Floating Island Macromycetes from the Carpatho-Pannonian Region in Europe

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Zoologisch-Botanische Datenbank/Zoological-Botanical Database

Digitale Literatur/Digital Literature

Zeitschrift/Journal: Sydowia

Jahr/Year: 2009

Band/Volume: 61

Autor(en)/Author(s): Zöld-Balogh A., Dima B., Albert László, Babos M., Balogh M., Bratek Zoltán

Artikel/Article: Floating island macromycetes from the Carpatho-Pannonian Region in Europe. 149-176 ©Verlag Ferdinand Berger & Söhne Ges.m.b.H., Horn, Austria, download unter www.biologiezentrum.at

Floating island macromycetes from the Carpatho- Pannonian Region in Europe

AÂ .ZoÈld-Balogh1, B. Dima2, L. Albert3, M. Babos4, M. Balogh5 & Z. Bratek1

1 Department of Plant Physiology and Molecular Plant Biology, EoÈtvoÈs LoraÂnd University, PaÂzmaÂny PeÂter seÂtaÂny 1/c, Budapest, H-1117 Hungary 2 Department of Nature Conservation and Landscape Ecology, Faculty of Agriculture and Environmental Sciences, Szent IstvaÂn University, PaÂter KaÂroly u. 1, GoÈdoÈlloÍ, H-2103 Hungary 3 Karthauzi u. 4/a, Budapest, H-1121 Hungary 4 Szentes u. 52/a, Budapest, H-1147 Hungary 5 Paluster Bt. for Ecology and Conservation, VoÈlgy u. 21, Budapest, H-1214, Hungary

ZoÈld-Balogh AÂ ., Dima B., Albert L., Babos M., Balogh M. & BratekZ. (2008) Floating island macromycetes from the Carpatho-Pannonian Region in Europe. ± Sydowia 61 (1): 149±176. This study summarizes research data of the last 50 years on basidiomes and ascomycetes from sudds in the Carpathian and Pannonian Regions in Europe. The 76 basidiomycetes taxa collected from Sphagnum sudds (282 collections) belong to 3 orders (Agaricales, Boletales and Russulales), 15 families, and 25 genera. The 77 collections from non-Sphagnum sudds contained 33 species representing 4 orders (the above mentioned and Cantharellales), 12 families and 18 genera. Only three species were found in both types of sudds: Cortinarius uliginosus, Psathyrella typhae and Russula laccata. The 69 collections of ascomycetes (collected on the Sphagnum-free habitats of both kinds of floating fens) contained 27 species belonging to 4 orders (Dothideales, Helotiales, Orbiliales, Pezizales) 11 families and 19 genera. Representative ascomycetes included Lachnum virgineum, Mollisia ligni and Scutellinia crinita. More than half of the basidiomycetes are mycorrhizal (66% in Sphagnum sudds and 55% in non-Sphagnum sudds), while 92% of the ascomycetes proved to be saprobes. The following species occurred exclusively in floating islands of the studied area: Cortinarius tubarius, C. uliginosus, Hygrocybe coccineocrenata, Inocybe lacera var. helobia, Omphalina gerardiana, Russula betularum and R. laccata. These strongly rarefied habitats are the last asylums of several rare and unique species. Keywords: Ascomycota, Basidiomycota, habitat conservation, non-Sphagnum sudd, Sphagnum sudd

Floating islands are various types of natural freshwater habitats that can support fungi. A floating island is an intermediate habitat type of freshwater environments; it includes the interfaces between land and water (Tsui & Hyde 2003). They have been less surveyed than any other type of freshwater habitats, which could also include

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basin, shore, slope, flat and raised bogs formed on waterlogged soil (Larson 1991, Mitsch & Gosselink1993). The ecological importance of wetlands is now widely recognised. Catallo (1993) called wetlands nature's lungs and nature's kidneys; along the same analogy, they could also be described as nature's womb (nursery areas for aquatic organisms) and according to Mitsch & Gosselink(1993) they function as liver (detoxification centre for pollutants and excess nutrients). Wetlands are known to be the home of a large number of recognized rare species; this is obviously partly due to the fact that wetland habitats themselves are rare (Doust & Doust 1995). Floating islands ± as interesting variants in freshwater marshes of wetlands ± are characterised by peat soil with a balanced cold soil temperature, a constantly cool and moist microclimate. They are nutrient-poor, partly anoxic, often acidic, and are often exposed to wind and sun. A floating island is covered by stands of reeds, high sedge or peat mosses. Successional development can lead to woody vegetation, either scrubforest or sometimes high forest (Balogh et al. 1980b, Rydin & Jeglum 2006). The origin and botanical succession of floating fens, their role in the succession of sedimentation, and the development of peat have been explored by Weber (1908), Pallis (1915), Ruttner (1963), Heil- man (1968), Junk(1970), Swan & Gill (1970), Moore & Bellamy (1974), Balogh & FrenyoÂ (1980), Balogh (1981, 1983, 2000a, b, 2001a, 2002, 2003a), Huffman & Lonard (1983), Verhoeven (1986), Anthony (1999), Hackney et al. (2000), Bauer et al. (2003) and Rydin & Jeglum (2006). Their positive effects on water quality expressed as halobity, saprobity, toxicity and trophity have been known for decades (Balogh 1980, 1983, Balogh et al. 1980a, 1982, 1992). Based on these features they can be used for water cleaning (Balogh 1983). Sudds are important in ecosystem functioning (Balogh 1980, 1983, Mitsch & Gosselink1993) but their nomenclature has not been standardised yet. The scientific terms describing them vary greatly, they are inconsistent, e.g. in Africa they have been referred to as floating raft, floating swamp, island of floating vegetation, mat of floating vegetation, floating island (Denny 1991), sudd (Finlayson & Moser 1991). In North America and Europe they are recognized as floating marsh, floating meadow, floating island, floating mat, floating bog, quaking bog, floating fen and quaking fen, quaking mat and quaking mire (Mitsch & Gosselink 1993, Rydin & Jeglum 2006) and spring fen (VasÏutovaÂ 2005). Floating islands have long been known in literature. In Europe the first known description was given by the French Father Fournier (1643±1667, cited by Merrien 1975). The common memory preserving the former large networkof Carpatho-Pannonian sudds has been

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immortalized by the ethnographic literature (Herman 1887/1888, Balassa 1975, Gunda 1977, 1980). On the basis of information from other continents as well, we can state that floating islands are numerous and diverse on Earth and they can be found everywhere from the arctic to the Antarctic Circle (Balogh 1983, Van Duzer 2004, 2006) and exist until intrusive and expansive water management is implemented. Crucial changes have occurred since pre-settlement times; the original floating islands (inland and coastal) have been decreasing because of drainage and other human development (Mitsch & Gosselink1993). There are very few available sources in the literature on sudd fungi. Stendlung & Charvat (1994) published on the colonisation of mycorrhizal fungi of Typha in floating mats. Moreau (2002) described Hygrocybe coccineocrenata, Hypholoma elongatum and Rick- enella fibula var. hydrina as typical Sphagnum sudd basidiomes in French floating fens. Zehfuss (2005) published a detailed report on 16 basidiomes from a Sphagnum sudd in the German Schwarzwald Mountains, also identified by the authors of the present study in the Carpatho-Pannonian Region. In the Czech Republic some sudds were researched in 14 localities of Moravian-Slovakian borderland area (VasÏutovaÂ 2005). During three years 592 records were found including 171 macrofungi species in 46 spring fens of all 4 fen types. Several studies have been published on the sudd fungi in the Carpatho-Pannonian Region; on basidiomes (Babos 1979, 1987, 1989, LaÂszloÂ et al. 1988, Balogh 1989, BathoÂ 1994, LaÂzaÂr et al. 2000; Albert et al. 2004, Albert & Dima 2005, 2007), on arbuscular mycorrhizas (ReÂpaÂs et al. 1998, ZoÈld-Balogh et al. 2002a, b, 2003), on ascomycetes (Bratekand ZoÈld-Balogh 2001, 2002) and on symbiotic fungi of sudd orchids (IllyeÂs et al. 2005a, b). The studied area (with largely continental climate) is located in East-Central Europe. It belongs to three countries: Hungary, Slovakia and Romania (N458±N508, E168±E258). The inventory of sudd macromycetes was carried out in the habitat fragments with submontane climate at altitudes between 100 m and 500 m in the Hungarian area. Fragments with montane climate were sampled in the Carpathians (in Slovakia and Romania) at elevations ranging from 1000 m to 1700 m. These studied areas are relatively small and they are rare geomorpho- logical formations in the Carpatho-Pannonian Region, reflecting the existing fragmentation due to natural or social causes.

