Contribution to the knowledge of fungi of the Kampinos National Park ( Poland ) with particular emphasis on the species occurring in burnt places

The paper presents 32 species of macrofungi new to the Kampinos National Park, found during the studies on fire-damaged areas after the forest fires in 2015. Three species new to Poland were described and illustrated (Calycellina leucella, Exobasidium juelianum, and Gymnopilus decipiens). Four species from Polish red list of macrofungi have been recorded in the Kampinos National Park for the first time: Botryobasidium vagum (R), Geastrum coronatum (V), Helicogloea farinacea (E), Inonotus cuticularis (R). During the current studies 17 pyrophilous species new to the Kampinos National Park were found.


Introduction
The Kampinos Forest is located in central Poland, on the left bank of the Vistula River, west of Warsaw.Almost all its area is protected as the Kampinos National Park (Kampinoski Park Narodowy, KPN).KPN was founded in 1959 to preserve the unique complex of inland dunes and marshy areas.It covers over 38 500 ha, therefore it is the second largest national park in Poland.The characteristic landscape of KPN shows the mosaic of dune areas, covered by pine and mixed forest, intersected by a swamp belts occupied by alder and riparian forests or vast, wet meadows [1][2][3][4].The fungi of the Kampinos Forest have been studied for over 140 years.The first information about the mycobiota of this area was published by Berdau [5], who listed 67 fungal species from the vicinity of Warsaw including at least one from environs of Wólka Węglowa village (east part of KPN).Further scarce data were provided by Chełchowski [6,7] and Błoński [8].First systematic investigation of the fungal diversity in the Kampinos Forest was conducted by Rudnicka-Jezierska in the 1960s.This study was focused on the psammophilic species, inhabiting bare and overgrown dunes [9][10][11].Additional data have been provided by other scientist (e.g., [12][13][14][15][16][17]). At the end of the twentieth century, the mycobiota of KPN was known to consist of 415 species.Intensive field works were conducted in the area of KPN in the years 2012-2014, reported in the monograph of fungi of the Park [18].This work lists 1533 taxa (species, varieties, and forms) of macromycetes, found by the authors (1407 taxa) or mentioned in the previous literature (425 taxa).Although KPN has the highest fire hazard category in Poland and incidental and intentional fires have been quite common in its area [19], the post-fire fungi of KPN have not been studied until now.Hitherto, only two non-obligatory pyrophilous species, Rhizina undulata Fr. and Sphaerosporella brunnea (Alb.& Schwein.)Svrček & Kubička, have been reported from the Kampinos Forest [18,20].At the end of the spring 2015 (May 7 and June 4), two intentional arsons occurred in the central part of KPN.The surface fire took the area of 10.92 ha in the Laski Forestry (forest compartments No. 76 and 77).This fire-incident made an opportunity for additional study of mycobiota of KPN, with special emphasis on the post-fire fungi.The paper presents the list of the fungi new to KPN found on the burned area.Some interesting findings from other places of the Park are also included.

Material and methods
The burned area (10.92 ha) is located in forest compartments No. 76 and 77 of the Laski Forestry, in the east part of KPN, ca.2.7 km S from Palmiry village (Fig. 1).About half of it is protected as a part of the Sieraków Strictly Protected Area (OOŚS -Obszar Ochrony Ścisłej Sieraków) (Fig. 2), the rest is subject to active protection.It is covered

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The studies were conducted on 45 permanent plots (10 × 10 m) in the Laski Forestry.Eleven of them were located in the interior of the area destroyed by fire, i.e., represented the part of the area damaged by fire to the highest degree.These plots were selected randomly from the 15 most burned sites of the area.The other plots were located systematically along the borders of the burned area and formed 17 pairs.One plot of each pair was located in the burned area and were characterized by lower degrees of fire damage, while the second one was adjacent to it, but in non-burned forest representing the same plant community.Only a small part of the north edge of the burned area was excluded from the study due to protection zone of the black stork's nest.Additionally, the studies were also made by the route method, both on the burned area as well as in the other parts of KPN.The sporocarps were collected once in 2015 (November) and every month between April and October in 2016.The specimens collected were identified by the standard methods used in fungal taxonomy, using light-microscope and staining in ammonia, KOH, Congo Red, Melzer's reagent, cotton blue in lactophenol, brilliant cresyl blue (CRB), and sulfovanilin.The drawings of microcharacters were made on the basis of microphotographs, taken with Bresser MicroCam 5.0 digital camera and Bresser Science TRM 301 light microscope.All measurements were made directly through the light microscope under an oil immersion objective (×100).The spore dimensions were established from measurements of 50 randomly selected, well-formed spores (the deformed or atrophied spores were excluded from analysis).The 95% confidence intervals of the mean were calculated and the lower and upper values are given.For other structures, the extreme size values were presented, obtained after measuring 25 elements.Species descriptions were prepared based on all collected specimens.The fungal names were cited after the Knudsen and Vesterholt [21] and MycoBank [22].Polish distribution data were taken from checklists [23][24][25] and online database of Polish fungal literature [26].Threat categories (E -endangered, R -rare, V -vulnerable) in Poland were given according to "Red list of the macrofungi in Poland" (RL) [27].Dried specimens were deposited in fungaria of the Institute for Agricultural and Forest Environment, Polish Academy of Sciences (IŚRiL), Division of Mycology and Forest Phytopathology of the Warsaw University of Life Sciences -SGGW (WAML), and private fungarium of B. Gierczyk (BGF); each fungarium number represents different collection.

