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Dudka I.O. Xylophilous fungi and fungi-like organisms in the virgin and oldaged beech forests of the Ukrainian Carpathians. The results of study of the xylophilous mycobiota species diversity in the virgin forests and old-aged beech forests presented in one reserve and two national nature parks of Ukrainian Carparthians (Carpathian Biosphere Reserve, national nature parks Zacharovanyi Krai (Bewitched Land) and Synevyr) are given. At first in the virgin beech forests observations on the species composition and distribution of wood-destroying fungal representatives from class Ascomycetes (group of discomycetes) that associated with the dead wood both trophically (active destructors) and topically (inhabitants on the dead trunks and fallen branches of beech and subdominant tree species of different size) were held. The analysis of distribution in the virgin beech forests of fungi-like organisms from class Myxomycetes was realized. The observations on the rare species in slime molds biota were held in dependence of the level of anthropogenic impression on certain localities. The review of literature data concerning species diversity of wooddestroying aphyllophoroid fungi revealed in the Uholka virgin beech forests (Carpathian biosphere reserve) was made.

.. . ( , ) Ascomycetes ( ), ( ), ( - ). Myxomycetes. . , ( ).

The dead wood old stumps, trees killed by insects and damaged by various fungal and bacterial diseases but remained to stand, fallen trunks, branches and twigs forms one of the richest microbiotopes for biological 116 diversity in forest ecosystems. The dead wood together with associated species diversity plays the key role in supporting of forest productivity and stable development and in maintenance of its ecological balance. This microbiotope is especially important for the organisms which are tightly connected trophically and/or topically with wood substrates: insects, fungi, lichens, mosses etc.

In spite of the tremendous meaning of the dead wood its volumes in economical forests are extremely low. The dead wood in managed forests of Europe fluctuated from 0,6 m/ha in Austria to 12 m/ha in Switzerland. Only protected virgin or old-aged natural forests guarantee securing of the dead wood volumes necessary for support of biological diversity in forest ecosystems at the high level. In Carpathians there are numerous protected areas which embrace the virgin beech forests. The volumes of dead wood in the reserves and nature parks with the virgin beech forests increase sharply in comparison with the managed ones. For example, it is known that in the virgin beech forests of the national park Semenik-Cheile-Karasului (Romania) accumulated mass of the dead wood varies in the ranges 78121 m/ha. In another European virgin beech forest Haveshova (nature park Poloniny, Slovakia) upon the average 121 m/ha of dead wood remains are collected. It is considered that the highest volumes of the dead wood for arpathians are concentrated in the Uholka virgin beech forest (Carpathian biosphere reserve, Ukraine): here the content of woody remains composes 200(300) m/ha (Commarmot, Bachofen, Bundziak et al., 2003).

This important component of reserve ecosystems provides the numerous forest inhabitants with food substances and specific biotopes, creating the favorable conditions for high and stable species diversity.

First of all it concerns so called xylophilous or xylobiont organisms whose development and further existence are impossible without woody substrates. The representatives of xylobiont ecological group are wide-spread among the fungi and fungi-like organisms. Unfortunately the investigation of ecosystems species composition, participation of living organisms in redistribution of substance and energy on different trophic levels are in the first turn limited with producers and consumers that is plants and animals. Numerous reducers in particular fungi which realize destruction of organic matter including the dead wood dont predominantly attract attention of researchers. In comparison with plants fungi from the point of view of their study may be considered as real outsiders.

Meanwhile it is known that fungi are one of the biggest group of organisms.

Today almost 100 000 fungal species are described (Dictionary of the Fungi, 2008); prognostic value of fungal species diversity is fluctuated from 200300 thousands (, , 1995) to 1, 5 millions of species (Hawksworth, 1991). In forest ecosystems there are many xylobiont fungal species that belong to the wood-destroying organisms taking part in destruction of lignin and cellulose of plant origin (, 1993).