Materials and Methods The present study is based on irregular ± though covering all seasons ± field workby the authors. Emphasis has been made on collecting representatives of all the appearing macromycetes species.

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Sudd basidiomes have been collected since 1959 and ascomycetes since 1994. For identification of collected species the following works were used: Dennis (1981), Moser (1983), Breitenbach & KraÈnzlich (1984), BaÂnhegyi et al. (1985), Brandrud et al. (1989±1998), Schumacher (1990), LizonÏ (1992), Courtecuisse & Duhem (1995), Hawksworth et al. (1995), Hengstmengel (1996), Yao & Spooner (1996), Krieglsteiner (2000±2003), Bratek& ZoÈld-Balogh (2001), BelluÂ et al. (2004) and Vesterholt (2005). Fungal taxonomies follow the categories of The Index Fungorum (CABI 2008). In case of fungal nomenclature The Index Fungorum (CABI 2008) was also used, except for the species of genus Leccinum where den Bakker & Noordeloos (2005) and of the genus Hebeloma where Vesterholt (2005) were consulted. The collected specimens can be found in the authors' own herbaria and partly in the Herbarium of the Botanical Department of the Hungarian Natural History Museum (BP). Herbarium acronyms follow Holmgren et al. (1990). For abbreviation of authors' names the workof Kirk& Ansell (1992) was followed.

Results A comprehensive description of basidiomes and ascomycetes found on floating islands in the studied area is presented here, including the evaluation of mostly previously published data. The attributes of habitats, the substrates in cases of saprobic fungi and the plant symbionts of mycorrhizal associations are also presented along with the authors and their published data (Tables 1, 2, 3). Collectors' names without years in the tables denote unpublished data. The collection sites are marked 1±34 in the map (Fig. 1.) and listed in the index (Table 4). Names of the host or substrate plant species in sudds are listed in Table 5. Table 6 contains the distribution of collections according to months, and Table 7 to seasons.

Basidiomycota Sudd basidiomes examined so far originate mainly from Sphag- num habitats: 282 records of 76 taxa (Table 1). In spite of the relatively low number of samples, non-Sphagnum sudds proved to be species rich: 33 species from 77 samples (Table 2). The data on Sphagnum sudds were collected in 22 localities (14 from Hungary, 5 from Romania and 3 from Slovakia, respectively); whereas those of non-Sphagnum sudds came from 17 localities (15 from Hungary and 2 from Romania) (Tables 1, 2).

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Fig. 1. ± The collection sites (1±34) in the Carpatho-Pannonian region. (Image reproduced with kind permission of ELTE TeÂrkeÂptudomaÂnyi eÂs Geoinformatikai TanszeÂk, Budapest, Hungary.)

Most basidiome samples from Sphagnum sudds were collected in Kis-Mohos (RN11 ± 53 samples) in Lucs (RN27 ± 46 samples) and in Mohos (RN29 ± 44 samples). Most basidiomes of non-Sphagnum sudds were collected in Lake Nagy (RN16 ± 15 samples) and in KereÂk Waters of Lake Velencei (RN19 ± 14 samples). A strong seasonal behavior was shown. No basidiomycetes were found between December and March in any of the surveyed sudds (Tables 6). In Sphagnum sudds basidiomycetes were detected during seven months; 60 % of the collections were found from August to Septem- ber. Basidiomycetes were collected in non-Sphagnum sudds during eight months; the most fruitful months were September and October (nearly half of the data). In Sphagnum sudds the summer data out- number those of the autumn; whereas in non-Sphagnum sudds the opposite was found (Tables 7). In Sphagnum sudds only a limited number of samples were collected in spring; whereas in non-Sphag- num sudds numerous records were collected in spite of the low number of species. Mycorrhizal fungi were mostly found under the same host plants both in the Sphagnum sudds and the non-Sphagnum sudds: Betula

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spp., Salix spp., Picea abies and Pinus sylvestris. The 103 collections suggest that Betula pubescens was the dominant host in Sphagnum sudds while Salix cinerea (22 collections) was the dominant host species in non-Sphagnum sudds. Sphagnum and Phragmites were the most common substrate types in Sphagnum sudds and in non-Sphagnum sudds, respectively. The representatives of Agaricales were found in the highest number in both sudd types followed by species of Russulales. As Tables 1 and 2 show, in Sphagnum sudds Galerina paludosa, Lactarius helvus and Russula emetica were the most commonly found species in summer, whereas Leccinum variicolor was common in autumn. Due to the relatively low number of collections, none of these species has been considered common so far. A notable difference was shown in species distribution between the fringe zones and the central areas (montane type) of a sudd. In the sudd fringe zones, where usually Salix species dominate, their mycorrhizal fungi occurred: Cortinarius cinnamomeoluteus, C. helobius, C. uliginosus, Hebeloma pusillum, Inocybe salicis and Russula laccata. Betula species and their mycorrhizal partners were abun- dant both in the fringe and the central zones of sudds. Characteristic species were Cortinarius pholideus, C. tubarius, Leccinum cyaneobasileucum, L. holopus, L. variicolor, and Russula betularum. Picea abies occurred only in the fringe zones of sudds in montane regions. Under Picea, which is ± due to the similarly favourable habitat conditions ± widespread on sudds and on terrestrial mires alike, the mycorrhizal fungi Cortinarius subtortus and Russula emetica, for example, occur more frequently. Pinus sylvestris and its mycorrhizal partners appeared mostly inside floating fens, e.g.: Cortinarius huronensis, Suillus bovinus, and Suillus variegatus. Occasionally Laccaria proxima became frequent in sudds under Betula as well as Salix and Pinus. Other ubiquites were also found in floating mats: Amanita fulva, Cortinarius semisanguineus, and Paxillus involutus. The mycorrhizal species (66 %) outnumbered the saprobes (34%) both in the Sphagnum sudds and in the non-Sphagnum sudds (55% and 45%). Having found too few data on sudds in the international literature, the distribution of fungal species according to their life styles was compared to the data of Finnish peatlands. The propor- tion of mycorrhizal and saprobic species is balanced in the Finnish peatlands (Salo 1993), just like the proportions between fungal species in Carpatho-Pannonian sudds. Lactarius helvus, Leccinum variicolor and Russula laccata can be considered the most widespread mycorrhizal species in the Sphagnum sudds of the Carpatho-Pannonian Region. In Sphagnum

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sudds Hypholoma elongatum, Galerina paludosa and G. tibiicystis proved to be the most common saprobic basidiomes in contrast with Mycena belliae in non-Sphagnum sudds.

