ISSN 1995�4255, Contemporary Problems of Ecology, 2013, Vol. 6, No. 4, pp. 390–395. © Pleiades Publishing, Ltd., 2013.Original Russian Text © V.A. Vlasenko, 2013, published in Sibirskii Ekologicheskii Zhurnal, 2013, No. 4, pp. 507–514.

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In modern understanding, fungi are a polyphyleticgroup of osmoheterotrophic spores forming eukary�otes with a vegetative body looking like a coenoeciumor divided into cells filaments with a solid coat [1].

The object of study is polypores of a xylobioniccomplex growing on wood or on a product of itsdestruction. Polypores have carpophores with differ�ent forms and structures, from resupinate to attachedor differentiated to pileus and vellus. They all have atubular hymenophore.

Bracket fungi play an important role in processes ofwood destruction, providing the circulation of ele�ments and energy transformation in forest ecosystems[2, 3]. Some bracket fungi are agents of trunk and rootdotes in woody plants [4, 5]. Tree�destroying fungi arenecessary elements of various types of forest commu�nities, and they reflect general principles of forestdevelopment and its condition [6]. Many species areused as bioindicators for estimating the condition ofthe forest ecosystem [7, 8]. They are also actively stud�ied in relation to their application in medicine,because they synthesize a wide range of biologicallydifferent active compounds [9, 10, 11]. The aim of thisstudy was to show the ecological characteristics ofpolypore biota in the forest steppe of Western Siberia.

MATERIALS AND METHODS

The species composition was observed during thestudy, and ecological characteristics of bracket fungifrom a xylobionic complex in the forest steppe of West�ern Siberia were studied. Ground species of polyporesfrom genus Albatrellus and Boletopsis found in WesternSiberia were not found in forest steppe. Polyporusrhizophilus (Pat.) Sacc. growing on roots of cereals in

steppes as well as fungus with aculeiform hymeno�phore Irpex lacteus (Fr.) Fr. and ground fungus Coltri�cia perennis (L.) Murrill were excluded from the anal�ysis.

Studies were conducted according to the routemethod in the forest steppe of Western Siberia on theterritory of Altai krai and Novosibirsk oblast. The lit�erature also including data about fungi from the West�ern Siberian forest steppe [5, 12, 13, 14, 15], includingthe Transural region [3]. The characters of biotope,substrate type and its condition (wood species, alive ordead, and stage of wood destruction), and topic niche(living tree, standing dead trees, fallen trees, stub,strap, and root) were also registered when collectingfungi. A specimen that is an assembly of fruit bodiesformed at the unit of wood substrate by one vegetativebody of fungi was considered a registered unit. Thetotal number of observed fungi specimens was esti�mated by the number of substrate units. Russian and for�eign instruments, an ML�2A microscope and a CarlZeiss Stemi DV4 stereomicroscope, were used for thetaxonomic identification of fungi. Nomenclature wasused according to the Index Fungorum (http://www.indexfungorum.org/Names/Names.asp). TheSorenson–Chekanovskii (KSC) coefficient was used todetect similarities and differences in species composi�tion on various substrates. An analysis was conductedusing Microsoft Office Excel 2003 (databases,matrixes, and coefficient calculation) and STATIS�TICA 6 (cluster analysis). Cluster analysis was con�ducted using consolidation rules according to theWard method. The minimal dispersion was optimizedwithin clusters and almost equal clusters form as aresult. The squared Euclidean distance was used as ameasure of closeness. The reliability of statistical data

Ecological Characteristics of Bracket Fungi in the ForestSteppe of Western Siberia

V. A. VlasenkoCentral Siberian Botanic Garden, Siberian Branch, Russian Academy of Sciences,

ul. Zolotodolinskaya 101, Novosibirsk, 630090e�mail: vlasenkomyces@mail.ru

Abstract—Bracket fungi (Polypores) of the forest steppe in Western Siberia were studied and their ecologicalcharacteristics were determined. The xylobiotic complex is presented with 128 species of bracket fungi; 120species develop on the main forest�forming woody plants. The distribution of bracket fungi species on plantspecies forming the substrate was studied and peculiarities of species distribution, their ratio on live trees andon dead wood, and the distribution of species in topic niches were analyzed.

Keywords: polypores, ecology of mushrooms, substrate specialization, forest steppe, Western Siberia

DOI: 10.1134/S1995425513040136

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was proven by controlling the clustering stability whenchanging methods.

