arranged (parallelin a section) hyphae 420 m broad, cells cylindrical to nar-rowly fusiform or narrowly ellipsoid, with distinct violet to

reddish brown incrustations arranged in a tiger pattern, ter-minal cells indistinct, subfusoid to narrowly clavate, lowerlayer thick, less coloured, yellow in KOH, of loosely ar-ranged parallel to slightly interwoven hyphae with less dis-tinct incrustations, this layer gradually passing into thepileus context made up of cylindrical, narrowly fusiform tonarrowly ellipsoid hyphae up to 25 m broad, hyaline orpale brownish in KOH. When a pileus scalp is observed, thepileus surface is covered with fascicles or a sparse net ofcells forming the upper layer of the pileus cuticle. Stipe cu-ticle a cutis of densely arranged, parallel, cylindrical hyphae38 m broad, yellow-brown with violet tinge, cells withyellow-brown to rusty brown incrustations, slightly thick-walled, clamped, terminal cells indistinct, cylindrical;

caulocystidia not observed, but a sparse net of interwoven,yellow-brown incrusted veil hyphae 28 m broad coveringthe cuticle. Stipe context made up of cylindrical to subellip-soid, slightly thick-walled, yellowish hyphae up to 15 m

broad, mixed with 510 m broad branched hyphae withyellow content in KOH. Clamp connections present in alltissues. Fragments of lamellae exuding a bright yellow pig-ment when mounted in 5% KOH.

Fructification: AprilMay (CR), February, Octo-ber, December (Belgium: see Deneyer et al. 2002).

E c o l o g y (for detailed data see Holec et al. 2003): In theCzech Republic, Gymnopilus igniculus was found on a burn-ing coal mine dump in the city of Ostrava where it grew ondecaying wood of Fraxinus. The dump is composed of siltyshales, claystones, siltstones and fossil soil with Stigmaria; toa lesser extent of fine-grained sandstones. At present, most

parts of the dump are reclaimed and planted mainly with Be-tula stands (3040 years), mixed stands (Betula, Tilia, Popu-lus, Quercus, Fagus) and somewhere also with stands ofPinus nigra or Quercus robur. In some places, heat andgasses escape from lower parts of the dump containing coalwith a relatively high content of sulphur. Consequently, thesoil of the collecting site was rather warm. During collectingdays, its surface reached a temperature of about 45 C.Although the site was insolated, it was moist due to the es-caping humid heat and gasses. The surface was overgrown bythe mossAulacomnium palustre, the grass Setaria pumila anda tree stand of Fraxinus excelsior. In Belgium, the speciesgrows in the same artificial habitat burning coal dump (40C on surface, see Bon et Roux 2002: 16).

9

Text-fig. 2. Gymnopilus igniculus: 1 Ostrava-Radvanice, 24April 1999 (BRNM 686264); 2 Ostrava-Radvanice, 1 May1999 (PRM 900986). For explanations see Material and Meth-ods. Scale bar = 10 m.

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Di st ri bu ti on : So far, Gymnopilus igniculus is knownonly from Belgium, where it was described from (Deneyeret al. 2002), the Czech Republic (Holec et al. 2003) andprobably also from France (see Discussion).

Discussion: The taxonomy, ecology and distributionof Gymnopilus igniculus was thoroughly discussed by

Holec et al. (2003). A shortened discussion is presentedhere.

The most important characters of fruitbodies of G. ig-niculus from the Czech Republic are given in the paragraphCharacteristics in brief. The fruitbodies agree in the most es-sential characters with those of Gymnopilus igniculus de-scribed from Belgium (Deneyer et al. 2002). However, thereare also differences. The fruitbodies from Belgium are slen-der and possess small purplish scales on a yellow ground, andare not completely purple-fibrillose like the fruitbodies fromOstrava. At first view, the fruitbodies from the two countriesare different. However, when the descriptions are compared,the differences with the original G. igniculus are only quanti-tative: the Belgian fruitbodies have a pileus cuticle thin andsoon differentiated into scales, with very few fibrils; thosefrom the Czech Republic have a thicker pileus cuticle whichdissociates less into scales or only with age. When the youngfruitbodies are compared, they are almost identical. Possibly,Deneyer et al. (2002) described in fact local populations withsmaller and slender fruitbodies with an underdevelopedpileus cuticle, whilst the material from the Czech Republicrepresents robust and more coloured fruitbodies.

In addition, such more robust fruitbodies were alsofound in France (burning dump Pinchonvalles, Avion, Pas-de-Calais) by J. Vast and R. Courtecuisse (see note by R.

Courtecuisse at the end of the paper by Deneyer et al. 2002:p. 16). In this case, the pileus surface was also fibrillose butlater divided (broken) into appressed scales (we saw a pho-tograph kindly provided by R. Courtecuisse). Maybe thisaspect was caused by insolation, as the fruitbodies were col-lected in June.