Ascomycota This study results in about 27 ascomycetes species found in 69 collections during 38 forays. The collected items came from 12 Hun- garian and 1 Romanian localities. As Table 3 shows, most ascomycete collections (24) came from KereÂkWaters of LakeVelencei (RN19) and from SzigetcseÂp (RN21, 16 collections). The data originated from Sphagnum sudds and non-Sphagnum sudds alike; from the peat soil or from putrescent remains of plants. Except for Mitrula paludosa the studied ascomycete species occurred in Sphagnum-free micro- habitats. Similarly to basidiomycetes, there were no ascomycetes found from December to February (Table 6). Most collections originated in October (16) followed by June (14 collections). As data in Table 7 demonstrate, similar numbers of collections were obtained from spring to autumn, indicating that sudds have a balanced cool microclimate throughout the growing season. Geopora tenuis was found most frequently with Lachnum virgineum in summer, Scutellinia crinita dominated in spring and Mollisia ligni was frequent in autumn. Scutellinia crinita was found in all three gath- ering seasons in five habitats. Although Geopora tenuis was found only in three habitats and in two seasons, it was so common there that it can be considered one of the most frequent ascomycetes of sudds. Most (70 %) of the ascomycetes collections were found on remains of woody plants, 19 % originated from herbs 3 % were collected on the ground. Geopora tenuis and Morchella elata are presumed to form mycorrhiza based on the descriptions of the mycorrhizas of closely related species (Agerer 1987±2002, Buscot & Kottke 1990, Duchesne & Weber 1993, Dahlstrom et al. 2000, Reinoso et al. 2005, IllyeÂs 2006, Tedersoo et al. 2006). The majority of the studied species live as saprobes (92 %) on dead leaves or stems of non-arboreal plants or in putrescent sub- stance of arboreal plants. Almost half of the sudd ascomycetes were associated with willow species (as substrate or host). Willows proved to be the most appropriate substrate for 43 % of the collections and for nearly half of the listed species. The following ascomycetes can be considered as common sudd species: Geopora tenuis, Lachnum virgineum, Mollisia ligni, Orbilia luteorubella and Scutellinia crinita. These five species were found in

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most examined habitats and gave the half of all records in this study. During our investigations we did not find any sudd-specialist ascomycetes.

Tab. 1. ± Basidiomycetes in Sphagnum sudds. (Date = month, year) (s.n. = sub. nom., n.d. = no data, rot. = rotted, lv. = leaves).

Species Locality Substrate Host Date Reference

Amanita fulva (Schaeff.) Fr. 29 Bet. pub. 08.1964 LaÂszloÂ (1970) 29 Pic. abi. 07.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) 27 Bet. pub. 08.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 07.1999 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) 27, 29 Bet. pub. 08.08.1999 LaÂzaÂr et al. (2000) 11 Bet. pub. 09.1990 Albert et al. (2004) 12, 11 Bet. pub. 09.10.1987 BathoÂ A. porphyria Alb. & Schwein. 1 Bet. pub. 08.2002 Albert et al. (2004) Cantharellula umbonata 29 Sph. 11.1967 LaÂszloÂ (1970) (J.F. Gmel.) Singer 30 Sph. 09.1980 LaÂszloÂ et al. (1981) 25 Pol. com. 07.1989 Albert et al. (2004) 1 Pol. com. 10.1999 Albert et al. (2004) Clitocybe fragrans (With.) 30 Sph. 09.1980 LaÂszloÂ et al. (1981) P. Kumm. Cortinarius acutus (Pers.) Fr. 11 Bet. pub. 09.1990 Albert et al. (2004) C. cinnamomeoluteus P.D. Orton 16 Sal. cin. 09.1988 Albert et al. (2004) (s.n.: C. salignus (M.M. Moser & Gerw. Keller) G. Garnier) C. cinnamomeus (L.) Fr. 27 Pin. syl. 09.1986 Albert et al. (2004) (s.n. C. cinnamomeobadius Rob. Henry) C. decipiens var. atrocoeruleus 19 Sal. cin. 05.2004 Albert et al. (2004) (M.M. Moser ex M.M. Moser) H. Lindstr. (s.n. C. atrocoeruleus M.M. Moser ex M.M. Moser) C. flexipes (Pers.) Fr. 11 Bet. pub. 09.1990 Albert et al. (2004) (s.n. C. paleiferus SvrcÏek) 11 Bet. pub. 09.1992 Albert et al. (2004) 11 Bet. pub. 09.1992 BathoÂ C. helobius Romagn. 21, 21 Sal. cin. 05.08.1990 Albert et al. (2004) C. huronensis Ammirati & 27 Pin. syl. 09.1986 Albert et al. (2004) A.H. Sm. 32 Pin. syl. 10.1986 Albert et al. (2004) C. multicolor M.M. Moser ex 28 Pic. abi. 06.1983 Albert et al. (2004) M.M. Moser C. pholideus (Fr.) Fr. 13, 13 Bet. pub. 10.10.1959 Babos (1989) C. semisanguineus (Fr.) Gillet 27 n.d. 08.1969 LaÂszloÂ (1972) (s.n. Dermocybe 1 Pin. syl. 10.1985 Albert et al. (2004) semisanguinea (Fr.) M.M. Moser) 14 Bet. pub. 08.2003 Dima C. subtortus (Pers.) Fr. 27 Pic. abi. 09.1986 LaÂszloÂ et al. (1988)

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Tab. 1. ± Continued

Species Locality Substrate Host Date Reference

C. tubarius Ammirati & A.H. Sm. 27 Pin. syl. 08.1969 LaÂszloÂ (1970) (s.n. Dermocybe palustris var. 11 Bet. pub. 08.1988 BathoÂ (1994) sphagneti (P.D. Orton) M.M. 29 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) Moser, C. sphagneti P.D. Orton) 29, 29 Pin. syl. 07.08.1999 LaÂzaÂr et al. (2000) 27 Pin. syl. 09.1986 Albert et al. (2004) 12 Bet. pub. 10.1987 Albert et al. (2004) 12 Bet. pub. 09.1992 Albert et al. (2004) C. uliginosus Berk. 1 Sal. aur. 08.2002 Albert et al. (2004) Entoloma cuspidiferum 11 Sph. 10.1987 Albert et al. (2004) Noordel. E. sphagneti Naveau 30 Sph. 09.1980 LaÂszloÂ et al. (1981) 27 Sph. 09.1986 Albert et al. (2004) Galerina gibbosa J. Favre 1 rot. Sph. 06.2001 Albert et al. (2004) G. paludosa (Fr.) KuÈ hner 11, 12 Sph. 07.07.1988 BathoÂ (1994) 11, 12 Sph. 08.09.1988 BathoÂ (1994) 29, 29 Sph. 07.08.1998 LaÂzaÂr et al. (2000) 27 Sph. 08.1998 LaÂzaÂr et al. (2000) 25 rot. Sph. 06.1988 Albert et al. (2004) 16 rot. Sph. 07.1988 Albert et al. (2004) 16 rot. Sph. 07.1990 Albert et al. (2004) 1 rot. Sph. 11.2000 Albert et al. (2004) 18 rot. Sph. 07.1988 Albert et al. (2004) 14 rot. Sph. 08.2003 Albert & Dima (2007) G. tibiicystis (G.F. Atk.) KuÈ hner 1 rot. Sph. 06.1985 Albert et al. (2004) 12 rot. Sph. 07.1988 Albert et al. (2004) 12 rot. Sph. 08.1992 Albert et al. (2004) 3 rot. Sph. 10.1989 Albert et al. (2004) 1 rot. Sph. 08.2004 Albert et al. (2004) 18 rot. Sph. 07.1988 Albert 14 rot. Sph. 08.2003 Dima Gymnopilus fulgens 25 rot. Sph. 07.1989 Albert et al. (2004) (J. Favre & Maire) Singer 16 rot. Sph. 09.1991 Albert et al. (2004) 1 rot. Sph. 10.2001 Albert et al. (2004) Hebeloma helodes J. Favre 27 n.d. 08.1969 LaÂszloÂ (1970) H. incarnatulum A.H. Sm. 16 Bet. pen. 09.1989 Albert et al. (2004) (s.n. H. longicaudum (Pers.) P. Kumm.) H. velutipes Bruchet 11 Bet. pub. 09.1990 Albert et al. (2004) (s.n. H. longicaudum (Pers.) P. Kumm.) Hygrocybe coccineocrenata 1, 5 Sph. 08.08.2002 Albert et al. (2004) (P.D. Orton) M.M. Moser 1, 5 Sph. 08.08.2003 ZoÈld-Balogh H. conica (Scop.) P. Kumm. 21 Sph. 09.2002 Albert et al. (2004) Hypholoma elongatum (Pers.) 29 Sph. 09.1968 LaÂszloÂ (1972) Ricken 27 Sph. 08.1969 LaÂszloÂ (1972) 12 Sph. 10.1987 Babos (1989) 11, 12 Sph. 06.09.1988 Babos (1989)