RESULTS AND DISCUSSION

Wood plants play an important role in fungi life;they form the substrate where fungi grow. It connectsthem trophically (nutritive connections) and topically(interactions associated with habit area) to woodplants. Analyses of fungi species distribution on woodplants make it possible to determine ecological char�acteristics in certain natural zones and in presentedbiotopes.

One hundred and twenty�eight species of polyporesfrom the xylobionic complex have been found in theforest steppe of Western Siberia. One hundred andtwenty species were found on forest�forming woodplants. An analysis of the species composition of poly�pores found on the main wood plants in the foreststeppe of Western Siberia showed that it is more spe�cific on soft woods and forms a specific cluster (Fig. 1).Broadleaved trees form another cluster, which isdivided into several branches. The largest number offungi species (120 species) develops on the main for�est�forming species: Pinus silvestris L., Betula pendulaRoth., and Populus tremula L. and on dominants ofbottomland biotopes, underbrush, and brush commu�nities presented with species from genus Salix, Padus,and Sorbus. Forty�nine species of polypores are pre�sented on common pine in the studied region. Eighty�five species develop on broadleaved trees, 58 on birch,55 on aspen, 25 on willow, 24 on cherry, and 12 moun�tain ash.

Aspen and birch (KSC = 0.58) and willow and aspen(KSC = 0.48) have the highest degree of similarity inpolypore composition among broadleaved trees.Cherry and mountain ash stand separately (KSC =0.56). The species composition of fungi on commonpine is close to the one on birch (KSC = 0.19), whichcan be explained by the widespread distribution ofthese two wood species on the studied territory andtheir common occurrence in numerous biotopes.

Thirty�four species (26.56% of all species) belongto those observed at the studied territory only on com�mon pine: Gloeophyllum abietinum (Bull.) P. Karst.,G. odoratum (Wulfen) Imazeki, G. protractum (Fr.)Imazeki, Onnia tomentosa (Fr.) P. Karst., Phellinus vit�icola (Schwein.) Donk., Ph. weirii (Murrill) Gilb.,Porodaedalea pini (Brot.) Murrill, Antrodia gossypium(Speg.) Ryvarden, A. serialis (Fr.) Donk., A. xantha(Fr.) Ryvarden, Climacocystis borealis (Fr.) Kotl.et Pouzar, Fomitopsis rosea (Alb. et Schwein.) P. Karst.,Postia caesia (Schrad.) P. Karst., P. fragilis (Fr.) Jülich,P. leucomallella (Murrill) Jülich, P. ptychogaster(F. Ludw.) Vesterh., P. sericeomollis (Romell) Jülich,P. stiptica (Pers.) Jülich, Pycnoporellus fulgens (Fr.)Donk., Loweomyces wynneae (Berk. et Broome)Jülich, Dichomitus squalens (P. Karst.) D.A. Reid,Diplomitoporus crustulinus (Bres.) Doman�ski, D. flave�

scens (Bres.) Doman�ski, Leptoporus mollis (Pers.)Quél., Skeletocutis amorpha (Fr.) Kotl. et Pouzar,Trichaptum abietinum (Dicks.) Ryvarden, Trichaptumfuscoviolaceum (Ehrenb.) Ryvarden as well as knownfrom literature data Anomoporia bombycina (Fr.)Pouzar [12], Phellinus chrysoloma (Fr.) Donk. [12],Fibroporia vaillantii (DC.) Parmasto [12, 13], Ph. har�tigii (Allesch. et Schnabl) Pat. [13], Ph. ferrugineofus�cum (P. Karst.) Fiasson et Niemelä [14], andGloeoporus taxicola (Pers.) Gilb. et Ryvarden [15].

