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Turczaninowia 19 (1): 5-18 (2016) DOI: 10.14258/turczaninowia.19.1.1 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
УДК 582.284(470)
The Meruliaceae of Russia. I. Bjerkandera
I. V. Zmitrovich, M. A. Bondartseva, N. P. Vasilyev
Komarov Botanical Institute, BINRAS, Prof. Popov, 2, St. Petersburg, 197376, Russia, Е-mail: IZmitrovich@binran.ru
Key words, basidiomycetes, aphyllophoralean fungi, meruliaceous fungi, modifications variability, broadleaf wood-associated fungi, Russia, Saint Petersburg parks.
Summary. This paper opens a taxonomical survey on the genera of Meruliaceae (Polyporales, Basidiomycota) presented in Russian mycobiota. All the meruliaceous fungi represent an obligate component of heterotrophic block of forest ecosystems and considerable demanded biotechnological resource. The purpose of the present elaboration is a revision of East European and North Asian material on Bjerkandera genus highlighting its species' and intraspecific morphological variability and substrate specialization. The macroscopic descriptions are based on a study of fresh and dried specimens. The material of the herbaria of Komarov Botanical Institute (St. Petersburg, LE) and Institute of Zoology and Botany of Estonian Agricultural University (TAA) is studied. Micromorphological analysis is included the hyphal system revealing, the hyphae, basidia/basidiospores morphometry, and microchemical tests of the structures in question. The genus Bjerkandera is accepted in its original Karstenian sense, although the concepts by Pilat, Corner, Pouzar, and Zmitrovich et al. were discussed. The genus is characterized by two-layered context with rather loose tomentum and dense layer above the hymenophore, monomitic to pseudodimitic hyphal system, clamped generative hyphae, and ellipsoid-cylindrical basidiospores not staining in Cotton blue and Melzer's reagent. Only two species, Bjerkandera adusta and B. fumosa were recognized in the genus, and a possible position of B. subsimulans and B. terebrans was discussed, too. The polymorphism of B. adusta is exhaustively presented and the form tegumentosa was epitypified and described. The polymorphism of B. fumosa is also presented, and the formflavipora was correctly published and epitypified. The relationships between two species are discussed and the key for species delimitation is presented here. Distributional patterns are presented for both species as well as their substrate range. The substrates of B. adusta and B. fumosa in old-growth arboreta of Saint Petersburg are presented.
Ботанический институт им. В. Л. Комарова, БИНРАН, Проф. Попова, 2, Санкт-Петербург, 197376, Россия
Ключевые слова: базидиомицеты, афиллофоровые грибы, мерулиевые грибы, модификационная изменчивость, грибы широколиственных пород, Россия, парки Санкт-Петербурга.
Аннотация. Статья открывает таксономический обзор родов семейства Meruliaceae (Polyporales, Basidiomycota), представленных в микобиоте России. Мерулиевые грибы представляют облигатный компонент гетеротрофного блока лесных экосистем и значительный биотехнологический ресурс. Цель данной работы - таксономическая обработка восточноевропейского и североазиатского материала по роду Bjerkandera, с особым вниманием к описанию внутривидового полиморфизма. Макроскопические описания основаны на исследовании свежих и высушенных образцов, хранящихся в гербариях Ботанического института им. В. Л. Комарова (г. Санкт-Петербург, Россия, LE) и Института Зоологии и ботаники Сельскохозяйственного университета г. Тарту (Эстония, TAA). Микроморфологический анализ включал выявление гифальной системы, морфоме-трию гиф, базидий и спор и микрохимические тесты структур. Род Bjerkandera принят в его оригинальной трактовке П. Карстеном, хотя обсуждены также концепции А. Пилата, Э. Корнера, З. Поузара и И. В. Змитро-вича и соавторов. Род характеризуется двухслойной тканью с рыхлым верхним слоем и темной линией над
Поступило в редакцию 29.07.2015 Submitted 29.07.2015
Принято к публикации 06.12.2015 Accepted 06.12.2015
Мерулиевые грибы России. I. Род Bjerkandera
И. В. Змитрович, М. А. Бондарцева, Н. П. Васильев
гименофором, мономитической или псевдодимитической гифальной системой, пряжками на генеративных гифах и эллипсоидально-цилиндрическими базидиоспорами, не изменяющими окраски в реактиве Мельцера и Хлопчатобумажном синем. В роде принимается два вида - Bjerkandera adusta и B. fumosa. Таксономическое положение B. subsimulans и B. terebrans требует корректировки. Подробно описан внутривидовой полиморфизм B. adusta, причем произведена эпитипификация f. tegumentosa. Представление полиморфизма B. fumosa сопровождается корректным оформлением и эпитипификацией f. flavipora. Подробно обсуждена проблема разграничения двух видов и представлен видовой ключ. Собран материал по географическому распространению двух видов и их субстратной приуроченности, включая полученные авторами данные о субстратной приуроченности видов, выявленной в старовозрастных посадках широколиственных пород на территории г. Санкт-Петербурга.
This paper opens a taxonomical survey on the genera of Meruliaceae (Polyporales, Basidiomycota) presented in Russian mycobiota. The Meruliaceae family unites a certain corticioid (Zmitrovich, 1997), some polyporoid (Binder et al., 2013) and even lentinoid (Zmitrovich, Malysheva, 2013) fungi associated to the wood in various stages of humification and causing a white rot. All the meruliaceous fungi represent an obligate component of heterotrophic block of forest ecosystems and considerable demanded biotechnological resource.
The genus Bjerkandera was described by P. Karsten (1879) for annual tyromycetoid polypores characterized by colored pore layer which is separated from the context by a dense zone. The generic name refers to Clas Bjerkander (1735-1795), a Swedish naturalist. During a long time, the genus kept two species, B. adusta (Willd.) P. Karst. and B. fumosa (Pers.) P. Karst., but initially it contained also B. dichroa (Fr.) P. Karst. [= Gloeoporus dichrous (Fr.) Bres.], B. amorpha (Fr.) P. Karst. [= Skeletocutis amorpha (Fr.) Kotl. et Pouzar], B. kymatodes (Fr.) P. Karst. (= Skeletocutis amorpha), B. diffusa (Fr.) P. Karst. (an ambiguous species), and B. isabellina (Fr.) P. Karst. (a synonym of B. adusta in current use). Ames (1913) and later Donk (1974), who have adhered two-species concept of the genus, wrote about the distinct layer which precedes a tubes formation and that this layer is highly characteristic.