The fruitbodies from the Czech Republic and France al-so differ from the Belgian ones by their smell and taste. Thesmell of the first ones was not very distinctive while aprominent farinaceous-herbaceous smell and taste was not-ed in the Belgian material. External conditions (temperatureor drought?) may be responsible for this discrepancy, simi-

larly as in the previous case.A trophic difference can also be pointed out between theoriginal localities and French and Czech sites: all collec-tions from Belgium, small and slender, grow between moss-es (Campylopus), the mycelium growing from a layer ofmoss litter. The robust fruitbodies collected by Courtecuisseand Graca are associated with wood remnants, perhaps amore favourable substrate for their development.

Gymnopilus igniculus obviously has a greater variabili-ty of macrocharacters than was observed in the original col-lecting sites in Belgium. The more robust forms from theCzech Republic and France having a fibrillose pileus cover-ing which later separates into scales may be somewhat dif-ferent due to fructification in spring (the Belgian fruitbodieswere collected from October to February). In this period the

insolation is higher and air humidity lower which may causea different development of the pileus cuticle.

Gymnopilus species with purplish, violaceous or vina-ceous tinged fruitbodies are rare in Europe. They mostlyrepresent species imported from the tropics or subtropics. Itis e.g. Gymnopilus purpuratus (Cooke et Massee) Singer,

described from tree fern stems in the Royal Botanic Gar-dens, Kew (Cooke et Massee, Grevillea 18: 73, 1890; Cooke1883: 375; coloured picture: Cooke 18811891: plate 964).Fruitbodies identified as G. purpuratus were also found inthe greenhouse of the Botanical Garden in Zrich (Breiten-bach and Krnzlin 2000: 140). The name G. purpuratus wasfurther used for collections from compost heaps of woodand bark remnants in the Ribnitz-Damgarten district in Ger-many (Kreisel and Lindequist 1988, Ludwig 2001: 154,coloured picture by Ludwig 2000: p. 45). Rllin (1998)published finds identified as Gymnopilus cf.peliolepis fromthe base of a palm tree in an office in Genve, Switzerland.A find of Gymnopilus dilepis (Berk. et Broome) Singerfrom a pot with Philodendron purchased from a supermar-ket in Great Britain was published by Watling (1998); amore recent, abundant find, on a heap of woodchips, hasbeen illustrated by T. Leech in Henrici (2002: back cover).Finally, Bon and Roux (2002) used the name Gymnopilusluteifolius (Peck) Singer for G. purpuratus s. Breitenbach etKrnzlin (2000) and the name Gymnopilus peliolepis(Speg.) Singer for G. purpuratus s. Ludwig (2000, 2001).

These finds of Gymnopilus having purplish or violetcolours differ from Gymnopilus igniculus in smaller sporesmostly measuring 68.5 46 m (average spores withoutextremely large ones which are often present in Gymnopilus)

and in more distinct, mostly erect scales covering the wholepileus surface. Moreover, Gymnopilus purpuratus s. Kreiseland Lindequist (1988) differs in the presence of abundantpleurocystidia and in blue to blue-greenish colour changes onthe stipe surface and in the context.

The discussion on violet-coloured species of Gymno-pilus in Europe (Holec et al. 2003) clearly showed how poorour knowledge of this group in Europe is and how difficultit is to identify the finds. The reasons are the rarity of suchfinds, evident tropical or subtropical origin of collectionsfrom indoors or greenhouses and the difficulty to judge thevariability of European records with respect to species de-

scribed from other continents.Quite recently, this group of Gymnopilus wasthouroughly revised by Rees et al. (2004), also using molec-ular data from ribosomal ITS sequences. The authors usedavailable collections of most taxa discussed above (and oth-ers) from Europe, Sri Lanka, Australia, South and NorthAmerica. They proved the existence of two separate butclosely related species: Gymnopilus purpuratus andGymnopilus dilepis, which include most of the collectionsreported above; moreover, Gymnopilus luteofolius andGymnopilus megasporus Grgurinovic also were recognisedas good species separated from G. purpuratus (but closelyrelated). However, no collections of G. igniculus were stud-ied and its relation to other purple to vinaceous-colouredspecies remains unresolved.

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C o l l e c t i o n s s t u d i e d :Belgium: Cuesmes, "Terril du Levant", among Campylopus

retroflexus on hot mineral ground (charcoal), 9 Dec. 2001, leg.Y. Deneyer, P.-A. Moreau, J. Nuytinck and J. J. Wuilbaut(herb. PAM 01120901: fruitbodies not formally designated asisotype, but originating from the 30 original fruitbodies fromwhich the holotype deposited in BR was selected; all these

specimens were collected at the same site (about 20 m2). Formore details see Holec et al. (2003).

Czech Republic: Silesia region, Ostrava-Radvanice, burning coalmine dump, on decaying wood of Fraxinus, 24 April 1999, leg.M. Graca (BRNM 686264); ibid., 1 May 1999 (PRM 900986).

Gymnopilus bellulus (Peck) Murrill(Text-fig. 3; Pl. 2; Pl. 3, fig. 1; Pl. 10)

Bas.: Agaricus bellulus Peck, Bull. Buffalo Soc. Nat. Sci. 1: 51,1873.

Gymnopilus bellulus (Peck) Murrill, North American Flora,vol. 10, part 3: 200, 1917.