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Tab. 1. ± Continued

Species Locality Substrate Host Date Reference

11, 11 Sph. 07.08.1988 BathoÂ (1994) 27 Sph. 09.1986 Albert et al. (2004) 12 Sph. 09.1992 Albert et al. (2004) 1, 1 Sph. 10.11.2000 Albert et al. (2004) H. myosotis (Fr.) M. Lange 1 Sph. 08.2002 Albert et al. (2004) H. polytrichi (Fr.) Ricken 30 Sph. 09.1980 LaÂszloÂ et al. (1981) H. udum (Pers.) KuÈ hner 30 Sph. 09.1980 LaÂszloÂ et al. (1981) 29, 29 Sph. 07.08.1999 LaÂzaÂr et al. (2000) Inocybe lacera (Fr.) P. Kumm. 27 n.d. 08.1970 LaÂszloÂ (1979) I. lacera var. helobia Kuyper 16 n.d. 09.1988 RimoÂczi (1994) 17 Sal. cin. 07.1988 Albert et al. (2004) 16 Bet. pen. 07.1988 Albert et al. (2004) 16 Pop. alb. 07.1990 Albert et al. (2004) I. napipes J.E. Lange. 29 n.d. 08.1965 LaÂszloÂ (1972) Laccaria bicolor (Maire) P.D 27 n.d. 08.1969 LaÂszloÂ (1970) Orton L. proxima (Boud.) Pat. 12, 11 Bet. pub. 09.10.1987 BathoÂ (1994) 11, 12 Bet. pub. 08.08.1988 BathoÂ (1994) 11 Bet. pub. 08.1988 BathoÂ (1994) 29 Pic. abi. 08.1998 LaÂzaÂr et al. (2000) (s.n. L. laccata var. proxima 12 Bet. pub. 09.1990 Albert et al. (2004) (Boud.) Maire) 1 Bet. pub. 10.2000 Albert et al. (2004) 14 Bet. pub. 09.2004 Albert & Dima Lactarius glyciosmus (Fr.) Fr. 11 Bet. pub. 09.1990 Albert et al. (2004) 12 Bet. 09.1992 BathoÂ L. helvus (Fr.) Fr. 27 Pin./Bet. 08.1970 LaÂszloÂ (1972) 30 Bet. ver. 09.1980 LaÂszloÂ et al. (1981) 25 Bet. 08.1960 Babos (1989) 11 Bet. pub. 07.1987 Babos (1989) 11, 15 Bet. pub. 07.09.1988 Babos (1989) 29 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) 29, 29 Pin. syl. 07.08.1999 LaÂzaÂr et al. (2000) 12 Bet. pub. 09.1990 Albert et al. (2004) 12 Bet. pub. 08.1992 Albert et al. (2004) 14 Bet. pub. 09.2004 Siller et al. (2006) 29 n.d. 08.1964 LaÂszloÂ/Babos (BP) 11, 12 Bet. pub. 07.07.1988 BathoÂ 11, 12 Bet. pub. 08.08.1989 BathoÂ L. mammosus Fr. 29 Pic. abi. 08.1965 LaÂszloÂ (1970) L. rufus (Scop.) Fr. 30 Bet. ver. 09.1980 LaÂszloÂ et al. (1981) 11 Bet. pub. 09.1990 Albert et al. (2004) 11 Bet. pub. 08.1987 BathoÂ 12, 11 Bet. pub. 09.10.1987 BathoÂ 11, 12 Bet. pub. 08.08.1988 BathoÂ 14 Bet. pub. 09.2004 Albert & Dima L. sphagneti (Fr.) Neuhoff 27 n.d. 08.1970 LaÂszloÂ (1972) 27 Bet. pub. 08.1998 LaÂzaÂr et al. (2000)

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Tab. 1. ± Continued

Species Locality Substrate Host Date Reference

27 Pic. abi. 07.1999 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) L. trivialis (Fr.) Fr. 27 Pic. abi. 08.1969 LaÂszloÂ (1970) L. vietus (Fr.) Fr. 27 Bet. pub. 08.1969 LaÂszloÂ (1972) 30 Bet. pub. 09.1980 LaÂszloÂ et al. (1981) 12, 11 Bet. pub. 09.10.1987 BathoÂ (1994) 27 Bet. pub. 07.1998 LaÂzaÂr et al. (2000) 29 Pic. abi. 08.1998 LaÂzaÂr et al. (2000) 27 Bet. pub. 08.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 07.1999 LaÂzaÂr et al. (2000) 27 Bet. pub. 07.08.1999 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) Leccinum cyaneobasileucum 11, 11 Bet. pub. 10.10.1987 Albert et al. (2004) Lannoy & EstadeÁs 11 Bet. pub. 07.1988 Albert et al. (2004) (s.n. L. brunneogriseolum var. 16 Bet. pen. 09.1988 Albert et al. (2004) sphagnorum Lannoy & EstadeÂs) 16 Bet. pen. 09.1989 Albert et al. (2004) 16 Bet. pen. 09.1991 Albert et al. (2004) 1 Bet. 10.1985 Albert et al. (2004) 1 Bet. 09.1988 Albert et al. (2004) 14 Bet. pub. 09.2004 Albert & Dima L. holopus (Rostk.) Watling 31 Bet. pub. 09.1977 LaÂszloÂ (1979) 11 Bet. pub. 10.1988 Babos (1989) 11, 12 Bet. pub. 10.10.1987 BathoÂ (1994) 1 Bet. pub. 09.1987 Albert et al. (2004) 1 Bet. pub. 08.2001 Albert et al. (2004) 11 Bet. pub. 08.2002 Albert et al. (2004) 1 Bet. pen. 08.1990 Albert et al. (2004) 1 Bet. pub. 09.1988 Vasas (BP) L. variicolor Watling 16, 11 Bet. 10.10.1988 Babos (1989) 12 Bet. 10.1988 Babos (1989) 11 Bet. 10.1987 BathoÂ (1994) 11 Bet. pub. 08.1988 BathoÂ (1994) 11 Bet. pub. 09.1992 BathoÂ (1994) 29 Bet. pub. 08.1999 LaÂzaÂr et al. (2000) 27 Bet. pub. 09.1986 Albert et al. (2004) 33, 34 Bet. pub. 10.10.1986 Albert et al. (2004) 11 Bet. pub. 10.1987 Albert et al. (2004) 11 Bet. pub. 09.1992 Albert et al. (2004) 16 Bet. pen. 09.1988 Albert et al. (2004) 16 Bet. pen. 09.1989 Albert et al. (2004) 14 Bet. pub. 09.2004 Siller et al. (2006) 11 n.d. 10.1987 Vasas (BP) Marasmius epiphyllus (Pers.) Fr. 21 rot. lv. 11.2000 Albert et al. (2004) 21 rot. lv. 09.2002 Albert et al. (2004) Mycena epipterygia (Scop.) Gray 1 rot. pine 10.2000 Albert et al. (2004) needles M. megaspora Kauffman 30 Sph. 09.1980 LaÂszloÂ et al. (1981) (s.n. M. dissimulabilis (Britzelm.) Lapl.)