Further, various authors included the type B. adusta (and, therefore, the genus as a whole) into such genera as Gloeoporus Mont. (Pilat, 1937; Corner, 1989), Tyromyces P. Karst. (Pouzar, 1966) or Grifola Gray (Zmitrovich et al., 2006).
The reasons for inclusion of Bjerkandera type to the Gloeoporus were two-layered nature of tyromycetoid basidiocarps and a certain degree of gelatinization of hymenophoral tissues. However, as it was noted by Corner (1989), the degree of tube gelatinization is rudimentary in G. adustus (Willd.) Pilat and G. fumosus (Pers.) Pilat - in contrast to G. dichrous and some tropical genus representatives.
The reasons for merging of Bjerkandera into Tyromyces were a monomitic hyphal system in both genera and rather comparable tyromycetoid morphotypes. Further, it was shown that the hyphal system in tyromycetoid fungi is rather diverse and that Bjerkandera basidiocarps have a morphotype intermediate between grifoloid and tyromycetoid ones, therefore, both B. adusta and B. fumosa were moved into the Grifola.
A recent molecular testing of polypore genera confirms a monophyletic nature of B. adusta/B. fumosa grouping as a sister lineage of indigo-colored corticioid Terana coerulea (Lam.) Kuntze (Floudas, Hibbett, 2015). A great work on corrections of sequences misidentifications in Bjerkandera adusta/B. fumosa pair was provided by Jung et al. (2014).
The purpose of the present elaboration is a revision of East European and North Asian material on B. adusta and B. fumosa highlighting their intraspecific morphological variability and substrate specialization.
Materials and Methods
The macroscopic descriptions were based on a study of fresh and dried specimens. The materials of the herbaria of Komarov Botanical Institute (St. Petersburg, Russia, LE) and Institute of Zoology and Botany of Estonian Agricultural University (Estonia, TAA) were studied. Microscopic preparations were mounted from dried material in Melzer's solution, 10 % ammoniacal Congo Red and 5 % aqueous solution of KOH, using a LOMO Micmed-6 light microscope. The hyphal system was revealed and described according to updated technique (Zmitrovich et al., 2009). The size of mature spores was measured on 30 spores in distilled water and Melzer's solution.
Results and Discussion
Meruliaceae Rea, 1922, British Basid.: 620.
Phanerochaetoideae (Jülich) Parmasto, 1986, Windahlia, 16: 17.
Bjerkandera P. Karst., 1879, Medd. Soc. Fauna Fl. Fenn., 5: 38.
= Myriadoporus Peck, 1884, Bull. Torrey Bot. Cl., 11, 3: 27.
Basidiocarp annual or wintering, pileate, decur-rent to prostrate, soft to pliable. Upperside when present subtomentose, matt or rugulose at drying, hygrophanous. Context two-layered with rather loose tomentum and dense layer above the hymeno-phore. Hymenophore as a single tube layer of ce-raceous consistency. Hyphal system monomitic in the tubes and pseudodimitic in the context. Generative hyphae with clamp connections. Cystidia none. Basidia clavate with median constriction, 4-spored, with a basal clamp. Basidiospores ellipsoid-cylin-dric, smooth, thin-walled, negative in Melzer's reagent. Worldwide distributed oligotypic genus associated to the white-rot of hardwoods, rarely conifers.
Type species: Polyporus adustus Willd.: Fr., 1821, Syst. Mycol., 1: 363 (selected by Murrill, 1903).
Type specimen is deposited in Friesian herbarium of the Uppsala University Museum of Evolution (Sweden, UPS) marked as "Finland, Mustiala, leg. P. Karsten" (Ryvarden, 1991).
Anamorph: Geotrichum-like (Romero et al., 2007).
In Eurasian continent the genus contains two polymorphic species characterized below. In Americas, two additional species, Bjerkandera atroalba (Rick) Westphalen et al. and B. centroamericana Westphalen et al. (Westphalen et al., 2015), plus some ambiguous Murrill's taxa were reported.
Key to species
1. Pores 6-7 per mm, pore surface combines whitish (tube mouths) and smoky-gray to grayish-black tinges, dense zone above tubes is grayish-black.................................................... 1. B. adusta
- Pores (1)2-4(5) per mm, pore surface combines whitish (tube mouths) and buff to isabelline tinges,
dense zone above tubes is brownish-cinnamon ......
.............................................................2. B. fumosa
1. Bjerkandera adusta (Willd.: Fr.) P. Karst., 1879, Medd. Soc. Fauna Fl. Fenn. 5: 38. = Boletus adustus Willd., 1787, Fl. Berol. Prodr.: 392. -Polyporus adustus Willd.: Fr., 1821, Syst. Mycol., 1: 363. - Leptoporus adustus (Willd.: Fr.) Quel., 1886, Enchir. Fung.: 177. - Polystictus adustus (Willd.: Fr.) Gillot et Lucand, 1890, Bull. Soc. Hist. Nat. Autun, 3: 173. - Gloeoporus adustus (Willd.: Fr.) Pilat in Kavina et Pilat, 1937, Atlas Champ.
Eur., 3: 137. - Tyromyces adustus (Willd.: Fr.) Pou-zar, 1966, Folia Geobot. Phytotax. Bohemoslov., 1: 370. - Grifóla adusta (Willd.: Fr.) Zmitr. et Maly-sheva in Zmitr., Malysheva et Spirin, 2006, Mycena, 6: 21.
= Boletus fuscoporus J.J. Planer, 1788, Ind. Pl. erfurt. Fung. add.: 26.
= B. pelloporus Bull., 1791, Hist. Champ. France: 365.
= B. carpineus Sowerby, 1799, Col. Fig. Engl. Fung. Mushr., 2: pl. 231.
= B. crispus Pers., 1800 ("1799"), Observ. Mycol., 2: 8.
= B. concentricus Schumach., 1803, Enum. Pl., 2: 387.
= Poria argentea Ehrenb., 1818, Sylv. Mycol. Berol.: 31.