Naucoria bellula (Peck) Sacc., Syll. fung. 5: 841, 1887. Flammula bellula (Peck) Pilt, Kl k urovn naichhub hibovitch a bedlovitch (Agaricalium europaeorumclavis dichotomica): 351, 1952 (1951).

= Gymnopilus microsporus (Singer) ex Singer, Lilloa 22: 561,1951 (1949).

Selected illustrations: Breitenbach et Krnzlin,vol. 5: fig. 141. Moser et Jlich: III Gymnopilus 2, bottomfigure. Ludwig: fig. 31.6. Bon et Roux: pl. 3-A.

Charac te r i s t i c s in b r ie f : F ru i tbod ies sma l l ,pileus 525 mm, hemisphaerical, conical-convex to convex,surface not glossy (mat) or slightly lustrous, smooth to fine-ly rugulose, but under lens distinctly tomentose-rugulose to

finely rugulose-rugged, rusty orange to rusty brown with or-ange tinge, margin paler, yellow to yellow-ochre, lamellaedeep yellow, then yellow-brown to yellow-rusty, stipe1035 13 mm, yellow-rusty in the upper part, towardsthe base yellow-brown, ochre-brown to rusty brown, surfacepale yellow, yellow to yellow-rusty fibrillose-tomentose,taste bitter, spores small, mostly 4.56.0 3.03.5 m, E =(1.3)1.41.9(2.0), Q = 1.63, ellipsoid to amygdaliform-ellipsoid with a distinct suprahilar depression, medium tocoarsely verrucose, cheilocystidia narrowly lageniform withmore or less prominent globose head which is not sharplyseparated from the neck, hyphae on pileus and stipe surface

with clavate terminal elements up to 14 m broad. Growingon strongly decayed wood (especially trunks) of conifers innatural or near-natural forests, preferably in the mountains.

D e s c r i p t i o n : Fruitbodies growing individually or insmall groups, never caespitose. Pileus 525 mm, hemis-phaerical to conical-hemisphaerical with inflexed marginwhen young, then campanulate-convex, conical-convex toconvex, sometimes with a low broad umbo, margin some-what overlapping lamellae, at most slightly hygrophanousbut mostly dry, not translucently striate, surface not glossyor only slightly lustrous, smooth to finely rugulose but un-der lens distinctly tomentose-rugulose to finely rugulose-rugged, not scaly, colour rusty orange to rusty brown (6D8),

mostly with orange tinge, margin paler, yellow, yellow-rusty or yellow-ochre. Lamellae crowded, L = 2230, l =

13, 24 mm high, more or less ventricose, near stipeemarginate and decurrent with a small tooth, at first paleyellow, then deep yellow, at maturity yellow-brown to yel-low-rusty, edge concolorous. Stipe 1035 13 mm, basesometimes slightly bulbous, cylindrical, often curved, atfirst with pale yellow cortinoid traces of velum (towardspileus margin), soon disappearing, ground colour yellow-rusty in upper part, towards the base yellow-brown, ochre-brown to rusty brown, surface pale yellow, yellow toyellow-rusty fibrillose-tomentose, base sometimes whitish

tomentose. Odour indistinct. Taste distinctly bitter.Spores small, 4.56.0(6.5)3.03.5(4.0) m, E =(1.3)1.41.9(2.0), Q = 1.63, yellow to rusty yellow in KOH,ellipsoid, amygdaliform-ellipsoid to ovoid ellipsoid in frontview, ellipsoid to amygdaliform-ellipsoid with distinctsuprahilar depression in side view, wall rusty brown, mediumto coarsely verrucose, without suprahilar disc, slightly dextri-noid (with pale reddish brown tinge in Melzers reagent), ma-ture spores acyanophilous or only slightly cyanophilous,immature ones or those with a broken wall distinctlycyanophilous. Basidia 4(2)-spored, 1624 56m, cylin-drical to narrowly clavate with median constriction. Basidioles1618 5 m, narrowly clavate. Cheilocystidia numerous,rarely intermixed with basidiolae and basidia, small,(16)2026 46 m, narrowly lageniform with inflated

11

Text-fig. 3. Gymnopilus bellulus: 1 Srn, Dra skly protect-ed area, JH 534/01 (PRM); 2 esk leby, Splenit moun-tain, JH 286/98 (PRM). For explanations see Material andMethods. Scale bar = 10 m.

