Macrofungi of the Bieszczady Mountains

The Bieszczady Mts, a mountain range in SE Poland, is a hot spot of fungal richness and diversity in Poland. This paper summarizes 5 years of studies in the Bieszczady Mts, as well as previously published research. A total of 1,377 macromycetes taxa were found, including many (464) which were protected, red-listed, or very rare in Poland. Thirty-eight taxa (nine Ascomycota and 29 Basidiomycota) have been reported in Poland for the first time: Agrocybe gibberosa, Auriporia aurulenta, Bolbitius variicolor, Bulgariella pulla, Chaetosphaerella phaeostroma, Clitocybe subspadicea, Clitopilus passeckerianus, Cortinarius anomalus var. subcaligatus, C. fervidus, C. flexipes var. inolens, C. sylvae-norvegicae, Cudoniella tenuispora, Entoloma bisporigerum, E. olorinum, E. poliopus var. parvisporigerum, E. sericeoides, Galerina caulocystidiata, Gymnopilus josserandii, Hymenoscyphus subferrugineus, Hypholoma olivaceotinctum, Inocybe queletii, Laccaria altaica, Lactarius romagnesii, L. rostratus, Mycena epipterygia var. atroviscosa, M. epipterygia var. candida, M. polygramma f. candida, Octavianina lutea, O. mutabilis, Pachyella violaceonigra, Panaeolus papilionaceus var. capitatocystis, Phaeocollybia jennyae, Psathyrella almerensis, Pyrenopeziza inornata, Scutellinia torrentis, Tricholoma basirubens, Tricholomopsis flammula, and Vibrissea decolorans. For all new taxa, short descriptions based on the collected material have been provided.


Study area
The Bieszczady Mts are a mountain range located in Poland, Slovakia, and Ukraine making up the western part of the Outer Eastern Carpathians. They are part of the Beskidy Mts, a series of the mountain ranges lying between the Czech Republic and Ukraine. The Polish Bieszczady Mts are classified as the Western Bieszczady Mts and are part of the mesoregion of the Beskidy Lesiste located in the eastern part of Podkarpackie Province [1,2]. The Polish part of this range occupies an area of ca. 1,450 km 2 and is bounded by Polish-Ukrainian and Polish-Slovakian borders in the south and east, respectively; the Łupkowska Pass, Osława valley, and the road between Szczawne and Czaszyn in the  feature for the Bieszczady Mts are the shrub communities with green alder (Fig. 4), Athyrio distentifoliae-Sorbetum alnetosum viridae, Pulmonario filarszkyanae-Alnetum viridis, Calamagrostis-Alnus viridis, or Salix silesiaca-Alnus viridis in subalpine areas. Hornbeam forests are rare and represented only by Tilio-Carpinetum; additionally, common alder communities are extremely rare. The nonforest plant communities in the Bieszczady Mts were strongly influenced by the pasturage of sheep, goats, and horses in the past. Shepherding has resulted in the formation of unique landforms in the subalpine zone (above 1,200 m a.s.l.), called polonynas (Polish: połoniny; Fig. 5), which are characteristic of the Eastern Beskidy Mts. Over 40 plant communities have been described in the subalpine zone of the Polish Bieszczady Mts. In the lower montane zone, over 50 types of nonforest plant communities are present. Of these, raised bogs (Oxycocco-Sphagnetea; Fig. 6), poor fens (Scheuchzerio-Caricetea), and poor meadows with Nardus stricta are the most valuable [3,5].
The soils of the Bieszczady Mts are derived from the Carpatian Flysch Belt rocks. The most common soils are eutrophic and dystrophic loamy brown earths (cambisols), shallow initial soils (leptospos, regospos), and soils characteristic of moist habitats, such as river banks (fluviosols), bogs (histosols), or meadows (gleysols) [3]. From a mycological point of view, the major soil characteristic influencing the fungal diversity is their pH. The soils of Bieszczady Mts are, in general, acidic or neutral (pH = 5.5-7.0 in water) with low concentrations of carbonates [Ca(Mg) CO 3 ]. This limits the number of fungal taxa, e.g., the calciphilous species of hypogeous or boletoid fungi do not occur in Bieszczady Mts.
Due to their unique character, and natural and cultural value, the Bieszczady Mts are protected. Over 290 km 2 are protected as the Bieszczady National Park (BdPN). Established in 1973, it has since been expanded a few times and is the third largest national park in Poland. The BdPN buffer zone is over 520 km 2 . BdPN covers the most valuable parts of the Polish Bieszczady Mts. In 1992, the East Carpathian Biosphere Reserve was   established by the UNESCO. It is a transboundary protection project that includes parts of Poland, Ukraine, and Slovakia. The entire Polish section is located in the Bieszczady Mts and is almost 1,090 km 2 large. It includes the BdPN and two landscape parks: Ciśniańsko-Wetliński Landscape Park and San Valley Landscape Park. The Bieszczady Mts are also protected as two Natura 2000 areas: Special Protection Area (Bieszczady SPA, PLC180001) and part of the Basin of the Upper San Special Area of Conservation (PLH180021) [6,7]. Fourteen natural reserves have been established in the Bieszczady Mts: Cisy na Górze Jawor, Gołoborze, Hulskie im. prof. Stefana Myczkowskiego, Krywe, Nad Jeziorem Myczkowieckim, Olsza kosa w Stężnicy, Olszyna Łęgowa w Kalnicy, Przełom Osławy pod Duszatynem, Przełom Sanu pod Grodziskiem, Sine Wiry, Śnieżyca wiosenna w Dwerniczku, Woronikówka, Zakole, and Zwiezło [6,8]. Moreover, seven landscape-nature complexes (Wieś Krywe, Młyn w Hulskiem, Cerkiew w Hulskiem, Cmentarz w Ruskiem, Młyn w Dwerniku, Wieś Smolnik, Cmentarz w Stuposianach) and 44 ecological sites have been established in the Bieszczady Mts [7].