= Boletus isabellinus Schwein., 1822, Schr. Naturf. Ges. Leipzig 1: 96.
= Polyporus murinus Rostk. in Sturm, 1838, Deutschl. Fl., 3, Abt. 4: 117.
= P. subcinereus Berk., 1839, Ann. Nat. Hist., Mag. Zool. Bot. Geol., 3: 391.
= P. halesiae Berk. et M.A. Curtis, 1853, Ann. Mag. Nat. Hist., Ser. 2, 12: 434.
= P. scanicus Fr., 1863, Monogr. Hymenomyc. Suec., 2, 2: 269.
= P. lindheimeri Berk. et M.A. Curtis, 1872, Gre-villea, 1, 4: 50.
= P. fumosogriseus Cooke et Ellis, 1881, Grevil-lea, 9, 51: 103.
= Daedalea oudemansii var. fennica P. Karst., 1882, Medd. Soc. Fauna Fl. Fenn., 9: 69.
= Myriadoporus adustus Peck, 1884, Bull. Torrey Bot. Cl., 11, 3: 27.
= Polystictus gloeoporoides Speg., 1889, Boln Acad. Nac. Cienc. Córdoba, 11, 4: 451.
= Polyporus macrosporus Britzelm., 1894, Ber. Naturw. Ver. Schwaben, 31: 174.
= P. ochraceocinereus Britzelm., 1895, Botan. Zbl., 62: 311.
= P. burtii Peck, 1897, Bull. Torrey Bot. Cl., 24: 146.
= Coriolus alabamensis Murrill, 1907, N. Amer. Fl., 9, 1: 19.
= Polyporus excavatus Velen., 1922, Ceské Hou-by, 4-5: 641.
= P. cinerascens Velen., ibid.: 642. =P. atropileus Velen., 1925, Mykologia (Prague), 2: 74.
= P. tegumentosus Velen., ibid.: 74. = Daedalea solubilis Velen., 1926, Mykologia (Prague), 3: 102.
Icon.: Bulliard (1790: pl. 501, 2, ut Boletus pelloporus); Rostkovius (1837: tab. 38 ut Polyporus adustus); Kennedy, Larcade (1971: figs 1-12, ut Polyporus adustus); Phillips (1981: p. 236); Breitenbach, Kranzlin (1986: fig. 329); Gilbertson, Ryvarden (1986: fig. 65); Ryvarden, Gilbertson (1993: fig. 72); Roy, De (1996: fig. 13); Bernicchia (2005: fig. 140; pl. p. 610); Niemela (2005: fig. 53); Jung et al. (2014: fig. 1A); Ryvarden, Melo (2014: fig. 76).
Basidiocarps 1-4.5 x 2-10 x 0.3-1.5 cm, annual (wintering), as sessile, decurrent or patch-like resupinate clustering pilei of tough-fleshy consistency. Upperside (when present) subtomentose to matt or rugulose at drying, cream to isabelline with gray tinges, often with pale or subochraceous obscure zonation, grayish or blue-grayish along the margin, rather loose and spongy. The margin as a rule acute, sometimes border-like, slightly undulating, white, then with cineraceous shades, sterile up to 2 mm at the maturity. Context two layered with rather loose light cream or grayish upper layer 0.2-1 cm thick and dense grayish-black layer (so-called "black line") above the hymenophore. Hymenophore as a single tube layer 0.02-0.5 cm thick of ceraceous consistency, initially cream, then smoky-gray to blackish mouse-gray. Pores 6-7 per mm, angular, rather thin-walled; pore surface combines whitish (tube mouths) and smoky-gray to grayish-black tinges (fig. 1).
Hyphal system monomitic in the tubes and pseudodimitic in context. Generative hyphae 2-4 ^m in diam., regularly branched at acute margin, with regular clamp connections. Pseudoskeletal hyphae 3-6(10) ^m in diam., fibrous or ramified, thick-walled to subsolid in KOH, in some parts swelling at 10 ^m in cross. Cystidia none. Basidia 10-15 x 4-5.5 ^m, clavate with median constriction, 4-spored, with a basal clamp. Basidiospores (4)4.3-5.5(6.5) x (2.2)2.5-3.5 ^m, ellipsoid-cylindric, smooth, thin-walled, negative in Melzer's reagent (fig. 2).
On dying trees, fallen logs and branches, stumps, buried wood and small debris of many hardwoods, rarely conifers, causing a white rot. For pathogenic significance - see Brooks (1925).
S ub strata: Acacia, Acer, Aesculus, Ailanthus, Alnus, Betula, Carpinus, Castanea, Celtis, Corylus, Crataegus, Cytisus, Eucalyptus, Fagus, Fraxinus, Juglans, Larix, Malus, Myoporum, Olea, Platanus, Populus, Prunus, Rhamnus, Robinia, Rosa, Quercus, Salix, Sambucus, Sorbus, Syringa, Tamarix, Tilia, Ulmus, Abies, Larix, Picea, Pinus (Jura et al., 2011; Bondartseva et al., 2014; Ryvarden, Melo, 2014).
General distribution: EUROPE (Austria, Belarus, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Italy, Latvia, Lithuania, the Netherlands, Norway, Poland, Portugal, Romania, Russia, Slovenia, Spain, Sweden, Switzerland, Ukraine, United Kingdom); SOUTH AMERICA (Argentina, Brazil, Chile); CENTRAL AMERICA (Costa Rica, Cuba); NORTH AMERICA (Canada, Mexico, USA); ASIA (Armenia, Azerbaijan, Georgia, Iran, Israel, Japan, Korea, Mongolia, Nepal, Russia, Sri Lanka, Turkey); SOUTHERN HEMISPHERE (Papua New Guinea, Australia, New Zealand); AFRICA (Ethiopia, Malawi) (Jura et al., 2011).
Distribution in Russia: see Tables 1, 2.
Cultural characteristics: Nobles (1965); Westhuizen (1971).
Note: In the field, the species can be easily identified due to smoke or mouse-gray hymenial fields with white-cream sterile border, by small pores 6-7 per mm, and black line above the hymenophore. Some variants of B. fumosa can come nearer to the discussed species, but they are easily separated by more obscure dense zone of cinnamomeous colors and (in median) larger pores which are as a rule anisodiametric. After wintering, its blackish-gray hymenophore can change the color to coffeate (resembling those of B. fumosa), but in this state white reticulum (glacing tube moths) absent and pores in median stay sufficiently smaller.