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basal part, long cylindrical neck (1.52.5 m) and more or lessprominent globose head (23.5(5) m) but head not sharplydivided from the neck, sometimes also without head, thin-walled, hyaline. Pleurocystidia absent. Lamellar trama regu-lar, of densely arranged hyphae 416 m broad, cellscylindrical or slightly fusiform, with yellow membranal pig-

ment, trama completely pale yellow, subhymenium thin, ofdensely arranged hyphae. Pileus cuticle (section) a cutis ofdensely arranged parallel hyphae 38 m broad, cells cylin-drical or slightly inflated, with rusty brown incrustations, en-tire layer yellow- to rusty brown, in scalp covered with a loosenet of interwoven cylindrical hyphae (veil hyphae?) 313 mbroad with distinctly clavate terminal elements up to 14 mbroad, rusty brown incrusted (in a tiger pattern), pileocys-tidia resembling cheilocystidia not observed. Stipe cuticle acutis of densely arranged cylindrical hyphae 38 m broad,cells rusty brown incrusted, terminal elements sometimesslightly capitate, this layer covered with loosely arranged pro-truding and curved hyphae (veil hyphae?), locally formingnests of interwoven hyphae 36 m broad, rusty brown in-crusted, with distinctly clavate to sphaeropedunculate terminalelements up to 12 m broad, caulocystidia rare, resemblingcheilocystidia, lageniform or cylindrical with capitate head.Clamp connections present in all tissues. Fragments of lamel-lae exuding yellow pigment when mounted in KOH.

Fr u ct if ic at i on : July October.Ecology: Based on data from the Czech Republic,

Gymnopilus bellulus is a saprophyte growing on thick fall-en trunks ofAbies alba and Picea abies. The trunks are inlater stages of decay characterised by the absence of bark,soft wood and mostly by the presence of moss covering. Ex-

cept for conifers, there are also rare finds from wood of Fa-gus. All my collections from the Czech Republic as well asthe herbarium specimens I have studied from this area orig-inate from natural to near-natural forests mostly designatedas protected areas. A first type are so-called mixed moun-tainous forests composed of Fagus, Picea andAbies (typi-cal example: the Boubnsk prales and ofnsk pralesvirgin forests) with admixture of Acer pseudoplatanus.Another habitat of G. bellulus is represented by naturalmountainous Picea abies forests (climax spruce forests). Inboth habitats, G. bellulus grows only at sites with a richpresence of fallen trunks of old thick trees. This means that

the species is lacking in cultural forests where all fallentrunks are removed. Concerning altitude, G. bellulus growsin the montane to supramontane zone (7301340 m a.s.l.,see collections studied).

I have also studied collections from Slovakia, Austria,Ukraine and Italy. In all cases, the data on ecology weresimilar (growth in natural to near-natural mountainousforests on wood of Picea or Abies). Also occurrence on

Larix and Pinus cembra are reported by Breitenbach etKrnzlin (2000) and on Taxus by Orton (1993). Preferenceof G. bellulus for mountainous areas is also confirmed bya map by Krieglsteiner (1991) from Germany which clearlyshows that the species grows there in the mountains only.On the other hand, Orton (1993) reports finds from GreatBritain which are from lowlands. In accordance with my da-

ta, Hiland (1990) also writes that G. bellulus is confinedto mature or even virgin forest types (to damp shady Piceaforests in Norway).

Distribution: In the Czech Republic, Gymnopilusbellulus is documented from several mountain ranges (u-mava, Novohradsk hory, Krkonoe, Beskydy). Generally, it

is a rare species which is, however, typical and scattered inappropriate locations (see Ecology). Its records in suitablehabitats of some other mountains are to be expected (e.g. Je-senky, Krlick Snnk, etc.). The species is also knownfrom the Carpathians in Slovakia and Ukraine (see kubla2003 and the collections studied). In Europe, it is furtherknown from e.g. Italy, Switzerland (Breitenbach et Krnzlin2000), Germany (Krieglsteiner 1991), Austria (Keller etMoser 2001) and France (e.g. Josserand 1948, Bon et Roux2002), mostly from the Alps, but also from the Massif Cen-tral, Jura, Schwarzwald, Schwbische and Frankische Alb.Except for the mountains, it is rarely found in lowlands (e.g.France, Great Britain, see Bon et Roux 2002, Orton 1993).In Scandinavia, G. bellulus is reported only from Norway(Hiland 1990). The species seems to be rare everywhere.

D i s c u s s i o n : Gymnopilus bellulus is well distinguish-able according to its small fruitbodies having relativelybright colours (pileus with orange tinge, lamellae at firstdeep yellow), bitter taste, small spores with distinct suprahi-lar depression, coarse ornamentation and length/width ratiomostly 1.41.9, small lageniform cheilocystidia with moreor less distinct head and other characters summarised in theparagraph Characteristics in brief. Gymnopilus josse-randii also has small fruitbodies and small spores but it dif-fers in the characters discussed under that species.

Some authors (e.g. Moreno et Esteve-Ravents 1990)are convinced that G. bellulus in the sense of European au-thors is different from the original American G. bellulus byPeck. They suggested to use the name Gymnopilus mi-crosporus (Singer) ex Singer for the European G. bellulus.The name G. microsporus is based on the Latin descriptionand illustration of Flammula liquiritiae (Pers.) P. Kumm.sensu Bresadola (Icon. mycol., vol. 16, text + tab. 783,1930). However, Flammula liquiritiae sensu Bresadola is afungus which is difficult to interpret.

