The general morphology and spore variability of Hyphoderma setigerumare discussed. The herbarium material from Belarus and northwest Russia (Leningrad region) has been involved in the study to build a generalized morphological description of the species

Yurchenko, E.O.

Zmitrovich, I.V.

Repository of Polessky State University (Репозиторий Полесского государственного университета)

Volume LXXVIII, pp.423-434 April-June 2001VARIABILITY OF HYPHODERMA SETIGERUM(CORTICIACEAE S.L., BASIDIOMYCETES) IN BELARUS AND NORTHWEST RUSSIAEugene 0. Yurchenko and Ivan V. Zmitrovich*Laboratory of Mycology, V.F. Kuprevich Institute of Experimental Botany, Akademichnaya str. 27,220733,Minsk, BELARUS E-mail: fiingi@biobel.bas-net.by ^Laboratory of Systematics and Geography of Fungi, V.L. KomarovBotanical Institute, Prof. Popov str. 2, 197376, St. Petersburg, RUSSIA E-mail: ivan@IZ6284.spb.eduAbstract: The general morphology and spore variability of Hyphoderma setigerum are discussed. The herbarium material from Belarus and northwest Russia (Leningrad region) has been involved in the study to build a generalized morphological description of the species.Key words: Corticioid fungi, Basidioma morphology, Cystidia forms, Spore size, Spore quotient.The corticioid fungus Hyphoderma setigerum (Fr. : Fr.) Donk is a widespread polytrophic saprobe, associated presumably with deciduous wood. The species is characterized by remarkable fruitbody variability (Eriksson, Ryvarden, 1975; Jiilich, 1984; Domanski, 1988, Hansen, Knudsen, 1997), and its morphological concept is sufficiently complex issue. As neutral “type” of the species some authors (Nikolayeva, 1961) consider an abstract setigerous grandinioid fungus, however in many cases such concept will be not applicable. In our opinion, the principal diagnostic characters are an association of clamped cystidia with diversely shaped hymenjal warts, while the limits of basidioma variability are principally extensive. The purpose of this work is to describe the variability in H. setigerum as known from the north-central Europe.The material from Belarus and northwest Russia (mainly Leningrad region) was studied. The vouchers taken in account are kept in the herbaria of V.F. Kuprevich Institute of Experimental Botany, Minsk (MSK, Fungi) and V.L. Komarov Botanical Institute, St. Petersburg (LE). Specimens from Belarus (MSK) have mainly been collected by E.O. Yurchenko. Collections from Russia were deposited by T.L. Nikolayeva and I V. Zmitrovich.424The data on geography and hosts of the species in Belarus are presented in Table 1. When the substrate was not reliably identified the host name is enclosed in brackets.Table 1. Substrates and spore variability in H. setigerum  (Belarusian material studied). Spore sizes are given as means of 30 measured spores and SD is given in parenthesis.Spore size and shape:MSK District of Substrate (host) Length x width Lengthno. Belarus (|im ) /w idth3812a Minsk (Fraxinus pennsylvanica)7.7 (1.6) x 3.3 (0.3) 2.43812b Minsk (Fraxinus pennsylvanica)7.5 (1.6) x 3.0 (0.4) 2.53870 Pruzhany (Quercus robur) 7.5 (1.0) x 2.8 (0.4) 2.73871 Mazyr Alnus glutinosa 8.9 (1.6) x 3.1 (0.3) 2.93872 Mazyr Alnus glutinosa 10.55(1.3) x 3.55 (0.4) 3.04095 Svislach Quercus robur 10.2 (1.1) x 3.3 (0.4) 3.14131 Lepel’ Salix pentandra 8.8 (1.4) x 3.3 (0.4) 2.74321 Lepel’ Alnus glutinosa 8.5 (1.6) x 2.9 (0.5) 3.04426 Lahoisk Sorbus aucuparia 10.6(1 .5) x 3.7 (0.5) 2.94431a Hlubokaye Sorbus aucuparia 8.2 (1.4) x 