The variability of the species concerns mode of growth organization and the degree of tubes development. A lot of forms were described during species history and the basic ones are described below.
Forma resupinata (Bourdot et Galzin) Domanski, Orlos et Skirg., 1967, Flora Polska, Grzyby (Myco-ta), Aphyllophorales: 114. - Leptoporus adustus f. resupinatus Bourdot et Galzin, 1928 ("1927") Hy-menomyc. France (Sceaux): 552.
Basidiocarps as resupinate patches 3-10 and more cm in diam. with well-developed mucedinous margin. The hymenophore is well-developed, but pores can be locally enlarged at 2-5 per mm (Bond-artsev, 1953). The micromorphology varies as in a neutral type. This is prostrate growth form of the fungus known on many hardwoods.
Forma tegumentosa (Velen.) Bondartsev, 1953, Tinder Fungi Europ. U.S.S.R. Caucasus: 240. -Polyporus tegumentosus Velen., 1925, Mykologia (Prague), 2: 74 (see fig. 3).
Basidiocarps as decurrent to resupinate prostrating pilei 4-6 cm in the largest dimension with-
Fig. 2. The microstructures of Bjerkandera adusta: a -the hyphal structure of a sterile colored «tegument» (f. tegumentosa, LE 287571), b - a typical hystion, composing a sterile tissues of the fungus, c - basidia, d - basidio-spores. Scale bar - 5 ^m.
Fig. 1. The most widespread («effused-reflexed») morphotype of Bjerkandera adusta presented by clustering pilei, developing from a common effused "stroma". Scale bar - 1 cm.
Fig. 3. Bjerkandera adusta f. tegumentosa (LE 287571), growing in a natural conditions. Scale bar - 1 cm.
out a hymenophore. A central fields of the patches ochraceous-gray, or mouse-gray, margin white. The fungus resembles a stromata of pyrenomycete Kretzschmaria deusta (Hoffm.) P. M. D. Martin. The micromorphology varies as in a neutral type. This is sterile and more or less prostrate, but negatively geotropic (with upward subhymenial filed) growth form of the fungus known on many hardwoods.
Epitype: Russia, Saint Petersburg, "Literator-skie mostki" museum necropolis, on stump of Acer platanoides, 12 VII 2015, leg. et det. I. V. Zmitro-vich (LE 287571).
Basidiocarps 2.1 x 1.5 x 0.05-0.2 cm, wrinkled due to basidiocarp initial centers, prostrate with in-rolling undulating margin and centrally sterile (teg-
umentose) negatively geotropic subhymenial fields of isabelline, then mouse-gray coloration. On marginal areas under lens can be allocated whitish net with cellars 7-10 per mm and ca. 0.1 mm deep. The micromorphology varies as in a neutral type. The tegument structure is pictured in fig. 2a. Basidia and basidiospores not developed.
Forma solubilis (Velen.) (Velen.) Bondartsev, 1953, Tinder Fungi Europ. U.S.S.R. Caucasus: 239. - Daedalea solubilis Velen., 1926, Mykologia (Prague), 3: 102.
Basidiocarps as easily separated from substrate dorsally attached resupinate patches 2-4 cm in diam. with well-developed bolster-like margin. The pores of normal sizes, but of daedaleoid appearance,
smoky-gray. The micromorphology varies as in a neutral type. This is resupinate growth form of the fungus which has a certainly disordered hymeno-phore. Known on hardwoods.
Exsiccates examined. F. O. Westerberg "Flora Suecica", no. 10, Sweden, 6 X 1912, leg. F. O. Westerberg (ut Polyporus stereoides), det. A. S. Bondartsev. - J. Lütkemüller "Kryptogamae exsic-catae, no. 308, Austria, on dry trunk in the garden (ut Polyporus adustus). - A. de Magocst-Dietz "Kryptogamae exsiccatae", no. 308b: Hungaria, Budapest, Botanical garden, trunk of Ailanthus glandulosa (ut Polyporus adustus). - "Farlow Herbarium" distr. Harvard Univ., "California Fungi" distr. Herb. Univ. California, no. 246, Los Gatos canyon, on Alnus rubra, 23 II 1924, leg. H. E. Parks (ut Polyporus adustus). - V. Litschauer et H. Lohwag "Fungi selecti ex-iccati europaei", no. 121: Austria, Wien, Botanical garden, on dry Fagus sylvatica, 12 VII 1929, leg. V. Litschauer (ut Poria canescens P. Karst.), det. A. S. Bondartsev ut Bjerkandera adusta f. resupinata. -E. Leppik "Fungi Estonici exsiccati, fasc. 1", no. 18, Tartumaa, on fallen Populus tremula, 15 VI 1930, leg. E. Leppik (ut Leptoporus adustus); ibid., no. 19, Tartu, Botanical garden, on drying Aesculus hippo-castanum, 6 VII 1930, leg. E. Leppik (ut Leptoporus adustus). - S. Lundell et J. A. Nannfeldt "Fungi Exsiccati Suecici, Praesertim Upsaliensis", no. 57, Upland, Uppsala, Carolinapark, on stumps of frondose tree, 8 VII 1933, leg. S. Lundell (ut Polyporus adustus). - K. E. Murashkinsky "Hymenomycetes Sibiri-ae", Kazakhstan, Borovoye, on Sambucus, I 1936 (ut Coriolus adustus f. resupinatus). - S. Lundell et J. A. Nannfeldt "Fungi Exsiccati Suecici, Praesertim Upsaliensis", no. 2616, Gästrikland, Gävle, Valls-hage, on dead trunk of Sambucus racemosa, 21 VIII 1956, leg. J. A. Nannfeldt (ut Polyporus adustus). - "Plantae Norvegicae" distr. Mus. Bot. Univ. Oslo, no. 95, Lyngdal, on Tilia cordata, 26 IX 1969, leg. L. Ryvarden. - "Herbier National de Mycologie du Canada", no. GCF 26, Quebec, Gatineau park, on wood, X 1979, leg. N. Binyamini. - "Fungi Ros-sici" distr. Mus. Bot. Univ. Helsinki, no. 2, Karelia, Vodlozero Nat. Park, on Populus tremula/Phellinis tremulae, 23 VIII 1994, leg. R. Penttilä. - "Fungi Rossici" distr. Mus. Bot. Univ. Helsinki, no. 36, Karelia, Vodlozero Nat. Park, on Populus tremula, 25 VIII 1994, leg. R. Penttilä.