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Tab. 1. ± Continued

Species Locality Substrate Host Date Reference

Omphalina ericetorum (Bull.) 30 Pol. com. 09.1980 LaÂszloÂ et al. (1981) M. Lange 29 Pol. com. 07.1998 LaÂzaÂr et al. (2000) Omphalina ericetorum (Bull.) 29, 27 Pol. com. 07.07.1999 LaÂzaÂr et al. (2000) M. Lange (s.n. Phytoconis erice- 1, 1 Pol. com. 11.11.2000 Albert et al. (2004) torum (Pers.) Redhead & Kuyper) O. gerardiana (Peck) Singer 1, 1 rot. Sph. 10.10.1985 Albert et al. (2004) O. oniscus (Fr.) QueÂl. 30 Sph. 09.1980 LaÂszloÂ et al. (1981) 11 Sph. 09.1990 Albert et al. (2004) Paxillus involutus (Batsch) Fr. 27 Pic./Bet. 08.1969 LaÂszloÂ (1970) 11 Bet. pub. 09.1990 Albert et al. (2004) Pholiota squarrosa (Weigel) 29 snag of Aln. 09.1971 LaÂszloÂ (1972) P. Kumm. glu. Pluteus cyanopus QueÂl. 21 Phr. aus. muck09.2002 Albert et al. (2004) Psathyrella cf. trepida (Fr.) Gillet 16 Phr. aus. 07.1988 Albert et al. (2004) muck P. typhae (Kalchbr.) 16 Typ. muck 07.1988 Albert et al. (2004) A. Pearson & Dennis 16 Typ. muck 09.1995 Albert et al. (2004) Russula atropurpurea 1 Bet. pub. 10.2000 Albert et al. (2004) (Krombh.) Britzelm. R. betularum Hora 11, 12 Bet. pub. 07.07.1988 BathoÂ (1994) (s.n. R. emetica var. betularum 12 Bet. pub. 10.1987 Albert et al. (2004) (Hora) Romagn.) 11 Bet. pub. 07.1988 Albert et al. (2004) 11, 12 Bet. pub. 08.09.1992 Albert et al. (2004) 11 Bet. pub. 07.1988 BathoÂ 11, 11 Bet. pub. 08.08.1988 BathoÂ 12 Bet. pub. 09.1992 BathoÂ R. claroflava Grove 11 Pin. syl. 09.1992 BathoÂ (1994) (s.n. R. flava Romell) 27 Pic. abi. 07.1999 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) 11 Bet. pub. 09.1990 Albert et al. (2004) R. decolorans (Fr.) Fr. 27 Pic. abi. 08.1970 LaÂszloÂ (1972) 27 Bet. pub. 07.1998 LaÂzaÂr et al. (2000) 29 Pic. abi. 07.1998 LaÂzaÂr et al. (2000) 27 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) 27 Bet. pub. 08.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 07.1999 LaÂzaÂr et al. (2000) 27 Bet. pub. 07.1999 LaÂzaÂr et al. (2000) R. cf. elaeodes (Bres. ex 16 Bet. pen. 09.1988 Albert et al. (2004) Romagn.) Bon 16 Bet. pen. 09.1991 Albert et al. (2004) R. emetica (Schaeff.) Pers. 29 Pin./Pic. 08.1964 LaÂszloÂ (1970) 27 Pic. abi. 08.1969 LaÂszloÂ (1970) 27 Pic. abi. 08.1969 LaÂszloÂ (1972) 30 Pin. syl. 09.1980 LaÂszloÂ et al. (1981) 29 Pic. abi. 08.1999 LaÂzaÂr et al. (2000) 27 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) 27,27 Pic. abi. 07.08.1998 LaÂzaÂr et al. (2000)

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Tab. 1. ± Continued

Species Locality Substrate Host Date Reference

27 Pic. abi. 07.1999 LaÂzaÂr et al. (2000) 29 Pic. abi. 08.1998 LaÂzaÂr et al. (2000) R. fragilis Fr. 30 n.d. 08.1964 LaÂszloÂ (1970) R. gracillima Jul. Schaeff. 1 Bet. pub. 10.1991 Albert et al. (2004) R. laccata Huijsman 16 Sal. cin. 07.1988 Babos (1989) (s.n. R. atrorubens sensu J.E. 12 Sal. cin. 10.1988 Babos (1989) Lange) (s.n. R. aquosa Leclair) 21 Sal. cin. 05.1989 Babos (1989) 15 n.d. 08.1976 Babos (1989) 16 Sal. cin. 09.1987 Albert (2001) 12 Bet. pub. 10.1987 Albert et al. (2004) 21 Sal. cin. 10.1989 Albert et al. (2004) 21 Sal. cin. 11.2000 Albert et al. (2004) R. laccata Huijsman 1 Sal. cin. 08.2002 Albert et al. (2004) 19 Bet. pub. 10.2002 Albert et al. (2004) 1 Sal. cin. 10.2003 Albert et al. (2004) 11 Bet. pub. 09.1992 BathoÂ R. paludosa Britzelm. 27 Pic. abi. 08.1970 LaÂszloÂ (1972) 29, 29 Pin. syl. 07.07.1999 LaÂzaÂr et al. (2000) 29, 29 Pin. syl. 08.08.1999 LaÂzaÂr et al. (2000) 29 Bet. pub. 07.1998 LaÂzaÂr et al. (2000) 27 Bet. pub. 07.1999 LaÂzaÂr et al. (2000) 27 Pin. syl. 08.1999 LaÂzaÂr et al. (2000) 27, 12 Bet. pub. 07.08.1998 LaÂzaÂr et al. (2000) 29 Pin. syl. 08.1998 LaÂzaÂr et al. (2000) R. puellaris Fr. 1 Pic. abi. 11.2000 Albert et al. (2004) 1 Bet. pub. 11.2000 Albert et al. (2004) R. sphagnophila Kauffman 11 Bet. pub. 09.1992 Albert et al. (2004) 11 Bet. pub. 09.1992 BathoÂ R.cf.versicolor Jul. Schaeff. 16 Bet. pen. 09.1991 Albert et al. (2004) R. vinosa Lindblad 27, 28 Pic. abi. 07.08.1999 LaÂzaÂr et al. (2000) Suillus bovinus (Pers.) Roussel 29 Pin. syl. 09.1968 LaÂszloÂ (1970) S. variegatus (Sw.) Kuntze 27 Pin. syl. 08.1970 LaÂszloÂ (1972) Tephrocybe palustris (Peck) 25 Sph. 06.1988 Albert et al. (2004) Donk 14 Sph. 09.2004 Albert & Dima Tylopilus felleus (Bull.) P. Karst. 1 Bet. pub. 08.2002 Albert et al. (2004)

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Tab. 2. ± Basidiomycetes in non-Sphagnum sudds. (Date = month, year) (s.n. = sub. nom., n.d. = no data, rot. = rotted, lv. = leaves, bran. = branches).