Current state of mycological studies in the Bieszczady Mts
Highly diverse habitats and their largely natural character results an above average macromycete richness. The first mycological data from the Bieszczady Mts were published by Gumińska as a result of an 11-day trip of Polish mycologists (S. Czyżewska, S. Domański, B. Gumińska, M. Lisiewska, A. Nespiak, A. Skirgiełło, and W. Truszkowska) to the mountain range, in the vicinity of the village Wetlina [9]. The interesting results obtained from this trip inspired further studies by a group of scientists from various academic centers and specializing in different systematic groups. T. Majewski and W. Wojewoda joined the previous participants. Three excursions were made between 1960 and 1970, and their findings were published in four papers [10][11][12][13]. A preliminary red list of macromycetes of the Polish Carpathians was published in 1991, while a checklist of the corticioid fungi of this region was published in 1999 [14,15]. Both were based on earlier published data including those from the Bieszczady Mts. No further systematic mycological studies were conducted in the area of the Bieszczady Mts until 2008; however, some data have been published in many chorological or taxonomic papers and monographs (see references list). In 2008, Gierczyk and coworkers started a 5-year study of the mycobiota of the Bieszczady Mts. The first results (together with all available literature data) were published in 2009 [16]. In 2011, a protection plan for the BdPN was prepared by Kujawa et al.; however, it was not published. A short description of the characteristics of the mycobiota of the BdPN was prepared and published in the monograph on fortieth anniversary of the Park [17]. Selected results from this study have also been published by the participants of this project in papers on particular species or systematic groups [18][19][20][21][22][23][24][25][26][27]. Some species from the Bieszczady Mts have also been reported in the internet Database of Protected and Endangered Fungi (GREJ) (e.g., [28][29][30][31][32][33][34]). The diversity of lichenized or lichenicolous macromycetous fungal species (Lichenomphalia, Multiclavula, Tremella) have also been published in papers and monographs on the Bieszczady Mts lichen biota [35][36][37][38][39][40][41][42]. In last few years, two forestry science reports were published, containing lists of the observed fungi [43,44].
In the current paper, all available data on the Bieszczady Mts mycobiota have been reviewed and published, including the published reports and the authors' unpublished data.

Results
The complete list of macrofungi species found by the authors in the Bieszczady Mts together with the associated literature data has been supplied as supportive material (Appendix S1). Data is presented according to the following scheme: species name; threat category, species protection; LR: literature data (with synonymous names used by cited authors); PP: data included in the Bieszczady National Park Protection Plan (2011; unpublished); UP: unpublished data from the years 2008-2013. Notes: additional information and comments (doubts, misconceptions, errors, etc.); description of the taxa new to Poland.
For PP and UP records, localities, phenological information, habitats, and substrata have been reported. In the LR section, the paper by Gierczyk et al. [16] has been cited only if this paper provided original data.