Wood-destroying fungi are widely distributed in Carpathian forest ecosystems, including the virgin and old-aged beech forests. Taking into account the presence of wood-destroying fungal xylobionts in many taxons of kingdom Fungi we have here reviewed representatives of Basidiomycetes and Ascomycetes. Both these classes of real fungi embrace species trophically and/or topically associated with the dead wood or with the wood of living trees. In addition we shall present the data on so called protozoan fungal analogues. These fungi-like organisms belong to class Myxomycetes that forms part of kingdom Protozoa (Mycetozoa). Relations of myxomycetes with the dead wood are topic in main but the development of generative stage of the slime molds in the absence of wood are impossible.

In the class Basidiomycetes the majority of wood-destroying species (75 % from all known species) is referred to aphyllophoroid fungi. Only 23 % species are the members of agaricoid fungi (, , 1979).

Species composition of aphyllophoroid fungi in the virgin beech forests of Ukrainian Carpathians was carefully studied in the forest massif Uholka (Carpathian biosphere reserve) in the frame of common project between the reserve and Swiss Federal Research Institute of Woods, Snow and Landscapes (, , 2006). The value of obtained data consists in the use of constant experimental plots for collection of fungal fruit bodies (basidiomata) and its combination with the results of route mycological observations at the territory of forest massif Uholka. In this famous virgin beech forest species composition of aphyllophoroid fungi turned out to be rather poor in comparison with the same composition of the economic beech forest in Sihlwald (Swiitzerland): in Uholka only 55 species of aphyllophoroid fungi.were revealed against 85 ones in Sihlwald. One more regularity was established for the Uholka virgin beech forest: low percent of aphyllophoroid fungi belonging to the ecologic group of xylotrophes (37 %, 41 species from 74). Correspondingly, in Sihlwald part of xylotrophes was significantly higher (67 %, 65 species from 74). It should be noted that Ganoderma applanatum (Pers: Wallr.) Pat., Bjerkandera adusta (Willd. et Fr.) P. Karst., Trametes versicolor (L.: Fr.) Pilt are the most wide-spread destructors of Fagus sylvatica L. dead wood in Uholka and Sihlwald. Fomes fomentarius (Fr.) Gillet joined those species as strongly distributed on the beech wood but only in Uholka. In Sihlwald F. fomentarius was not revealed. It is necessary to add that during our mycological observations in the supposed virgin beech forest Kamianka (national nature park Bewitched Land, 2011), Kvasovets and Krasna (national nature park Synevyr, 2013) we have repeatedly observed F.

140 .

fomentarius fruit bodies on dead beech trees that remain to stand, on fallen trunks in different stages of decay.

Except fungal destructors of the dead wood in the Uholka virgin beech forest some parasitic aphyllophoroid fungi infecting the living trees with the following development of heart or root rots were noted, among them Phellinus igniarius (L.: Fr.) Qul., Fomitopsis pinicola (Sw.: Fr.) P. Karst., Laetiporus 118 sulphureus (Bull.: Fr.) Murrill, Pleurotus ostreatus (Jacq.: Fr.) P. Kumm., Stereum hirsutum (Willd.) Fr. and others. On the whole it is proposed to consider the species composition of aphyllophoroid fungi in the virgin beech forests as normal for regions with protected vegetation and minimal anthropogenic stress (, , 2006).

In the class Ascomycetes a number of xylobiont fungal species is concentrated in the morphological group of discomycetes. This group is an assemblage of ascomycetes with the apothecia: cupulate, discoid or saucershaped fruit bodies. Some ideas on the participation of discomycetes associated with the dead wood in the local mycobiota of this fungal group may be received from literature data. So in the coniferous and deciduous forests of Russian Far East 640 species of discomycetes are known; 277 species of them belong to the ecological group of wood-destroying fungi (, 1998, 1999). It means that in the similar forests of Ukrainian Carpathians (the virgin beech forests with inset of Abies alba Mill.) not less than one third of discomycetous species will be connected with woody substrates. We begun to study of discomycete species diversity in the supposed virgin beech forests Kamianka (national nature park Bewitched Land, 2011), Kvasovets and Krasna (national nature park Synevyr, 2013). At the first stage of mycological observations 124 species of discomycetes associated with the dead wood were collected and identified. The wood-destroying species Mollisia cinerea (Batsch) P. Karst., M. melaleuca (Fr.) Sacc., Bisporella citrina Korf et S.E. Camp., Chlorociboria aeruginosa (Oeder) Seaver ex C.S.