Species Locality Substrate Host Date Reference

Clavulina coralloides (L.) J. SchroÈt. 14 Sal./Que. 08.2003 Dima Collybia tuberosa (Bull.) P. 20 rot. Phr. 10.1978 Babos (1989) Kumm. aus. Coprinus friesii QueÂl. 20 rot. Phr. aus. 08.1978 Babos (1989) 19 rot. Phr. aus. 08.2004 Albert et al. (2004) Cortinarius uliginosus Berk. 16 Sal. cin. 09.1990 RimoÂczi (1994) 16, 16 Sal. cin. 07.09.1988 Albert et al. (2004) 16 Sal. cin. 09.1989 Albert et al. (2004) 16 Sal. cin. 09.1995 Albert et al. (2004) 16 Sal. cin. 09.1997 Albert et al. (2004) Gymnopus dryophilus (Bull.) 2 rot. lv. of 10.2002 Albert et al. (2004) Murrill (s.n. Collybia dryophila Aln. glu. (Bull.) P. Kumm.) Hebeloma birrus (Fr.) Gillet 14 Sal./Que. 08.2003 Dima H. leucosarx P.D. Orton 20, 20 Sal. cin. 10.10.1978 Babos (1989) 18 Sal. cin. 10.2002 Albert et al. (2004) H. pusillum J.E. Lange 21 Sal. cin. 10.1989 Albert et al. (2004) 16 Sal. cin. 09.1995 Albert et al. (2004) Hypholoma laeticolor 12 rot. Phr. 10.1987 Albert et al. (2004) (F.H. Mùller) P.D. Orton aus. 9 rot. Phr. 09.1992 Albert et al. (2004) aus. H. polytrichi (Fr.) Ricken 1 Phr. aus. 10.1985 Albert et al. (2004) 11 Phr. aus. 10.1987 Albert et al. (2004) Inocybe acuta Boud. 11 Bet. pub. 09.1992 Albert et al. (2004) I. fuscomarginata KuÈ hner 19 Sal. cin. 05.2001 Albert et al. (2004) 19 Sal. cin. 05.2004 Albert et al. (2004) I. malencËonii R. Heim 21 Sal. cin. 05.1988 Albert et al. (2004) 21 Sal. cin. 05.1990 Albert et al. (2004) I. salicis KuÈ hner 16 Sal. cin. 09.1995 Albert et al. (2004) 16 Sal. cin. 09.1997 Albert et al. (2004) 19 Sal. cin. 06.2002 Albert et al. (2004) Lactarius lacunarum Romagn. 18 Sal. 07.1988 Albert & Dima (2005) ex Hora 14 Sal. cin. 08.2003 Albert & Dima (2005) 14 Sal. cin. 09.2004 Albert & Dima (2005) L. lignyotus Fr. 27 Pin. syl. 07.1998 LaÂzaÂr et al. (2000) 27, 27 Pic. abi. 07.08.1999 LaÂzaÂr et al. (2000) 28 Pic. abi. 08.1999 LaÂzaÂr et al. (2000) L. omphaliformis Romagn. 25 Aln. glu. 06.1988 Albert et al. (2004) L. serifluus (DC.) Fr. 14 Sal./Que. 08.2003 Dima Leccinum scabrum (Bull.) 27 Bet. pub. 09.1986. Albert et al. (2004) Gray 11 Bet. pub. 09.1990 Albert et al. (2004) [s.n. L. molle (Bon) Bon] 16 Bet. pub. 09.1991 Albert et al. (2004) Marasmius limosus Boud. & QueÂl. 19 Phr. aus. muck10.2002 Albert et al. (2004)

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Tab. 2. ± Continued

Species Locality Substrate Host Date Reference

Mycena belliae (Johnst.) P.D. 20, 20 rot. Phr. 11.11.1977 Babos (1989) Orton aus. stalk 20, 20 rot. Phr. 10.10.1978 Babos (1989) aus. 21 rot. Phr. 05.1988 Albert et al. (2004) aus. 21 rot. Phr. 04.1990 Albert et al. (2004) aus. 26 rot. Phr. 05.2002 Albert et al. (2004) aus. 19 rot. Phr. 10.2002 Albert et al. (2004) aus. 19 rot. Phr. 11.2003 Albert et al. (2004) aus. M. cf. metata (Secr. ex Fr.) 19 rot. debris 10.2002 Albert et al. (2004) P. Kumm. of Sal. M. sanguinolenta (Alb. & 16 Phr. aus. 07.1990 Albert et al. (2004) Schwein.) P. Kumm. muck 16 Phr. aus. 09.1995 Albert et al. (2004) muck Phaeomarasmius erinaceus 21 rot. bran. 05.1989 Albert et al. (2004) (Pers.) Scherff. ex Romagn. of Sal. 16 rot. bran. 09.1995 Albert et al. (2004) of Sal. 19 rot. bran. 11.2003 Albert et al. (2004) of Sal. sp. Pholiota henningsii (Bres.) 10, 17 rot. debris 05.07.1988 Albert et al. (2004) P.D. Orton of Sal. 16 rot. debris 07.1988 Albert et al. (2004) of Sal. 16 rot. debris 09.1997 Albert et al. (2004) of Sal. Pluteus chrysophaeus (Schaeff.) 19 rot. debris 08.2002 Albert et al. (2004) QueÂl. (s.n. P. luteovirens Rea) of Sal. Psathyrella typhae (Kalchbr.) A. 20 rot. Typ. lax. 10.1978 Babos (1989) Pearson & Dennis 19 rot. Typ. 05.2001 Albert et al. (2004) 19 rot. Typ. 06.2002 Albert et al. (2004) 19, 19 rot. Typ. 06.06.2004 Albert et al. (2004) Rickenella fibula (Bull.) Raithelh. 19 n.d. 05.2001 Albert et al. (2004) R. swartzii (Fr.) Kuyper 19 rot. Phr. aus. 08.2004 Albert et al. (2004) Russula cf. amoenicolor Romagn. 27 Pic. abi. 08.1998 LaÂzaÂr et al. (2000) R. insignis QueÂl. (s.n. R. livescens 27 Pic. abi. 07.1998 LaÂzaÂr et al. (2000) sensu J.E. Lange) R. integra (L.) Fr. 27 Pic. abi. 08.1998 LaÂzaÂr et al. (2000) 27 Pic. abi. 07.1999 LaÂzaÂr et al. (2000) 27 Pin. syl. 08.1999 LaÂzaÂr et al. (2000)

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Tab. 3. ± Ascomycetes in sudds. (Date = month, year) (s.n. = sub. nom., rot. = rotted, bran. = branches, ZB = ZoÈld-Balogh, rem. = remains, mos. = mossy, upr. = uprightly, bot. = bottom).

Species Locality Substrate Host Date Reference

Ascocoryne sarcoides (Jacq.) J.W. 7 dead bran. 10.1994 Bratek& ZB (2002) Groves & D.E. Wilson Ascophanus globosopulvinatus 21 rot. leaf of 07.1994 Bratek& ZB (2002) (Crossl.) Boud. ex Ramsb. Sal. cin. (s.n. cf. A. globoso-pulvinata (Crossl.) Boud.) Geopora tenuis (Fuckel) 21 rot. leaf of 07.1994 Bratek& ZB (2002) T. Schumach. Sal. cin. 24, 21 Sal. cin. 09.09.1994 Bratek& ZB (2002) 19 Sal. cin. 06.2000 Bratek& ZB (2002) 19, 19 Sal. cin. 06.09.2001 Bratek& ZB (2002) 19 rot. block of 11.2001 Bratek& ZB (2002) Sal. cin. 19 dead stalkof 06.2002 Bratek& ZB (2002) Typ. ang. 19 dead bran. of 06.2002 Bratek& ZB (2002) Sal. cin. 19 dead root of 06.2002 Bratek& ZB (2002) Sal. cin. Hymenoscyphus calyculus 1 barkof dead 11.2000 Bratek& ZB (2002) (Sowerby) W. Phillips bran. of Sal. (s.n. H. virgultorum (Vahl) W. Phillips) H. phyllophilus (Desm.) 21 dead leaf of 09.2002 Bratek& ZB (2002) O. Kuntze Sal. cin. H. salicellus (Fr.) Dennis 21 dead bran. of 07.1994 Bratek& ZB (2002) Sal. cin. 19 dead bran. of 10.2002 Bratek& ZB (2002) Sal. cin. H. scutula (Pers.) W. Phillips 19 rot. stem of 10.2002 Bratek& ZB (2002) Men. aqu. H. vitellinus (Rehm) O. Kuntze 21, 21 dead petiole of 07.09.1994 Bratek& ZB (2002) The. pal. 21 dead petiole of 09.2002 Bratek& ZB (2002) The. pal. Karschia bloxamii (Berk. & 19 dead bran. of 05.2001 Bratek& ZB (2002) W. Phillips) Sacc. Sal. cin Lachnum controversum (Cooke) 19 dead stalks of 05.2001 Bratek& ZB (2002) Rehm Phr.aus. (s.n. Dasyscyphus controversus 19, 19 dead stalks of 06.10.2002 Bratek& ZB (2002) (Cooke) Rehm) Phr.aus. 19 dead stalks of 10.2002 Bratek& ZB (2002) Phr.aus. L. virgineum (Batsch) P. Karst. 21 dead bran. of 07.1994 Bratek& ZB (2002) (s.n. Dasyscyphus virgineus Sal. cin. (Batsch) Gray)