Among broadleaved trees, birch and aspen, whichare the main forest�forming species, have the mostspecific species composition of fungi. There are14 species (10.93%) that are found only on aspen:Inonotus radiates (Sowerby) P. Karst., I. rheades(Pers.) Bondartsev et Singer, Phellinus tremulae(Bondartsev) Bondartsev et P.N. Borisov, Oxyporuspopulinus (Schumach.) Donk., Antrodia macra (Som�merf.) Niemelä, Abortiporus biennis (Bull.) Singer,Radulodon aneirinus (Sommerf.) Spirin, Antrodiellaromellii (Donk) Niemelä, Ceriporia excelsa S. Lundellex Parmasto, C. reticulata (Hoffm.) Doman�ski,Cinereomyces lenis (P. Karst.) Spirin, Polyporus pseudo�betulinus (Murashk. ex Pilát) Thorn, Kotir. etNiemelä, Skeletocutis subincarnata (Peck) Jean Keller,and Aurantiporus alborubescens (Bourdot et Galzin)H. Jahn. [3]. Eleven species (8.59% of all species) arespecific for birch: Inonotus obliquus (Ach. ex Pers.)Pilát, Piptoporus betulinus (Bull.) P. Karst., Postia flo�riformis (Quél.) Jülich, P. hibernica (Berk. et Broome)Jülich, P. tephroleuca (Fr.) Jülich, Junghuhniapseudozilingiana (Parmasto) Ryvarden, Antrodiellahoehnelii (Bres.) Niemelä, Daedaleopsis septentrionalis(P. Karst.) Niemelä, Trichaptum biforme (Fr.)Ryvarden, Trechispora mollusca (Pers.) Liberta andPhellinus laevigatus (Fr.) Bourdot et Galzin [3]. Jung�huhnia lacera (P. Karst.) Niemelä et Kinnunen,

PineBirch

Aspen Bird cherryMountain ash.Willow

Fig. 1. Diagram of similarity in the species composition ofpolypores in the forest steppe of Western Siberia on forest�forming wood plants.

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Schizopora flavipora (Berk. et M.A. Curtis ex Cooke)Ryvarden (1.56% of all species) were observed only oncherry. Trametes ljubarskyi Pilät (0.78% of all species)was found only on aspen. No specific species werefound on mountain ash. During further studies, thepercentage of narrow specialized species can bedecreased in the case of their detection on new sub�strates, except for obligate specialized species such asPiptoporus betulinus, Polyporus pseudobetulinus, Phell�inus tremulae, and some others.

Bracket fungi were also found on trees and brushesless widespread or growing at the border with othernatural zones and in parks and botanic gardens fromgenus Caragana, Hippophae, Lonicera, Populus spp.,Crataegus, Ribis, Juglans, Tilia, Quercus, Siringa,Malus, Acer, Morus, Fraxinus, Abies, and Larix, butthe species composition of fungi on them in the foreststeppe does not differ with variety. It should be notedthat Laricifomes officinalis (Vill.) Kotl. et Pouzar, Gan�oderma valesiacum Boud., Trichaptum laricinum(P. Karst.) Ryvarden [12], and Bondarcevomyces taxi(Bondartsev) Parmasto [15] are specialized species onlarch. They are specific for mountain forests on thesouth of Western Siberia. According to the literaturedata, one species was found for fir tree: Tyromyceskmetii (Bres.) Bondartsev et Singer; it is described forthe forest steppe of the Transural region in WesternSiberia [3]. In other natural zones this species wasobserved only on broadleaved trees [15, 17, 18]. Fungispecies on honeyberry are Phellinus lonicerinus(Bondartsev) Bondartsev et Singer; on sea buckthornthey are Phellinus hippophaëicola H. Jahn. [14] andPhellinus robustus (P. Karst.) Bourdot et Galzin [12,14], which are parasites�specific for these plants andare not observed on other substrates. On other wood

species, fungi that were specific for them were notfound.

Species such as Phellinus igniarius (L.) Quél.,Ph. punctatus (Fr.) Pilát, Ganoderma applanatum(Pers.) Pat., Bjerkandera fumosa (Pers ex. Fr.) Karst.,Gloeoporus dichrous (Fr.) Bres., Cerrena unicolor(Bull.) Murrill, Fomes fomentarius (L.) Polyporusbadius (Pers.) Schwein., P. brumalis (Pers.) Fr., Pyc�noporus cinnabarinus (Jacq.) P. Karst., Trametes hir�suta (Wulfen) Lloyd., T. ochracea (Pers.) Gilb. etRyvarden, T. pubescens (Schumach.) Pilát, T. trogiiBerk., and T. versicolor (L.) Lloyd, which are able todevelop on various broadleaved substrates, and speciesgrowing both on pine and broadleaved trees such asGloeophyllum sepiarium (Wulfen) P. Karst., G. trabeum(Pers.) Murrill, Schizopora paradoxa (Schrad.) Donk.,Oxyporus latemarginatus (Durieu et Mont.) Donk,Fomitopsis pinicola (Sw.) P. Karst., Phaeolus schwein�itzii (Fr.) Pat., Postia alni Niemelä et Vampola, P. bal�samea (Peck) Jülich, Bjerkandera adusta (Willd.)P. Karst., Ceriporiopsis mucida (Pers.) Gilb. etRyvarden, and Heterobasidion annosum (Fr.) Bref.have high a degree of trophic plasticity among poly�pores of the forest steppe.