To clear up all these discrepancies, I made a critical studyof Gymnopilus bellulus and G. microsporus, including the

type studies and detailed analysis of the original and recentconcepts of these species. The results will be published sepa-rately (probably in Mycotaxon in 2005 or 2006). The basicconclusions are that the American and European collections ofG. bellulus seem to be identical, and that G. microsporus typ-ified by the illustration by Bresadola is a hardly interpretablename which should be rejected. As Singers material of G. mi-crosporus is identical with G. bellulus, Gymnopilus mi-crosporus can be considered a synonym of G. bellulus.

C o l l e c t i o n s s t u d i e d :Austria Niedersterreich, Hohenberg, Lahnsattel, Abies, 30 June

1992, leg. A. Hausknecht (WU 10824). Niedersterreich,

Lunz/See, Rotwald: NSG Kleiner Urwald, wood of conifer(Abies?), 29 Aug 1996, leg. A. Hausknecht (WU 16262). Obersterreich, St. Konrad, Picea abies, on wood, 5 Oct 1984, leg.

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I. Krisai (herb. I. Krisai 3074). Steiermark, Pllau, Hllgraben,decayed conifer trunk (Picea?,Abies?), 12 Sep 2002, leg. J. Holec,JH 172/02 (PRM). Ditto,Abies alba, decayed trunk covered withmosses, 12 Sep 2002, leg. J. Holec, JH 171/02 (PRM).

Czech Republic Krkonoe Mts., pindlerv Mln, on stronglydecayed wood (Picea-Fagus forest), 8 Sep 1946, leg. J. Ku-bika (PRM 520916). umava Mts., Boubn mountain,

Boubnsk prales virgin forest, Aug 1936, leg. J. Herink (PRM27788). umava Mts., esk leby, Splenit mountain,Abies alba, on decayed trunk among mosses, 26 Oct 2002, leg.J. Holec (PRM 900962). Ditto, Abies alba , on decayingtrunk, 13 Oct 1997, leg. J. Holec, JH 763/97 (PRM 891944). Ditto, Abies alba , on decayed trunk, 26 Oct 2002, leg. J.Holec, JH 522/02 (PRM). Ditto, Picea abies, on decayingtrunk among mosses, 14 July 1998, leg. J. Holec, JH 286/98(PRM 897048). Ditto,Abies alba, on decayed trunk amongmosses, 3 Aug 1998, leg. J. Holec, JH 347/98 (PRM 897099). umava Mts., Lenora, Radvanovick hbet mountain ridge,Abies alba, on decaying trunk among mosses, 13 July 1998,leg. J. Holec, JH 261/98 (PRM 897028). umava Mts., NovPec, Plech mountain, Picea abies, on decayed trunk, 26 Aug1996, leg. J. Holec, JH 317/96 (PRM 889100). umava Mts.,Nov Pec, between Plech mountain and Trojmez, Piceaabies, on decayed trunk among mosses, 15 July 1998, leg. J.Holec, JH 299/98 (PRM 897058). umava Mts., Prily,danidla mountain, Picea abies, on decaying trunk amongmosses, 9 July 1998, leg. J. Holec, JH 211/98 (PRM 896984). umava Mts., Srn, Dra skly protected area, Abies alba,on decaying trunk, 29 Sep 2001, leg. J. Holec, JH 534/01(PRM). Ditto,Abies alba, on decaying trunk, 10 Oct 2002,leg. J. Holec, JH 461/02 (PRM). umava Mts., Zto nearLenora, Boubnsk prales virgin forest, Abies alba, on decay-ing trunk, 2 Oct 2001, leg. J. Holec, JH 589/01 (PRM). u-mava Mts., elezn Ruda, Debrnk protected area, on decaying

trunk of a conifer (Picea?,Abies?), 8 July 1998, leg. J. Holec,JH 181/98 (PRM 896964). Novohradsk hory Mts., ofnskprales virgin forest, Fagus, on wood, 29 Sep 1969, leg. J. Ku-bika (PRM 830924). Ditto, Fagus, on trunk, 21 Sep 1991,leg. V. Antonn (BRNM 553290). Beskydy Mts., Morvka,Travn, Picea abies, on decayed trunk, 17 Sep 1987, leg. Z.Pouzar (PRM 852262). Moravskoslezsk Beskydy Mts., dis-trict Frdek-Mstek, Bl, Salajka nature reserve, Abies alba,on dead trunk, 15 Aug 1970, leg. J. Kuthan (BRA). Mo-ravskoslezsk Beskydy Mts., Velk Karlovice-Leskov, Razu-la nature reserve,Abies alba, on decayed trunk, 28 Aug 1972,leg. J. Kuthan (BRA).

Italy Trento, Paneveggio, wood of a conifer, 4 Oct 1989, leg. R.Schtz (WU 8143). Trento, Passo di Manghen, on twigs, 11

Sep 1987, leg. I. Krisai (herb. I. Krisai 4332).Ukraine Eastern Carpathians, near Dilove (Trebuany), Ber-

lebash stream valley (Berleba), Picea abies, Aug 1937, leg. A.Pilt (PRM 488229). Eastern Carpathians, near Dilove (Tre-buany), Biliyi stream valley (Bl potok), Picea abies, Aug1935, leg. A. Pilt (PRM 20492). Eastern Carpathians, nearDilove (Trebuany), Biliyi stream valley (Bl potok), Piceaabies, Aug 1935, leg. A. Pilt (PRM 20232).