2. Bjerkandera fumosa (Pers.: Fr.) P. Karst., 1879, Medd. Soc. Fauna Fl. Fenn. 5: 38. = Boletus fumosus Pers., 1801, Syn. Meth. Fung., 2: 530. -Polyporus fumosus Pers.: Fr., 1821, Syst. Mycol.,
1: 363. - Gloeoporus fumosus (Pers.: Fr.) Pilat in Kavina et Pilat, 1937, Atlas Champ. Eur., 3: 149. -Tyromyces fumosus (Pers.: Fr.) Pouzar, 1966, Folia Geobot. Phytotax. Bohemoslov., 1: 370. - Polyst-ictoides fumosus (Pers.: Fr.) Teixeira, 1986, Rev. Brasil. Bot., 9, 1: 43. - Grifola fumosa (Pers.: Fr.) Zmitr. et Malysheva in Zmitr., Malysheva et Spirin, 2006, Mycena, 6: 21.
= Boletus imberbis Bull., 1791, Hist. Champ. France, 10: 339.
= Daedalea saligna Fr., 1818, Observ. Mycol., 2: 241.
= Polyporus pallescens Fr., 1818, Observ. Mycol., 2: 256.
= P. demissus Berk., 1845, London J. Bot., 4: 345.
= P. salignus var. holmiensis Fr., 1874, Hymeno-myc. Eur.: 544.
= P. fragrans Peck, 1878, ("1877"), Ann. Rep. N. Y. St. Mus. Nat. Hist., 30: 45.
= Bjerkandera pallescens subsp. pura P. Karst., 1882, Medd. Soc. Fauna Fl. Fenn., 9: 69.
= Polyporus hederae Ade, 1911, Mitt. Bayer. Bot. Ges., 2: 371.
= P. decurrens Velen., 1922, Ceske Houby, 4-5: 657.
= P. emergens Velen., 1922, ibid.: 657.
= P. eminens Velen., 1922, ibid.: 639.
P. robiniae Velen., 1922, ibid.: 658.
P. tyttlianus Velen., 1922, ibid.: 686.
P. aberrans Velen., 1925, Mykologia (Prague), 2, 5: 73.
Icon.: Sowerby (1799: tab. 230, ut Boletus pel-loporus); Bulliard (1798: tab. 445, ut Boletus imberbis); Breitenbach, Kränzlin (1986: fig. 330); Gilbert-son, Ryvarden (1986: fig. 66); Ryvarden, Gilbertson (1993: fig. 73); Roy, De (1996: fig. 14); Bernicchia (2005: fig. 141; pl. p. 611); Niemelä (2005: fig. 54); Jung et al. (2014: fig. 1B); Ryvarden, Melo (2014: fig. 77).
Basidiocarps 1-7 * 1.5-10 * 0.3-2 cm, annual (wintering), as sessile, decurrent or subresupinate clustering pilei of tough-fleshy consistency. Up-perside tomentose to matt or appressedly hispid-squamulose, whitish, pale-cream to isabelline, or tan, sometimes with cinnamomeous obscure median zone, rather loose and spongy. The margin more or less obtuse, border-like on resupinate parts, slightly undulating, white-cream, then with isabelline and cinnamomeous shades, sterile up to 2 mm at the maturity. Context with anise odor when fresh, two-layered with buff soft fibrous upper layer 0.3-1.5 cm thick and dense brownish-cinnamon layer above the
hymenophore. Hymenophore as a single tube layer 0.02-0.7 cm thick of ceraceous consistency, initially cream, then tan to coffeate. Pores (1)2-4(5) per mm, round or angular, isodiametric or anisodiametric; pore surface combines whitish, cream or yellowish (tube mouths) and buff to isabelline (pore substructure) tinges (fig. 4).
Hyphal system monomitic in the tubes and pseu-dodimitic in context. Generative hyphae 2.5-4.5 |m in diam., regularly branched at acute margin, with regular clamp connections. Pseudoskeletal hyphae 3.2-7(12) |m in diam., fibrous or ramified, thick-walled to subsolid in KOH, in some parts swelling at 12 |m in cross. Cystidia none. Basidia 19-23 x 4.5-7 |m, clavate with median constriction, 4-spored, with a basal clamp. Basidiospores 5.5-7.2 x 2.5-3.7 |m, ellipsoid-cylindric, smooth, thin-walled, negative in Melzer's reagent (fig. 5).
On dying trees, fallen logs and branches, stumps, buried wood and small debris of many hardwoods, causing a white rot. For pathogenic significance -see Bondartsev (1924).
Substrata: Acer, Aesculus, Ailanthus, Alnus, Betula, Corylus, Cytisus, Eucalyptus, Fagus, Fraxinus, Juglans, Malus, Myoporum, Philadelphus, Populus, Prunus, Quercus, Salix, Sambucus, Sorbus, Tilia, Ulmus (Ryvarden, Melo, 2014), Cedrus deodara (Roy, De, 1996).
General distribution: EUROPE (Austria, Belarus, Belgium, Bulgaria, Czech Republic, Denmark, Estonia, Finland, France, Georgia, Germany, Hungary, Italy, Latvia, Lithuania, the Netherlands, Norway, Poland, Romania, Russia, Slovakia, Sweden, Switzerland, Ukraine, United Kingdom), NORTH AMERICA (Canada, USA), ASIA (China, India, Kazakhstan, Russia, Thailand, Uzbekistan), NORTH AFRICA (Bondartsev, 1953; Gilbertson, Ryvarden, 1986; Ryvarden, Gilbertson, 1993; Roy, De, 1996; Bondartseva, 1998).
Distribution in Russia: see Tables 3, 4.
Cultural characteristics: Nobles (1965).