According to nutrition type, polypores from the xylo�bionic complex are divided into two trophic groups:xylosaprotrophs (developing on dead wood) and biotro�phs (growing on living trees) with different gradationsand switches [19]. For this reason, biotrophs are some�times divided into obligate and facultative [20].

The distribution of polypores on living and deadwood was analyzed for the main wood species of theregion, for which 120 species of polypores wereobserved. The species composition of polypores grow�ing on living and dead wood is clearly divided into twoclusters (Fig. 2) presented by groups of biotrophs andxylosaprotrophs, respectively. The cull wood of pine isan exception. Here, the species composition of poly�pores is close to that growing on living trees of com�mon pine. This tendency is associated with the highspecificity of pine wood as a substrate�forming specieswhich differs in having high acidity (pH = 5.6–6.1). Itdetermines the specialness of the species compositionof polypores, which is embodied in clustering. In gen�eral, 15 species of fungi can develop on living wood;117 species, including 12 species of xylosaprotrophswhich have biotrophic activity under certain condi�tions, can grow on dead wood. Among biotrophs, sixspecies belong to fungi growing on living pine: Onniatomentosa, Phellinus chrysoloma, Phellinus hartigii,Porodaedalea pini, Phaeolus schweinitzii, Heterobasid�ion annosum; five species grow on birch: Inonotusobliquus, Phellinus igniarius, Daedaleopsis septentrion�alis, Polyporus melanopus, Heterobasidion annosum;four species grow on willow: Phellinus igniarius, Laeti�porus sulphureus, Haploporus odorus, Polyporus melan�opus; two species grow on aspen: Phellinus tremulae,Oxyporus populinus; two species grow on bird cherry:

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dFig. 2. Diagram of similarity in the species composition ofpolypores in the forest steppe of Western Siberia on livingand dead wood of forest�forming plants. (l) Living trees;(d) dead wood.

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Heterobasidion annosum, Phellinus punctatus; and onespecies grows on mountain ash: Phellinus punctatus.

Principles of biotroph and xylotroph distributionby wood species and similarities in fungi species com�position on living and dead wood, in general, corre�spond to common factors of polypore species distribu�tion on the main woods of the forest steppe of WesternSiberia presented on Fig. 1, but with a slight preferen�dum. Thus, clusters were analogous according to sim�ilarities in fungi species composition on dead wood ofbroadleaved trees, while they differed little accordingto species composition on live trees. The species com�position of fungi on birch is close to that on willow(KSC = 0.44). Fungi growing on living aspen were notobserved on other trees. For this characteristic, theaspen cluster belongs to the isolated cluster of com�mon pine, although they have no common species.

It is known that the substrate is the main factorwhich limits the expansion of wood�destroying fungi[15]; among abiotic factors, temperature and humid�ity play an important role. However, the substrateshould be in suitable condition for the expansion ofone or another fungus species. For this reason, exceptfor the factor of trophic association and biotope char�acter, the character of the microenvironment deter�mining the fungi position and its connections withplant parts; the condition of tree, bark, and wood tis�sue; and the degree of their destruction are important.Topic connections between plants and fungi consist ofsubstrate formation by one species for another species,wherefore plants forming substrates play an important

role in fungi life, not only as a nutrition source, butalso as a microenvironment, because the vegetativebody of fungus, mycelium, is inside the substrate. Fruitbodies are necessary for spore spreading; their form,position of spore layer, and its modifications in differ�ent species, as well as the situation of basidiomas onthe substrate, are different. For this reason, its orienta�tion in space is also important. Association with themicroenvironment in polypores varies, and it appearsto be connected with trophic specialization, the way inwhich the substrate is populated, the stage of wooddestruction, the wide ecological amplitude of species,its biological characteristics, and its belonging to cer�tain seral group. Therefore, the topic niche determinesor excludes the possibility of one or another speciesexistence in biocoenosis due to their association withspecific microenvironments.