Gymnopilus josserandii Antonn(Text-fig. 4, Pl. 1, Pl. 11)

Gymnopilus josserandii Antonn, Fungi non delineati 11: 13, 2000.=

Naucoria subsphaerospora Joss., Bull. Soc. Mycol. France 64:21, 1948 (invalid name: published without Latin diagnosis). Gymnopilus subsphaerosporus (Joss.) Khner et Ro-

magn., Fl. anal. champ. supr.: 323, 1953 (invalid combina-tion: based on invalidly published basionym).

Se l ec te d il lu st ra t io ns : Moser et Jlich: III Gym-nopilus 4, top figure (as G. subsphaerosporus). Antonn etkubla, Fungi non delineati 11: photo no. 5, 2000. Breit-enbach et Krnzlin, vol. 5: fig. 150 (as G. subbellulus).

Holec, Czech Mycol. 53(2): fig. 3 between pages 138 and139, 2001. Ludwig: fig. 31.3. (as G. subsphaerosporus).Charac te r i s t i c s in b r ie f : F ru i tbod ies sma l l

(pileus up to 25 mm), surface fibrillose-tomentose, colourochre-brown, dark brown to rusty brown, lamellae typicallydark brown at maturity, spores mostly 4.56.0 3.54.5m,E = 1.11.4(1.5), Q about 1.25, subglobose but also broad-ly ellipsoid to broadly obovoid in side view, withoutsuprahilar disc and suprahilar depression, verrucose, cheilo-cystidia typically tibiiform with narrowly lageniform, cylin-drical or narrowly fusiform basal part, long narrow neck,distinct globose head and often thickened, rusty brown wall,pleurocystidia absent, caulocystidia of the same shape ascheilocystidia. Rare mountainous species growing mostly innatural and seminatural but also in man-made mixed orconiferous forests on strongly decayed wood of Picea or

Abies, especially on old stumps.D e s c r i p t i o n : Fruitbodies growing singly or in small

groups. Pileus 325 mm, at first subglobose to conical-sub-globose with involute margin, then convex to plano-convexor broadly campanulate, mostly with a broad low umbo,surface mat, dry but rarely slightly hygrophanous, finelyfibrillose-tomentose to distinctly tomentose, sometimes to-

13

Text-fig. 4. Gymnopilus josserandii: 1 Zto near Lenora,Paen mountain, JH 216/01 (PRM); 2 Prily, danidla

mountain, JH 52/02 (PRM); 3 JH Zto near Lenora,Boubnsk prales virgin forest, 580/01 (PRM). For explana-tions see Material and Methods. Scale bar = 10 m.

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mentose-scaly, colour variable, pale ochre, ochre to brown-ochre (5D7) when dry, but dark brown to rusty brown (6D8)when moist, old fruitbodies with rusty brown to purplish or-ange tinge, fibrillose-tomentose surface being yellow to yel-low-brown, drying surface being yellow-rusty (5B7).Lamellae rather sparse, L = 2435, l = 13, 23 mm high,

ventricose, near the stipe with a small decurrent tooth, yel-low with orange tinge when young, then sordid ochre-brownish (5D8), at maturity brown (6DE6), rusty brown(6D8) to typically dark brown (7E7), edge concolorous.Stipe 1035 0.150.4 mm, cylindrical or gradually thick-ened towards base, concolorous with pileus: beige-ochre,ochre-brown, ochre-rusty to rusty brown, surface distinctlyfibrillose-tomentose, covering pale yellow to yellow. Tastemild (without traces of bitterish taste). Smell indistinct.

Spores (4.0)4.56.0(6.4) (3.2)3.54.5(4.8) m, E =1.11.4(1.5), Q = 1.171.33 (variability measured in 4 spec-imens), mostly subglobose but also broadly ellipsoid to broad-

ly obovoid in side view, without suprahilar depression, in frontview subglobose, broadly ellipsoid to broadly lacrymoid;sometimes with a slightly polygonate outline, rusty ochre inKOH, wall rusty brown, medium thick, distinctly but notcoarsely verrucose, normal spores acyanophilous, those withbroken wall cyanophilous without any reaction in Melzersreagent or very slightly dextrinoid (with reddish brown hue onmature spores and spores with a broken wall). Basidia2030 57 m, narrowly cylindrical to narrowly clavate,often with median constriction, 4(2-) spored, sterigmata long,thin, 46 m. Cheilocystidia long, 2545 48(10) m,forming a sterile band on the edge, tibiiform with narrowly la-geniform, cylindrical or narrowly fusiform basal part, long

narrow neck (1.02.0 m) and distinct globose head(3.55 m), sometimes with slightly thickened and rustybrown wall, especially in the head (up to 1 m), content some-times homogeneously yellow-brown; rarely with hyaline la-geniform cystidia non-capitate. Pleurocystidia absent.Lamellar trama regular, hyphae 410(14) m broad, near thesubhymenium only 24 m, cells cylindrical to slightly inflat-ed (somewhere almost barrel-shaped), with distinct yellow-brown membranal and incrusting pigment, subhymenium notgelatinous, of densely arranged interwoven hyphae. Pileus cu-ticle (section) a cutis, not gelatinised, 3050m thick, formedby densely arranged parallel hyphae 412(14) m broad,