Note: This is rather variable species, easily recognized by two-layered context with obscure lower dense strate and strongly developed whitish reticu-
Fig. 4. The most widespread («imbricate») ecotype of Bjerkandera fumosa presented by clustering pilei, developing from a common «stroma», deeply rooted into substrate (Salix fragilis): a - a general view, b - pore surface with labyrinthine areas, c - wintered basidiocarps. Scale bar: a - 10 mm, b, c - 5 mm.
Fig. 5. The microstructures of Bjerkandera fumosa: a - a typical hystion, composing the subhymenium, b - a typical hystion, composing the context, c - basidia, d - basid-iospores. Scale bar - 5 ^m.
lum over tube moths. In some cases, the hymeno-phore looks to be whitish-cream without deep coloration, but usually tan or coffeate shades are well expressed. The cap clusters reminiscent those of B. adusta, but two species easily distinguishable due to diverse pore shape and sizes, coloration of the context and hymenophoral fields. B. fumosa has typically more thick basidiocarps with more or less obtuse margin and haven't a mouse-gray tinges in the hy-menophore coloration. The coffeate wintered specimens of B. adusta differs by smaller pores (6-7 per mm vs. 1-5 per mm in B. fumosa) and dense cera-ceous black line above the tube (dense zone in B. fumosa is looser and of cinnamomeous color). Several chromatic and growth forms were described during species history, the basic ones are listed below.
Forma flavipora (Bourdot et Galzin) Zmitr. et Bondartseva, comb. nov. (MB 813429). - Leptopo-rus imberbis f. flaviporus Bourdot et Galzin, 1925, Bull. trimest. Soc. mycol. France, 41, 1: 131 (see fig. 6).
As a neutral type, but with yellowish-cream to lemon-yellow pore surface without a coffeate substructure.
Epitype: Russia, Saint Petersburg, Botanical garden of the Komarov Botanical Institute, on drying Sorbus aucuparia, 15 IX 2014, leg. & det. I. V. Zmitrovich (LE 287572).
4
Fig. 6. Bjerkandera fumosa f. flavipora (LE 287572). Scale bar - 5 mm.
Basidiocarps 4.5 * 2.5 * 1.7 cm, as sessile de-current clustering pilei of tough-fleshy consistency. Upperside tomentose, slightly scrupose, pale-cream with isabelline margin, rather loose and spongy. The margin more or less obtuse, undulating, isabelline 2.5 mm. Context two-layered with buff soft fibrous upper layer ~1.3 cm thick with weak coffeate streak above the hymenophore. Hymenophore as a single tube layer ~0.05 cm thick, of ceraceous consistency, buff in section. Pores 3-4 per mm, angular and anisodiametric; pore surface yellowish-cream. The micromorphology varies as in a neutral type. Basid-iospores 5.5-6.5 * 2.5-3.0 ^m.
Forma saligna (Fr.) Donk, 1933, Med. Bot. Mus. Univ. Utrecht, 9: 164. - Polyporus salignus Fr., 1838, Epicr.: 452.
As a neutral type with decurrent base of cap clusters, daedaleoid pores and almost white upperside. Known on Salix sp. in the Netherlands, Sweden, Belarus and Orel Region of the Russia (Donk, 1933; Bondartsev, 1953).
Forma alba (Huds.) Donk, 1933, Med. Bot. Mus. Univ. Utrecht, 9: 164. - Boletus albus Huds., 1762, Fl. Angl.: 626.
Resembling the previous form, but with sessile (not decurrent) caps with more intensively colored uppeside. Known on Salix sp. in the Netherlands, France and Sweden (Donk, 1933).
Exsiccates examined. L. Romell "Fungi Exsiccati Praesertim Scandinavici", no. 11, Sweden, Holm, on trunk of Quercus sp., 24 XI 1889, leg. L. Romell (ut Polyporus holmensis). - "Terek District Station of Plant Protection" (dupl. herbarium), no. 634, on stumps of Juglans regia, 25 X 1926, leg. A. Lobik (ut Polyporus imberbis). - J. Smarods "Fungi Latvici", Vidzeme, on Fraxinus excelsior, 19 XI 1935, leg. J. Smarods. - E. Leppik, V. Litschauer "Fungi Estonici" (Mycotheca Lab. Phytopath. Univ. Tartuensis), dupl., on fallen trunk of Sambucus nigra, 9 XI 1939, leg. E. Leppik (ut Septoporus imber-
bis). - S. Lundell et J.A. Nannfeldt "Fungi Exsiccati Suecici, Praesertim Upsalienses", no. 440, Upland, Bondkyrka parish, on stump of Populus tremula (?), 28 IX 1935, leg. S. Lundell (ut Polyporus fu-mosus). - A. Jones "Mycological Collections" distr. Herb. Univ Illinois, no. 95, on dead Ulmus sp., 22 XI 1959, leg. A. Jones (ut Polyporus fumosus). - A. Jones "Mycological Collections" distr. Herb. Univ Illinois, no. 157, on dead Ulmus sp., 13 XI 1960, leg. A. Jones (ut Polyporus fumosus). S. Lundell, J.A. Nannfeldt et L. Holm "Fungi Exsiccati Suecici, Praesertim Upsalienses", no. 3088, Vâstmaland, Norberg, on stump of Ulmus glabra, 27 VIII 1963, leg. I. Nordin.
Apart from aforementioned species, Murrill (1907) gives two additional descriptions which hardly correlated to any known taxa. Their diagnoses are presented below.
Bjerkandera terebrans (Berk. et M. A. Curtis) Murrill, 1907, N. Amer. Fl., 9, 1: 42. = Polyporus terebrans Berk. et M. A. Curtis, 1869 ("1868"), J. Linn. Soc. Bot., 10: 306.
"Pileus subfleshy, thick, flabelliform, convex, 4 * 4-5 * 1 cm, attached by a thick, laterally-compressed, concolorous, pubescent elongation resembling a stipe, but probably the result of an effort of the part of sporophore to escape from the substratum; surface isabelline to luteous, pubescent-scabrous, azonate, smooth; margin obtuse, entire: context white to isabelline, homogeneous, soft-corky, nearly 1 cm thick; tubes whitish when young, fuliginous in dried specimens, less than 1 mm, mouths 4 to a mm, edges obtuse, entire. Spores not examined" (Murrill, 1907).
Type locality: Cuba (without exact region indication).