Several main topic niches can be separated forpolypores: trunks and roots of living trees, dry wood,strap, windfall, and stub. Individuals from differenttrophic groups, including both biotrophs andxylosaprotrophs, can be found on living wood. Biotro�phs on tree trunks (trunk parasites) destroying woodcore or saps are presented with such species as Phelli�nus chrysoloma, Ph. hartigii, Ph. igniarius, Ph. puncta�tus, Ph. tremulae, Porodaedalea pini, Phaeolus schwei�nitzii, Oxyporus populinus, Haploporus odorus, andLaetiporus sulphureus. Biotrophs are also observed onroots, including species such as Onnia tomentosa,Phaeolus schweinitzii, and Heterobasidion annosum,which are characteristic for conifers. Xylosaprotrophs

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Fig. 3. Diagram of similarity in the species composition of polypores in the forest steppe of Western Siberia in different microen�vironments. (l) Trunks of live trees, (d) standing dead trees, (w) fallen trees, (st) strap, (sb) stub, and (r) root.

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Polyporus melanopus and P. ciliatus, which havebiotrophic properties, are present on roots. Xylosapro�trophs occupy other topic niches.

The distribution of fungi species according tomicroenvironments varies depending on substrate�forming plants (Fig. 3). A high similarity in composi�tion of fungi species developing on roots and trunks ofliving trees due to the fact that these topic niches arepopulated with biotrophic active species destroyingaged, weaken, and wounded trees is the main featurefor all substrate�forming species. Biotrophs are thefirst among seral fungi in stage�by�stage changing aswood is destroyed. Fungi on strap form distinct grouppresented by xylosaprotrophs, but many of them pop�ulate dead wood or fallen substrate. The third group ispresented with xylosaprophits developing on deadwood, windfall, and stub. It should be noted that manyspecies from this group are observed on different sub�strates, but most species grow on windfall, most oftenon large trunks of forest�forming trees. Such species asPhellinus igniarius, Ph. punctatus, Fomitopsis pinicola,Ganoderma applanatum, Bjerkandera adusta, Daeda�leopsis confragosa, D. tricolor, Datronia mollis, Fomesfomentarius, Trametes hirsuta, T. trogii, and T. versi�color can develop in different microenvironments;they are also differ with high frequency and abundancewith a wide trophic association.

CONCLUSIONS

There are 128 species of polypores from the xylo�bionic complex observed in the forest steppe of West�ern Siberia. On the main�forest forming trees,120 species from genus Pinus, Betula, Populus, Salix,Padus, and Sorbus develop. Conifers differ with thehighest specificity of polypore species compositionamong trees. Forty�nine fungi species are observed inthe studied region on common pine among forest�forming wood; 34 species are specific. On broadleavedtrees, in general, 85 are species observed; 58 speciesare on birch, 55 species are on aspen, 25 species are onwillow, 24 species are on bird cherry, and 12 species areon mountain ash. Among broadleaved trees, birch andaspen have the highest species specificity. Fourteenspecies are observed only on aspen; 11 species arefound only on birch. Polypores on introduced woodand wood rare in the forest steppe in general differ withlow species diversity and are presented with widespread species.

Species of polypores such as Fomitopsis pinicola,Ganoderma applanatum, Fomes fomentarius, Phellinusigniarius, Ph. punctatus, Trametes trogii, and T. versi�color have a high degree of trophic plasticity in the for�est steppe. Highly specialized fungi species such asPhellinus lonicerinus on honeyberry or Ph. hippophaëi�cola on buckthorn are species�specific parasites andare not found on other substrates. Among the bracketfungi of the forest steppe, 15 species are able to growon living trees, 117 species develop on dead trees, and

12 species can also display biotrophic activity. Themain substrate�forming trees are divided into twogroups according to similarities in fungal species com�position: conifer and broadleaved trees. Common pineis similar to birch in the species composition of fungi,which can be explained by the widespread nature ofthese two wood species on the studied territory andtheir combined existence in numerous biotopes.

The distribution of fungal species according tomicroenvironments varies depending on substrate�forming plants and is associated with a complex of fac�tors such as trophic specialization, the way the sub�strate is populated, the stage of wood destruction, etc.Due to the fact that biotrophic active species populateliving trees, a high similarity in the species composi�tion of fungi on roots and trunks is observed. Thegroup of biotrophic species is the first among seralgroups changing stage by stage in the process of wooddestruction. Xylosaprotrophs populate other topicniches presented with dead wood, windfall, strap, andstub. Species populating various topic niches differwith high frequency, abundance, and wide trophicassociation.

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Translated by N. Barykina