cells cylindrical, with yellow membranal pigment and coarserusty brown incrustations, under it a hypodermium of parallelto slightly flexuously interwoven hyphae 48 m broad, withsame type of pigmentation, in scalp visible as a loose net of in-terwoven hyphae, terminal cells sometimes slightly clavate,pileocystidia absent. Stipe cuticle 2-layered, lower layer acutis of parallel, densely arranged, cylindrical hyphae 28 mbroad, with yellow-rusty membranal pigment and rusty brownincrustations, from which emerge nests of loosely arrangedand interwoven hyphae 26 m broad, cylindrical but with la-geniform-fusiform outgrowths or terminal elements and withnumerous caulocystidia resembling cheilocystidia in shapeand size but often narrower (with cylindrical body). Clampconnections present in all tissues.

Fructification: July October (CR). The speciesproduces fruitbodies already in summer.

Ecology: In the Czech Republic, G. josserandii isknown from montane forests at an altitude of about 750 to1150 m. The forests are stands of Fagus, Picea,Abies and

Acer pseudoplatanus (so called mixed montane forest) or

pure Picea forests (or with admixed Fagus). The species ismostly found in natural stands (e.g. the Boubnsk pralesvirgin forest in the umava Mts. and ofnsk prales vir-gin forest in the Novohradsk hory Mts.) or near-naturalforests, but finds from man-made spruce forests are alsoknown (see Holec 2001b). Gymnopilus josserandii preferswood of Picea abies but it was also found on Abies alba.Records from broadleaved trees are also reported (e.g.Josserand 1948, but with a question mark). The species istypical of strongly decayed stumps of old trees, mostly cov-ered with mosses. I have never seen it on fallen trunks.However, Keller et Moser (2001) and Beran (personal com-munication) report that G. josserandii grows on decayedfallen trunks of conifers in Austria.

D i s t r i b u t i o n : Gymnopilus josserandii is relativelyrare in the Czech Republic as it is known only from three re-gions the umava Mts. and Novohradsk hory Mts. in Bo-hemia and the Beskydy Mts. in Moravia (see Antonn etkubla 2000, Holec 2001b and the collections studied). Inall these regions the species is rare but regularly occurs insuitable habitats natural, seminatural to man-made mixedor coniferous forests with presence of old and strongly de-cayed stumps (or trunks) of Picea orAbies.

In Europe, G. josserandii is well documented fromFrance (Josserand 1948, as Naucoria subsphaerospora),

Switzerland (Breitenbach et Krnzlin 2000: 140, as G. sub-bellulus), the Netherlands (Arnolds et al. 1995, as G. sub-sphaerosporus), Germany (Krieglsteiner 1991; Luschka1993: 197; Ludwig 2001: 152; in all cases as G. sub-sphaerosporus) and Austria (Keller et Moser 2001, as G.subsphaerosporus), mostly from mountains. The species isnot reported from Great Britain (Orton 1993) and Nordiccountries (Hiland 1990, Ryman 1992). Generally, G.

josserandii can be characterised as a rare mountainousspecies (but rarely growing also in lowlands, e.g. in theNetherlands) known above all from mountains of CentralEurope (the Alps, Bayerischer Wald + the umava Mts.,

Beskydy Mts.). Its occurrence in suitable habitats of theCarpathians and Balkan mountains is expected. The findfrom Corsica published by Bon et Roux (2002) as G.

josserandii represents G. bellulus (results of a microscopicrevision of the depicted specimen: coll. no. 3582 from theprivate herbarium of P. Roux) as it has a moist and lustrouspileus surface whereas the true G. josserandii typically hasdry and fibrillose-tomentose surface of the whole fruitbody(compare also plate 4-A by Bon and Roux with my pho-tographs). Moreover, the shape of cheilocystidia depictedby Bon and Roux (2002: p. 31, fig. G) is atypical of G.

josserandii, which has cystidia with more prominent glo-bose head and narrow neck.

D i s c u s s i o n : Gymnopilus josserandii is well recog-

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nisable by its small fruitbodies with fibrillose-tomentosesurface, dark brown colour of mature lamellae, mild taste,small and almost globose spores having no suprahilar de-pression, cheilocystidia of a distinct shape (tibiiform with anarrowly lageniform or cylindrical basal part, long narrowneck and globose head, often with slightly thickened and

rusty brown wall) and growth on strongly decayed wood ofconifers (Picea, Abies), typically on old decayed stumps.Gymnopilus bellulus also has small fruitbodies and smallspores but differs in spore shape (more ellipsoid with dis-tinct suprahilar depression), brighter colour of fruitbodies(pileus rusty orange to rusty brown with orange tinge,lamellae deep yellow, then yellow-rusty), slightly lustrouspileus surface and different appearance of cheilocystidia(thin-walled, head not so prominent).