Substrata: on dead trees.
Distribution: known only from the type locality.
It is probably a certain form of B. fumosa, but stipe-like base not allow to reject also possibility of variant of Osteina obducta (Berk.) Donk.
Bjerkandera subsimulans (Berk. et M. A. Curtis) Murrill, 1907, N. Amer. Fl., 9, 1: 42. = Polyporus simulans Berk. et M. A. Curtis in Sacc., 1888, Syll. Fung., 6: 117; nec Bjerkandera simulans P. Karst., 1888, Rev. Mycol., 10, 37: 73 = Postia tephroleuca (Fr.) Jülich, 1982.
"Pileus explanate, fleshy-tough, sessile, dimidiate of fan-shaped, often attached by a narrow base, 5-10 * 10-15 * 0.3-0.7 cm, surface smooth, partially glabrous and partially clothed with scanty, flexible hairs; margin thin, acute, broadly sterile, lobed, with a zone of appressed hairs and blackish as tough scorched for 5-10 mm: context fibrous, hard and corky when dry, white to isabelline; tubes 2-5 mm long, white to fuliginous, mouths angular, irregular, 1 to 3 a mm, edges thin. Spores not examined" (Murrill, 1907).
Type locality: Cuba (without exact region indication).
Substrata: dead trunks.
Distribution: known only from the type locality.
This superficial description corresponds in a certain respects to Abortiporus biennis (Bull.) Singer.
This work was carried out in the Laboratory of Systematics and Geography of Fungi of the Koma-rov Botanical Institute of the Russian Academy of Sciences in canvas of the State task N 01201255602.
Table 1
Herbarium data on distribution of Bjerkandera adusta over Russia territory
and its substrate preferences
Region Substrata Date of collection Collector Herbarium numbers
European part
Bryansk Region, Bryansk vic. Fraxinus excelsior VII 1907 A. S. Bondartsev LE 26254
Kaliningrad Region, Golubaya river - 25VI2010 V. M. Kotkova LE 268941
Kaluga Region, Kaluga vic. Salix sp. 10 X 1909 Chernyshov LE 26141
Kaluga Region, Ugra National Park Fraxinus excelsior 28 VII 2013 S. V. Volobuev LE 299084
Karelia Republic, Lakh-denpokhja Betula sp. 10 X 1950 A. S. Bondartsev LE 26286
Continuation of Table 1
Region Substrata Date of collection Collector Herbarium numbers
Karelia Republic Populus trémula 20 IX1995 V. M. Lositskaya LE S5364
Karelia Republic, Matrosy Populus trémula V M. Kotkova LE 203909
Kursk Region, Kursk timber wood 1907 A. S. Bondartsev LE 26191
Kursk region, Korochi Acer sp. 2S VIII 1945 L. A. Lebedeva LE 26219
Leningrad Region, Nadevitsy Betula sp. 31 VII 1932 T. L. Nikolaeva LE 26255
Leningrad Region, Siverskaya Picea abies 4 VII 1936 R. Singer LE 26Ш
Leningrad Region, Yanega Populus tremula 19 IX1961 M. A. Bondartseva LE 26212
Leningrad Region, Berezovye Ostrova protected area Populus tremula 23 VII 2003 V M. Kotkova LE 242293
Leningrad Region, Berezovye Ostrova protected area Alnus incana/ Gloeoporus dichrous S VII 2004 V M. Kotkova LE 26S4S5
Leningrad Region, Cheremenetskiy protected area Populus tremula 7 IX 2002 V M. Kotkova LE 2S3S75
Leningrad Region, Kur-galsky protected area Populus tremula 11 IX1997 I. V Zmitrovich LE 2034S4
Leningrad Region, Kur-galsky protected area Quercus robur 12 IX1997 I. V Zmitrovich LE 20340S
Mari El, Kuvshin forestry - 20 VII 1937 B. P. Vassilkov LE 2625S
Moscow Region, Push-kino Betula sp. 27 VIII ? A. S. Bondartsev LE 2624S
Moscow Region, Mikhailovskoye Populus tremula 1 V 1907 A. S. Bondartsev LE 26164
Moscow Region, Goro-dishche Populus sp. 20 IX 200S N. V Psurtseva LE 265214
Nizhegorod Region Pinus sylvestris cones/ litter 5 VIII 1997 W. A. Spirin LE 20S432
Orel Region, Lov-chikovo Tilia cordata 1 VIII 2011 S. V. Volobuev LE 29S944
Orel Region, Turovka Corylus avellana 11 VIII 2012 S. V. Volobuev LE 29S777
Orel Region, Naryshkino Populus tremula б X 2012 S. V. Volobuev LE 291153
Rostov Region, Rostov-on-Don Betula pendula 15V2009 Yu. A. Rebriev LE 2S7095
Saint Petersburg, Botanical Garden of Komarov Botanical Institute - VIII 190S A. S. Bondartsev LE 26147
Tver Region, Tsentralno-Lesnoi reserve Betula sp. 5 IX2011 V M. Kotkova LE 2S4240
Tula Region, Tula vic. timber wood 1911 Trusova LE 26203
Udmurtia Republic, Izhevsk vic. Tilia cordata 15 IX1964 Kyganova LE 26237
Caucasia
Chechen Republic, Terek Vitis sp. 20 V 1926 L. Guseva LE 26079
Chechen Republic, Naurskaya - 20 V 1926 L. Guseva LE 26Ш
End of Table 1
Region Substrata Date of collection Collector Herbarium numbers
Karachaevo-Cherkessia Republic, Teberda reserve Carpinus betulus 19 VIII 2012 N. V Psurtseva LE 2SS32S
Stavropol Territory, Pyatigorsk Fagus sylvatica 12 IX1926 A. Lobik LE 26257
Stavropol Territory, Zheleznovodsk Fagus sylvatica 13 VII 1926 G. Lagadidze LE 26234
Siberia
Gorno-Altai Autonomous Republic, Kaitanak Betula sp. 19 VII 196S M. A. Bondartseva LE 26146/162
Gorno-Altai Autonomous Republic, Kaita-nak Salix sp. 19 VII 196S M. A. Bondartseva LE 26156
Irkutsk Region, Orlenga Betula sp. 2 IX1967 M. A. Bondartseva LE 26167/68
Irkutsk Region, Irkutsk vic. - 26 VIII 1911 Alexandrov LE 26166
Krasnoyarsk Territory, Monastyrskoye Betula sp. 2S VI 1920 A. L. Yavorskiy LE 26179/190
Krasnoyarsk Territory, Stolby Betula sp. S XI196S M. A. Bondartseva LE 26152
Far East
Khabarovsk Territory, Vinogradovka - XI1929 A. S. Bondartsev LE 26144
Khabarovsk Territory, Khabarovsk vic. Acer sp. 9 V 1935 L. V. Ljubarskiy LE 26213
Khabarovsk Territory, Bolshekhekhtsyrskiy reserve Betula sp./Trichaptum 20 VIII 1979 M. A. Bondartseva LE 26135
Khabarovsk Territory, Bastak reserve Corylus avellana 21 VII 2011 N. V. Bukharova LE 290715
Primorye Territory, Kangauz Carpinus sp. 192S A. S. Bondartsev LE 26184
Table 2
The substrate preferences of Bjerkandera adusta in old-growth arboreta on Saint Petesburg territory
Old-growth arboreta Substrata
The Peter the Great Botanical Garden of the Komarov Botanical Institute of RAS Acer platanoides, Alnus incana, Amelanchier canadensis, Betula pendula, Crataegus oxyacantha, Duschekia fruticosa, Fraxinus excelsior, Juglans regia, Larix sibirica, Malus baccata, Phellodendron amurense, Populus alba, Quercus robur, Sambucus racemosa, Salix alba, S. fragilis, Sorbus aucuparia, S. intermedia, Tilia cordata, T. platyphyllos, Ulmus laevis, Ulmus scabra.