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Tab. 3. ± Continued

Species Locality Substrate Host Date Reference

19 dead bran. of 06.2000 Bratek& ZB (2002) Sal. cin. 1, 19 dead bran. of 04.05.2001 Bratek& ZB (2002) Sal. aur. / cin. 1 dead bran. of 06.2001 Bratek& ZB (2002) Bet. pub. 5, 3 dead bran. of 06.06.2001 Bratek& ZB (2002) Sal. aur. 19 dead bran. of 05.2002 Bratek& ZB (2002) Sal. cin. Mitrula paludosa Fr. 27 rot. wood of 07.1998 LaÂzaÂr et al. (2000) Pic. abi. 27, 27 rot. wood of 07.08.1999 LaÂzaÂr et al. (2000) Pic. Mollisia ligni (Desm.) P. Karst. 1, 6 dead bran. 10.10.1994 Bratek& ZB (2002) 7 on rot. wood 10.1994 Bratek& ZB (2002) 8 dead bran. 10.1994 Bratek& ZB (2002) 19 dead bran. of 05.2002 Bratek& ZB (2002) Sal. cin. M. cinerea (Batsch) P. Karst. 1 rot. wood of 11.2001 Bratek& ZB (2002) Sal. M. palustris (Roberge ex Desm.) 19 dead stalkof 06.2002 Bratek& ZB (2002) P. Karst. Phr. aus. Morchella elata Fr. 19 ground 05.2002 Bratek& ZB (2002) Orbilia cf. curvatispora Boud. 7 rot. wood of 10.1994 Bratek& ZB (2002) Aln. glu. O. luteorubella (Nyl.) P. Karst. 21 dead bran. 09.1994 Bratek& ZB (2002) 6, 8 dead bran. 10.10.1994 Bratek& ZB (2002) 7 dead bran. 10.1994 Bratek& ZB (2002) Peziza limnaea Maas Geest. 5 rem. of wood 11.2000 Bratek& ZB (2002) 5 rem. of wood 11.11.2001 Bratek& ZB (2002) 5 rem. of wood 06.2002 Bratek& ZB (2002) Phaeohelotium umbilicatum 1 rot. leaf of 10.1994 Bratek& ZB (2002) (Le Gal) Dennis Que. pet. Psilachnum chrysostigmum (Fr.) 19 dead petiole of 06.2002 Bratek& ZB (2002) Raitv. (s.n. Pezizella cf. chrysos- The. pal. tigma (Fr.) Sacc.) Pulvinula convexella (P. Karst.) 21 rem. of wood 09.1994 Bratek& ZB (2002) Pfister 23 dead bran. of 03.2002 Bratek& ZB (2002) (s.n. P. constellatio (Berk. & Br.) Sal. cin. Boud.) Pyronema domesticum (Sowerby) 19 rot. wood of 03.2002 Bratek& ZB (2002) Sacc. Sal. cin. Rutstroemia sydowiana (Rehm) 21 rot. leaf of 07.2001 Bratek& ZB (2002) W.L. White Sal. cin. Scutellinia crinita (Bull.) 21 dead bran. of 04.1994 Bratek& ZB (2002) Lambotte Sal. cin.

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Tab. 3. ± Continued

Species Locality Substrate Host Date Reference

21 barkof dead 04.1994 Bratek& ZB (2002) bran. of Pop. 21 dead bran. of 07.1994 Bratek& ZB (2002) Sal. cin. 7 mos. rot. wood 10.1994 Bratek& ZB (2002) 1 rot. wood 10.1994 Bratek& ZB (2002) 19 dead stalkof 05.2001 Bratek& ZB (2002) Phr. aus. 21 dead bran. of 06.2001 Bratek& ZB (2002) Sal. cin. 26 dead bran. of 05.2002 Bratek& ZB (2002) cf. Vib. opu. Tapesia kneiffii (Wallr.) J. Kunze 22 on bot. of upr. 04.2001 Bratek& ZB (2002) (s.n. T. retincola P. Karst.) dead stalkof Phr. aus. Verpa conica (O.F. MuÈ ll.) Sw. 19 ground 03.2002 Bratek& ZB (2002)

Tab. 4. ± Collection sites in sudds and their Registry Numbers (RN)

Country Region Locality (RN)

Hungary OÍ rseÂg Hills Lake Fekete 1 Valley of Kis Stream 2 Lake OÈ rdoÈg 3 Lake VadkacsaÂs 4 Lake SaÂs 5 DoÂ Forest 6 Lake at ApaÂtistvaÂnfalva 7 Lake BaÂrkaÂs 8 Borsod-AbauÂj-ZempleÂn County Lake Hangonyi 9 Lake at Muhi 10 Lake Kis Mohos at KelemeÂr 11 Lake Nagy Mohos at KelemeÂr 12 MaÂtra Mountains Lake at Egerbakta 13 Lake NyõÂrjes at Sirok14 Pest County Lake at CsoÈmoÈr 15 Bakony Mountains Lake Nagy at OÈ cs 16 Lake Monostori at SzentbeÂkkaÂlla 17 Lake at KovaÂcs Mountain at ZalaszaÂntoÂ18 Velencei Lake KereÂkWaters 19 AgaÂrd 20 stagnant waters of RaÂckeve- SzigetcseÂp 21 SoroksaÂri-DunaaÂg

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Tab. 4. ± Continued

Country Region Locality (RN)

Dunaharaszti 22 SzigetszentmikloÂs 23 Szigetbecse 24 Szabolcs-SzatmaÂr-Bereg County NyõÂres Lake at Csaroda 25 Lake Vajai 26 Romania Eastern Carpathians Lake Lucs 27 Lake OÈ rdoÈg 28 Lake Mohos 29 VeresvõÂz Stream 30 Romania Eastern Carpathians BaÂlvaÂnyos 31 Slovakia Lower Tatras Vychodna 32 CÏ ingov 33 Henclova 34

Tab. 5. ± The plant species names of hosts and substrates and their abbreviations.

Names of plant species Abbreviations

Alnus glutinosa Aln. glu. Betula pendula Bet. pen. Betula pubescens Bet. pub. Betula verrucosa Bet. ver. Betula sp. Bet. Carex sp. Car. Frangula alnus Fra. aln. Mentha aquatica Men. aqu. Picea abies Pic. abi. Pinus sylvestris Pin. syl. Polytrichum commune Pol. com. Populus alba Pop. alb. Populus sp. Pop. Phragmites australis Phr. aus. Quercus petraea Que. pet. Salix aurita Sal. aur. Salix cinerea Sal. cin. Salix sp. Sal. Sphagnum sp. Sph. Thelypteris palustris The. pal. Typha angustifolia Typ. ang. Typha laxmannii Typ. lax. Typha sp. Typ. Viburnum opulus Vib. opu.

Tab. 6. ± Distribution of collections according to months authors and collections.

Months I II III IV V VI VII VIII IX X XI XII Sum.

Basidiomycetes in 0000365192804280 282 Sphagnum sudds Basidiomycetes in 0001105913171840 77 non-Sphagnum sudds Ascomycetes in sudds 00348149181660 69

Tab. 7. ± Distribution of collections according to seasons.

Seasons Winter Spring Summer Autumn Sum.