Discussion
The available literature on the fungal diversity in the Polish Bieszczady Mts has been reviewed and the new data collected by the authors in the years 2008-2015 is presented. The current list of macrofungi species identified in the Bieszczady Mts includes 1,377 species, subspecies, varieties, and forms: 235 taxa of Ascomycota and 1,142 taxa of Basidiomycota (in the Bieszczady National Park 1,206 species: 196 of Ascomycota and 1,010 of Basidiomycota) comprising 30% of the total number of macromycetes known in Poland. A further 71 taxa are supported by dubious data (16 Ascomycota and 55 Basidiomycota). The number of taxa from the Bieszczady NP listed by Kujawa [130] is higher (1,304), because it includes some unpublished data on corticioid fungi collected by Dariusz Karasiński and some dubious data. This mycobiota richness makes the Bieszczady Mts the third richest region in macrofungi in Poland, after Białowieża Primeval Forest (ca. 2,000 known taxa) [131] and Kampinos NP (ca. 1,600 taxa) [132]. In comparison, 1,233 taxa of macrofungi have been reported from the Slovakian Poloniny National Park [133][134][135]. Unfortunately, the mycological data for Uzhanian National Park (Ukraine) are not available. The official website of the Nature Reserve Fund of Ukraine mentions a total of 66 fungal species from Uzhanian National Park [136]. Many taxa recorded in the Bieszczady Mts are very valuable (Tab. 1). Four species (Hericium erinaceum, Hydnellum concrescens, Corticium polygonioides, and Skeletocutis odora) are strictly protected and another 14 (Bondarzewia mesenterica, Clavariadelphus pistillaris, Gomphus clavatus, Hericium alpestre, H. coralloides, Inonotus obliquus, Morchella conica, M. esculenta, Ptychoverpa bohemica, Sparassis brevipes, Strobilomyces strobilaceus, Tremiscus helvelloides, Verpa conica, and Xerocomus pelletieri) are partially protected in Poland. Two hundred and ninety taxa recorded in the Bieszczady Mts are red-listed comprising 30% of all species in the Polish red list. In the current paper, 319 taxa not previously known to be present in the Bieszczady Mts are reported. They include 38 taxa (nine Ascomycota and 29 Basidiomycota) hitherto not known in Poland: Agrocybe gibberosa, Auriporia aurulenta (Fig. 7), Bolbitius variicolor, Bulgariella pulla (Fig. 8), Chaetosphaerella phaeostroma, Clitocybe subspadicea, Clitopilus passeckerianus ( Fig. 9), Cortinarius anomalus var. subcaligatus, C. fervidus, C. flexipes var. inolens, C. sylvae-norvegicae, Cudoniella tenuispora (Fig. 10 (Fig. 11), Pachyella violaceonigra, Panaeolus papilionaceus var. capitatocystis, Phaeocollybia jennyae (Fig. 12), Psathyrella almerensis, Pyrenopeziza inornata, Scutellinia torrentis (Fig. 13), Tricholoma basirubens, Tricholomopsis flammula, Vibrissea decolorans (Fig. 14). This makes Polish Bieszczady Mts a hot spot of fungal diversity, in both Poland and Europe. Sixteen taxa listed on the list of indicators of the nature value of beech forests in Europe [137] have been found in the Bieszczady Mts. This is higher than that reported for other beech forests of European importance including Poloniny National Park (Slovakia) [134,138], the forest with the hitherto highest value published (15). The number of the taxa designated as species of special interest (SSI), which are selected from wood-inhabiting fungi in European beech forests [139], is also high (62). Reported values of this parameter for other beech forests in Europe are lower varying from 23 (Poloniny National Park, Slovakia) to 51 (UNESCO World Natural Heritage Kellerwald-Edersee National Park, Germany) [134]. Many SSI species observed in the Bieszczady Mts are common within this area, e.g., Hericium coralloides or Dentipellis fragilis. Many other taxa selected by Slovakian mycologists as species of special interest during the study of beech forests in that country (e.g., Antrodiella pallescens or Perenniporia narymica) [140] have also been found in the Polish Bieszczady Mts. One should note that the evaluation systems proposed by Christensen et al. [137] and Ódor et al. [139] were based on Western and Southern European data, therefore need to be tested and adjusted for Central European forest conditions. The Bieszczady Mts are the only place where 77 taxa are found in Poland making it an important refuge of macrofungi. Another 114 taxa reported in the Bieszczady Mts are extremely rare in Poland and have only been found in three or fewer localities in this country. Its unique ecosystems (e.g., Alnus viridis communities or postpasture habitats) must be protected to preserve the macrofungi richness of this mountain range. The unusually high availability of forest biomass in the Bieszczady Mts region [141], the most important parameter determining the presence of many rare, wood-inhabiting fungi, should be especially emphasized. It is difficult to indicate the regions in the Bieszczady Mts with the highest mycological value. The most interesting findings have been made on the northern slopes, in humid stream valleys, e.g., Hylaty Valley or Górna Solinka Valley, and in places with high concentrations of large, dead tree trunks on the forest floor. One could expect further valuable findings, as many fungal groups (genera Russula and Cortinarius, corticioid and hypogeous fungi, small Ascomycota) are still undercollected and underrepresented on the current species lists. Moreover, although field work has been conducted throughout the Bieszczady Mts, the greatest emphasis has been placed on the study of the Bieszczady National Park mycobiota. In consequence, the northern and western parts of the Bieszczady Mts have only been cursorily studied. Analysis of data on the Poloniny NP has showed that 462 species of macrofungi known from the Slovakian part of the Eastern Carpathians have not been observed yet in the Bieszczady Mts. Some of these, for example fungi inhabiting old oak trees and logs (e.g., Buglossoporus quercinus), are unlikely to be collected in the Bieszczady Mts due to the absence of suitable habitats and substratum; however, most of these 462 taxa probably also grow on the Polish side of the border. Almost 1,840 macrofungi taxa are known from the Polish and Slovakian parts of the East Carpathian Biosphere Reserve.