"Literatorskie mostki" museum necropolis Acer platanoides, Philadelphus coronarius, Quercus robur, Sorbus aucuparia, Syringa josikaea, S. vulgaris, Tilia cordata, T. platyphyllos, Ulmus scabra
Volkovskoye cemetery Acer platanoides, A. negundo, Betula pendula, Fraxinus excelsior, Philadelphus coronarius, Populus nigra, Quercus robur, Sorbus aucuparia, Syringa josikaea, Tilia cordata, T. platyphyllos, Ulmus laevis, U. scabra
The dendrarium of the Kirov Forest Management Academy Acer platanoides, Alnus glutinosa, Corylus avellana, Crataegus sanguinea, Fraxinus excelsior, Malus baccata, Salix fragilis, Sorbus aucuparia, Tilia cordata, Ulmus laevis.
Table 3
Herbarium data on distribution of Bjerkandera fumosa over Russia territory and its substrate preferences
Region Substrata Date of collection Collector Herbarium numbers
European part
Bryansk Region, Bryansk Betula sp. 20 VII 190S A. S. Bondartsev LE 26325
Karelia Republic, Sortavala forestry Sorbus aucuparia 21 VIII 1993 A. V. Ruokolainen LE 20S915
Kostroma Region, Vasilyevka - 1907 A. S. Bondartsev LE 2635
Kursk Region, Kursk Acer sp. 25 V1906 A. S. Bondartsev LE 2631S
Kursk Region, Kursk, Lazaretny garden Fraxinus excelsior 25 IX1906 A. S. Bondartsev LE 26349
Kursk Region, Les-na-Vorskle Betula sp. - I. E. Brezhnev LE 26339
Leningrad Region, Nizhnesvirsky reserve Populus tremula 4 VIII 2014 N. I. Kalinovskaya LE 303S09
Mordovia Republic, Mordovsky reserve Ulmus glabra 10 IX2013 S. Yu. Bolshakov LE 301234
Moscow Region Malus domestica 22 X 19S4 M. V. Gordienko LE 26295
Orel Region, Alexandrovka Populus tremula 5 X 2012 S. V. Volobuev LE 292076
Orel Region, Tureika Quercus robur 2 IX2012 S. V. Volobuev LE 29S611
Rostov Region, Veshenskaya Betula sp. 1 X 2004 Yu. A. Rebriev LE 227757/76S
Saint Petersburg, Botanical garden of Komarov Botanical Institute Ulmus glabra - A. S. Bondartsev LE 26352
Saint Petersburg, Volkovskoye cemetery Ulmus laevis 16 IX 2003 W. A. Spirin LE 20S17S
Udmurtia Republic, Kigbay Salix sp. 5 IX1965 Kychanova LE 26344
Udmurtia Republic, Izhevsk Salix sp. 9 IV 2009 V. I. Kapitonov LE 247363
Caucasia
Stavropol Territory, Pyatigorsk, Beshtau Carpinus betulus IX 1934 A. S. Bondartsev LE 26355
Urals
Sverdlovsk Region, Serginsky Ulmus sp. 1S IX1957 N. T. Stepanova-Kartavenko LE 26319
Far East
Sakhalin Territory, Krasnopolye Ulmus japonica 31 VIII 1954 V N. Ljubarskiy LE 26367
Table 4
The substrate preferences of Bjerkandera fumosa in old-growth arboreta on Saint Petesburg territory
Old-growth arboreta Substrata
The Peter the Great Botanical Garden of the Komarov Botanical Institute of RAS Acer platanoides, Crataegus sanguinea, Malus domestica, baccata, Salix fragilis, Sorbus aucuparia, S. intermedia, Tilia cordata, Ulmus laevis, U. scabra.
"Literatorskie mostki" museum necropolis Acer platanoides, Salix alba, Sorbus aucuparia, Syringa vulgaris, Tilia cordata
End of Table 4
Old-growth arboreta Substrata
Volkovskoye cemetery Acer negundo, Fraxinus excelsior, Salix alba, Sorbus aucuparia, Syringa josikaea, Tilia cordata, Ulmus scabra
The dendrarium of the Kirov Forest Management Academy Acer platanoides, A. negundo, Alnus incana, Betula pendula, Cornus sanguinea, Crataegus sanguinea, Fraxinus excelsior, Larix sibirica, Malus baccata, Picea glauca, Populus nigra, Quercus robur, Salix fragilis, Sambucus racemosa, Sorbaria sorbifolia, Sorbus aucuparia, Syringa josikaea, Tilia cordata, T. platyphyllos, Ulmus laevis.
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