In European literature, G. josserandii has been known asGymnopilus subsphaerosporus (Joss.) Khner et Romagn.for a long time. Unfortunately, this name is invalid becauseof a lacking Latin diagnosis. Consequently, Antonn (in An-tonn et kubla 2000) described the species validly asGymnopilus josserandii in honour of Marcel Josserand whorecognised it for the first time and published a perfect de-scription with exact line drawings.

My records well agree with the description by Josserand(1948: 2123) and the later description and colour photo-graph by Antonn (in Antonn et kubla 2000: 1316).However, I did not observe so many types of caulocystidiaas Antonn did, but only those resembling the cheilocys-tidia. The fruitbodies found by Josserand and Antonn havelonger stems (up to 5 cm).

The record described and photographed by Breitenbach

et Krnzlin (2000: p. 140141) and identified asGymnopilus subbellulus Hesler certainly representsGymnopilus josserandii. Almost all characters well agreewith the descriptions mentioned above. The only exceptionsrepresent the bitterish taste and presence of pileocystidiagiven by Breitenbach and Krnzlin. The authors obviouslyknew the invalid status of the name G. subsphaerosporus(which is cited by them as a synonym of G. subbellulus) anddecided to use the valid name by Hesler. The correctness ofthis conclusion is discussed below.

Gymnopilus subbellulus Hesler, North American species ofGymnopilus: 46, 1969 (in Mycologia Memoir no. 3) was de-

scribed from Michigan and California as a species of Gymno-pilus sect. Microspori. According to Hesler (1969), it isdistinguished by the following characters: non-dextrinoid, ellip-soid, ovoid to subglobose spores reaching 3.55.0 2.43.8 m,pleuro- and cheilocystidia both present, furfuraceous pileusand mild taste. If only the data from the book by Hesler (1969)are considered, many characters of G. subbellulus are reallyvery close to G. josserandii. However, the presence of pleuro-cystidia is in conflict with all European descriptions of thisspecies as well as with the records presented here, where nopleurocystidia were observed in spite of long and carefulsearch. For these reasons I considered Gymnopilus subbellu-lus Hesler a species different from Gymnopilus josserandiiAntonn (see Holec 2001b).

Later I studied the original material of G. subbellulusfrom Michigan herbarium (MICH). Surprisingly, the holo-type (coll. Smith 49838) is different from the paratype (coll.Smith 56336), which represents a species close or identicalwith G. bellulus (it has quite different spores: ellipsoid witha distinct suprahilar depression). This means that Hesler

(1969) did not have a clear concept of his new species.However, the use of the name G. subbellulus is fixed by theholotype which is microscopically very similar to the Euro-pean G. josserandii except for a rare presence of pleurocys-tidia and more ellipsoid (E = 1.151.62, Q = 1.31) and moredistinctly verrucose spores. In contrast with my previousopinion (Holec 2001b), G. subbellulus could eventually beconspecific with G. josserandii. However, more Americancollections must be compared with the European ones tojudge the variability of the American population and con-firm or refuse conspecifity. At this moment, I prefer to usethe European name G. josserandii for European collections.

C o l l e c t i o n s s t u d i e d :Czech Republic umava Mts., esk leby, Splenit mountain,on decayed stump of a conifer (Picea?, Abies?) covered withmosses, 26 Oct 2002, leg. J. Holec (PRM 900961). Ditto,Abies alba, on decayed stump, 13 Oct 1997, leg. J. Holec, JH755/97 (PRM 891945). Ditto, decayed stump covered withmosses (Abies?, Picea?), 26 Oct 2002, leg. J. Holec, JH 521/02(PRM). umava Mts., Jelen Vrchy near Nov Pec, Podkanlem protected area, Picea abies, on decayed stump, 30 Sep2000, leg. J. Holec, JH 173/00 (PRM 897842). umava Mts.,Prily, danidla mountain, Picea abies, on decayed stumpamong mosses, 16 July 2002, leg. J. Holec, JH 52/02 (PRM898682). umava Mts., Strn, Strn mountain, Piceaabies, on decaying stump among mosses, 28 Aug 2001, leg. J.

Holec, JH 187/01 (PRM). umava Mts., Zto near Lenora,Paen mountain (Boubn mountain group), Picea abies, on de-cayed stump among mosses, 30 Aug 2001, leg. J. Holec, JH216/01 (PRM). umava Mts., Zto near Lenora, Boubnskprales virgin forest, Picea abies, on decayed stump amongmosses, 2 Oct 2001, leg. J. Holec, JH 580/01 (PRM). Novohradsk hory Mts., ofnsk prales virgin forest, 19 Sep2004, leg. M. Beran (CB). Ditto, Picea abies?, in cavity of astrongly decayed stump, 27 Aug 2004, leg. M. Beran (CB).

Gymnopilus flavus (Bres.) Singer(Text-fig. 5, Pl. 12)

Bas.: Naucoria flava Bresadola, Ann. Mycol. 3: 162, 1905. Gymnopilus flavus (Bresadola) Singer, Lilloa 22: 561, 1951

(1949). Flammula flava (Br

gymnopilus republica tcheca

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