Ramamurthi, Korf et I.R. Batra, Scutellinia scutellata (L.) Lambotte, Peziza badia (Hedw.) Fr. have dominated in accordance with the frequency of its occurrence on all investigated plots of the virgin beech forests.

During the research of fungal species composition in the virgin beech forests of Ukrainian Carpathians we paid attention on the distribution of the wood-destroying basidiomycetes and ascomycetes at the woody substrates of various sizes. It is found the most number of the wood-destroying basidiomycetes develops its basidiomata on tree trunks (diameter 30cm 1m and more) and fallen branches of trees with middle diameter 10 20 cm. The wood-destroying basidiomycetes dont use thin branches and twigs as substrate since they are not able to realize full cycle of development and to form basidiomata on woody remains of small size. While discomycetes as against the wood-destroying basidiomycetes colonize both big trunks and thin twigs with diameter not less 3 mm forming its ascomata on them. It is considered the species of the wood-destroying discomycetes have distinct preference in relation of the dead wood of coniferous trees (, 2001). However, the first results of our observations in Ukrainian Carpathians dont support the regularity registered in the forest ecosystems of Russian Far East.

It should be noted some peculiarities in distribution of discomycetes on the dead wood in the virgin beech forests of Ukrainian Carpathians.

Discomycetes on this substrate may be divided between three subgroups in dependence of state and origin of woody remains. The first subgroup with the most number of discomycete species includes those ones that are habituated to the dead wood on different stages of decomposition.

The species belonging to this subgroup are characterized with wide ecological valence: some of them develop ascomata not only on wood but on soil, leaf litter and even on the dry rests of herbs. The species connected with the wood show adaptation to the rotten wood and to the undestructed wood. Among the wood-destroying discomycetes of the Carpathian virgin beech forests there is a lot of species associated with the tree bark. The members of that subgroup has narrower ecological valence: sometimes it is possible to observe specialization in relation of the bark of certain tree species. The highest species diversity of discomycetes on the tree bark as it was demonstrated by our researches in the forest ecosystems of Ukrainian Left-Bank (, , .,

2009) is typical for Quercus robur L. the bark of which is covered with the deep folds and cracks. The low species diversity of discomycetes revealed in Carpathian virgin beech forests on the bark of Fagus sylvatica L. may be explained by smooth and flat structure of beech bark surface. The bark of silver fir (Abies alba Mill.) that often plays the role of subdominant in Carpathian virgin beech forests in spite of folded and fissured structure is not suitable for the development of discomycetes because of acid value pH. The third subgroup of the wood-destroying discomycetes includes the species developing on cones and seeds (fruits) of tree plants. This subgroup is the most insignificant concerning the number of species. Its representatives are differed from previous subgroups with the very limited ecological valence.

Investigations of species diversity of fungi-like organisms (class Myxomycetes) from Carpathian virgin beech forests were held in Carpathian biosphere reserve (2006) and in national nature park Bewitched Land (2011). In June and July 2013 the first collections of myxomycetes were made in the supposed virgin beech forests of national nature park Synevyr. It is necessary to point that participation of myxomycetes in the process of the dead wood utilization is not finally ascertained till now: there is opinion that slime molds feed on bacteria from woody substrates (, 1993;

Stephenson, Stempen, 1994). Therefore it is assumed to consider the mutual relations between myxomycetes and woody substrates as topic ones.

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