Results: list of species
During the current studies (2015-2016) in KPN, ca.220 species of macrofungi were identified.In this paper, 32 species hitherto not mentioned from this area have been presented.The most attention has been paid to the area damaged by surface fire in 2015.The arson in KPN resulted in the fire-damage of 10.92 ha of pine forest.On the burned sites, 21 species new to KPN have been recorded, among them 17 are pyrophilous fungi.Three species hitherto not recorded from Poland have been described.Four of the species identified are red-listed, while the other 12 species are currently known from less than 10 contemporary (found after 1945) localities.
Notes.Species hitherto mentioned from over 10 localities in Poland.
Notes.Species very rare in Poland, known only from the Długie Lake Reserve [55] and Cieszyn [56].

Agrocybe dura (Bolton) Singer
Species description.This fungus is a parasite of Vaccinium vitis-idaea.The infected plants form stunted, less stout shoots.Their leaves are somewhat rolled or wavy, smaller, softer, and thinner than that of healthy plants.All infected parts are light red or pinkish.The hymenium forms on the underside of the leaves as a whitish, thin, farinose layer.The (sub)cylindrical, somewhat sinuous, 25-50 × 5-6 μm, 4-6 sterigmate basidia grow out from between the cells of host cuticle.Basidiospores cylindrical, often irregularly curved, hyaline, smooth, 1-5 septate, (8)9-12( 14) × (2.2)2.5-3(3.2) μm, without iodine reaction.Conidia few, forming at the basidiospores ends, cylindrical, hyaline, smooth, 4-5 × 1-1.2 μm.Clamps absent.Exobasidium juelianum differs from E. vaccini (Fuckel) Woronin by the type of infection, which is systemic in the former species (all parts of the shoots are infected) and local in the latter (the fungus forms red galls on individual leaves).The spores and conidia of E. juelianum are somewhat shorter than that of E. vaccini.

Notes.
Species not rare in Poland, known from ca. 20 localities.

Discussion
The current studies increases the number of macrofungal taxa known from KPN up to 1565 (1374 Basidiomycota and 191 Ascomycota).Among the recorded taxa, there are a few species quite common in Poland (e.g., Colpoma quercinum or Mycena septentrionalis), which are not pyrophilous and are quite abundant in the studied area, but had not been found in KPN previously.This indicates that even intensive, 3-year-long field works are too short to complete the fungal species list of a certain area.Although in the previous studies conducted in KPN the post-fire habitats (e.g., places after campfires or trees damaged by lightning) were carefully examined for the presence of pyrophilous fungi, the list of these species included only two taxa [18].During the current studies, 19 species of fungi associated with charcoal and burned places were found, i.e., 17 new to KPN (Anthracobia macrocystis, A. melanoma, A. nitida, Coprinellus angulatus, Daldinia vernicosa, Geopyxis carbonaria, Gymnopilus decipiens, Inermisia ?fusispora,Lyophyllum anthracophilum, Peziza echinospora, P. pseudoviolacea, Plicaria carbonaria, P. endocarpoides, P. trachycarpa, Pholiota highlandensis, Psathyrella pennata, Trichophaea abundans) and both taxa mentioned earlier (Rhizina undulata, Sphaerosporella brunnea).Moreover, most of them produced numerous sporocarps, even thousands of specimens (Gymnopilus decipiens, Lyophyllum anthracophilum, Pholiota highlandensis, Psathyrella pennata, Rhizina undulata).Therefore, one could conclude that for the occurrence and development of these species, large-scale fires are essential.After the first year of study, the recorded number of post-fire fungi in KPN (19) is the highest compared to other published results from Poland.Adamczyk et al. [34] recorded 18 species on 20 old camp-fires (a 3-year-long study), 17 taxa were found on 10 burned places in the Gorce Mts [31] (over 2-year-long observations), while 16 species were found in the fire-destroyed part of Jelonka Reserve [75] (6-year-long observations).Results obtained on large-area fire places in managed forests have shown a distinctly lower diversity of pyrophilous mycobiota (species number between 8 and 13) in these ecosystems [33,58,76].These differences and uniqueness of KPN may be the result of a few factors.Firstly, the burned area is, in part, a long-term protected one as the Sieraków Strictly Protected Area and it is overgrown by a ca.200-year-old forest.Moreover, the KPN is very often damaged by fires of different range, ca.80 times every year (about half of all fires recorded in national parks in Poland).These give post-fire organisms a good opportunity to outlast and propagate.
Three species new to Poland were collected in the studied area.One of them are post-fire species, i.e., Gymnopilus decipiens.Next two species, Calycellina leucella and Exobasidium juelianum, are not associated with burnt places.A few of the species recorded are very rare in Poland, hitherto mentioned from single localities (Coprinopsis laanii, Helicogloea farinacea).Peziza pseudoviolacea has been hitherto mentioned from Poland only as a endophyte of roots of Picea abies [54].Current findings are the first observations of the apothecia formation by this species in this area.Lachnellula suecica was found on the new substratum (Pinus sylvestris), since the past findings from Poland were associated with Pinus mugo and Picea abies [23,53].
The studies on fire-damaged areas of KPN will be continued in the coming years and the results and analyses of changes in the fungal biota during the habitat regeneration will be published.

Fig. 1
Fig. 1 Localization of the study area in Kampinos National Park.Yellow circle -fire-damaged wood.

Fig. 2
Fig. 2 Localization of burned area in the Kampinos National Park.Red line -boarders of burned forest; green line -boarders of the Sieraków Strictly Protected Area (OOŚS).

Fig. 3
Fig. 3 Appearance of fire-damaged wood in forest compartment No. 77 of the Laski Forestry, the Kampinos National Park; April 27, 2016.Photograph by A. Szczepkowski.
Notes.Species rather common in Poland and known from over 40 localities.