Basidiomycetes in Sphagnum sudds 0 3 149 130 282 Basidiomycetes in non-Sphagnum sudds 0 11 27 39 77 Ascomycetes in sudds 0 15 24 30 69

Discussion The special importance of floating islands The quantity of the sudds in Europe has considerably decreased since the large-scale regulation of natural waters. The surface waters and watersides that have been straightened, dredged and deprived of floating islands for the sake of unobstructed sailing and flood con- trol give technocrats an impression of a cultivated and well-ordered landscape. The decay and the dissolution of the rest of the sudds are accelerated by the ever growing pollution of the waters (SzaboÂ 1999). It is a vicious circle: the decreasing sudds are natural water cleaners themselves and they are less and less capable of eliminating the ever increasing pollution of living waters. The floating islands are changing and varied pseudo-lands. Especially in years and seasons with extreme precipitation it is gen- erally dangerous or impossible to walkon sudds, and perhaps that is why they are less visited by mycologists, and there is only a limited number of scientific papers on describing mycota. There are floating islands where the water has evaporated from the waterbed because of the extremely dry weather, its peat has crumbled, and it is only the fungus samples of the herbaria that keep the memory of the former habitat, e.g. RN4, 15 and 20 habitats (Table 4). The exploration of the mycoflora of the large unexplored wetlands in Canada (Red- head 1989) and other parts of North America is delayed. The explorer is overcome by the inconceivably large territories as well as

the harsh field conditions and some technical problems during the identification of fungi (Roberts et al. 2004). Therefore, the protection of sudds as varied but unique habitats has not come into the limelight properly. Terrestrial peatlands and floating islands are rarely differentiated and valued in the international literature on wetlands-mires-peatlands. Consequently, there might be sudd data in the fungi literature of all three areas but they have not been separated from the terrestrial mire data (e.g. Kotlaba & KubicÏka 1960). The development of terrestrial mires means the encroachment of grounds, whereas floating islands gradually decrease the mass of water (KaÂdaÂr 1965). The concept that sudds are habitat types distinctively different from terrestrial mires is reinforced by the following fact: these fungal species have occurred exclusively in the milieu of floating mats in the studied area so far: Cortinarius tubarius, C. uliginosus, Hygro- cybe coccineocrenata, Inocybe lacera var. helobia, Omphalina gerardiana, Russula betularum and R. laccata (Albert et al. 2004). Floating islands as rapidly decaying habitats are not only capable of cleaning living waters, but are also crucial refugia for many rare organisms (Balogh 1980, 1983, 2001b). The diversity of sudd fungi is largely dependent on the quality and quantity of vegetation. According to the General Habitat Clas- sification System (G-NHCS) the formal name of our studied habitat is floating fen (identification code: B1). Floating fens can only belong to marshes of Phragmites or Typha beds category (Fekete et al. 1997); CORINE-code: 53.11 and 53.13 (according to CORINE Bio- topes, Natura 2000, Biosphere Reserves Integrated Monitoring Sys- tem). The latest phytosociological account of Hungarian vegetation contains only six types of floating fens in the marsh and moor vegetation category: Betulo pubescenti-Sphagnetum recurvi, Cicuto-Car- icetum pseudocyperi, Phragmitetum communis, Salici cinereae- Sphagnetum recurvi, Sphagno squarrosi-Alnetum, Thelypteridi- Typhaetum angustifoliae (Borhidi 2003). By virtue of the law on Nature Conservation in Hungary [Article 23(2), Act N8 LIII of 1996] all mires ± sudds included ± are protected in Hungary. In contrast to the low numbers in the formal categories, sudds support a high diversity of habitat types and plant assem- blages (Balogh 2000a, b; 2001a). The floating fen with a few types of plant association is forced into the official category of habitat. The similarities of the two sudd types are reinforced by the fact that the orders, families and genera that have been found in both types overlap considerably. However, it does not describe the uniqueness and much greater variety of the floating island as a habitat and its diversity or the state of succession. Consequently, it is not covered by conservation legislation. The

defining role of specific habitats (e.g. plant association) is high- lighted by the fact that the species pools of the Sphagnum sudd and the non-Sphagnum sudd basidiomycetes were congruent only in three identical species: Cortinarius uliginosus, Psathyrella typhae and Russula laccata (Albert et al. 2004). The distribution of the studied sudd fungi in the examined region shows similarity to findings of Redhead (1989). The habitats we studied are characterized by the significant similarity of plant associations, cool climate and balanced cold soil temperature. The high number of summer collections in the present publication can be explained by the cool habitat's balanced climate. According to Red- head's observations the distribution of fungi seems to be wider than that of flowering plants. This fact has been verified by our own observations. Hygrocybe coccineocrenata (Boertmann 2000) ± a typical boreal species ± appeared as a first record from Hungary in two sudds (RN1, 5) in two successive extremely dry years (2002, 2003). Published data on wetlands refer to Cantharellula umbonata, Galerina paludosa, Marasmius epiphyllus, Phaeomarasmius erinaceus, Psathyrella typhae (Redhead 1981, 1989), Cortinarius uliginosus (Bujakiewicz 1993), Galerina paludosa and G. tibiicystis (Gulden & HallgrõÂmsson 2000) as characteristic species in circumboreal taiga and subarctic wetlands. The majority of basidiomes living in sudds can be considered as typically montane or boreal species hence they are rare in the Carpatho-Pannonian Region. The specifically non- zonal sudd habitats may allow niches of cryophilic fungi.

The protection of the habitats and the fungi of floating islands The European environmentalist pacts have not formulated the protection of fungi yet (BoÍsze & Fodor 2005). The experts of the ECCF (European Council for the Conservation of Fungi) have been working on the construction of a list of fungal species that could be included in the Bern Convention as well as later in the addenda of EU directives for habitat protection (Dahlberg & Croneborg 2003) since 1991. 15 out of the Member States (EU 25) have an official red list of fungi, 14 of them have fungal species protected by laws (Siller et al. 2006). The previously proposed red list of Hungarian fungi (RimoÂczi et al. 1999) includes 118 genera, 280 species and 5 sub- species, of which only 35 fungi species have gained official protection so far (Siller et al. 2005). Three of these 35 species occur mostly in sudds in the Carpatho-Pannonian region: Lactarius helvus, Lec- cinum variicolor and Russula claroflava. Out of the three countries covered in this paper, it is Romania that has not taken protective measures so far. In Slovakia the practical protection is legally regu-

lated. Although there are 94 fungal species protected by law or reg- ulations, no sudd fungi are included (ECCF 2006). Floating islands may be good indicators of habitat quality, especially indicating stress caused by civilization. The floating mats are exposed to extremely high level of stress; at edges of ponds, lakes and rivers; expansion and contraction of a peat body owing to changes in groundwater storage; stress from tidal inundation; freeze- thaw cycles under permafrost or seasonal frost; shear stresses from lateral water flow; atmospheric pressure changes and ebullition

fluxes of methane (CH4) bubbles formed deep in the anoxic peat (Rydin & Jeglum 2006). Conservation is of vital importance as many species that can be potential gene banks are on the verge of extinction. There is a need for priority ranking of sudd habitats and species so that we can plan and implement conservation programs for their protection (Balogh 2003b). Such programs aiming at enhancing full protection of the quality and quantity of sudds can only be implemented if international cooperation is established, because it is important to remem- ber that rarity is a consequence of ecological factors, not a cause (Doust & Doust 1995). With this study from Europe we would like to draw attention to the importance of protecting these habitats and to the gradually increasing number of identified species they contain that have been vanishing worldwide.

Acknowledgments We would like say thanks to Katalin HajduÂ for helping with the English translation of the text, and ZoltaÂn IllyeÂs for helping with the map of collection sites. We are grateful to Dr. Tibor StandovaÂr for supervising the technical terminology on conservation.

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(Manuscript accepted 27 April 2009; Corresponding Editor: R. PoÈder)

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