3.25 (0.4) 2.64431b Hlubokaye Sorbus aucuparia 8.0 (1.4) x 3.2 (0.4) 2.54576 Petrykau Pinus sylvestris 8.1 (1.4) x 3.7 (0.6) 2.24675 Pukhavichy Sorbus aucuparia 8.7 (1.5) x 3.4 (0.5) 2.64700 Minsk Pinus sylvestris 8.7 (1.8) x 3.8 (1.0) 2.34733 Lepel’ Betula pubescens 8.3 (1.6) x 3.0 (0.4) 2.84843 Lepel’ Alnus glutinosa 8.3 (1.2) x 2.8 (0.4) 3.04866 Lepel’ Alnus glutinosa 9.2 (1.6) x 3.0 (0.4) 3.14975 Rechytsa M alus sylvestris 8.1 (1.3) x 2.9 (0.4) 2.84976 Minsk Cerasus vulgaris 8.8 (1.2) x 3.2 (0.4) 2.84977 Minsk (Quercus robur) 8.7 (1.4) x 3.0 (0.4) 2.94978 Minsk Alnus glutinosa 7.7 (1.2) x 3.6 (0.8) 2.24979 Minsk Cerasus vulgaris 10.0(1 .3) x 3.5 (0.2) 2.94980 Khoiniki M alus domestica 7.8 (1.2) x 3.3 (0.4) 2.44981a Hrodna Betula pendula 7.3 (0.9) x 2.7 (0.2) 2.74981b Hrodna Betula pendula 8.8 (1.8) x 2.8 (0.3) 3.24983 Lahoisk M alus sylvestris 8.4 (1.6) x 3.0 (0.5) 2.84984 Minsk M alus sylvestris 11.1 (1.8) x 3.8 (0.3) 3.04254985 Pukhavichy Malus domestica 9.6 (1.5) x 3.3 (0.4) 2.94986 Lahoisk Malus sylvestris 8.7 (1.7) x 3.5 (0.3) 2.54987 Minsk Sorbus aucuparia 9.8 (1.9) x 3.6 (0.3) 2.74988 Minsk Cerasus vulgaris 9.0 (1.6) x 3.6 (0.4) 2.54989 Hrodna Tilia cordata 11.2 (1.6) x 3.6 (0.5) 3.14991 Asipovichy Alnus glutinosa 8.9 (1.8) x 2.9 (0.5) 3.24992 As ipo vichy Alnus glutinosa 9.0 (1.9) x 3.1 (0.3) 3.05076 Lepel’ Alnus glutinosa 9.2 (1.7) x 3.4 (0.4) 2.75077 Lepel’ Populus tremula 9.9 (1.4) x 3.6 (0.5) 2.85103 Dzyarzhynsk Betula pendula 7.6 (1.3) x 2.9 (0.3) 2.7Substrate. The fungus develops both on bark and wood periphery of different deciduous trees, rarely on conifers (Picea abies, Pinus sylvestris). Less frequent substrates recorded from the area are dead lichen thalli (Hypogymnia physod.es, MSK 4417, LE 201726; Parmelia omphalodes, LE 202182), pyrenomycete stromata (Daldinia concentrica, LE 206856), small plant debris, and burnt wood (conifer twigs, LE 206729), whereas the commonest one is Alnus wood. The dispersal of the species from nemoral to taiga vegetation zones is probably associated with aider rich vegetation.Macromorphology. The fungus forms resupinate fructifications of diverse morphology, and its field identification in many cases is problematic. Young basidiomata are as a rule arachnoid-farinaceous, then continuous, on decorticating wood typically effused. When found on the bark, basidiomata may arise from lenticells as initial warts, then orbicular and confluent, repeating or not the relief of substrate, reaching 5 x 15 cm and more in extent, becoming deeply and densely cracked with age. Consistency mainly soft ceraceous to slightly cartilaginous, in some cases floccose and reminding of Hyphodontia species. Thickness varies from 0.07 - 0.3 to 1 - 5 mm, sometimes even thicker (see below). Thicker parts as a rule occupy depressions in the substrate and include particles from the bark. Thickening of orbicular fruitbodies sometimes lead to semiglobose formation. Hymenial surface is initially farinaceous to granulose but becomes tuberculate, warted, or toothed with age (teeth mostly 0.25 - 1.5 mm diam and 0.2 - 1 mm high). Parmasto (1969) mentioned, that teeth in some specimens may be furnished by dendroid outgrowths (in our collections this phenomenon was observed too, e.g. LE 209772). Color varies from pure white to cream, yellow, isabelline or even ochraceous, in some cases a weak pinkish tinge was present. Margin thinning out or bolster-like, arachnoid to continuous and thickening, abrupt, in most cases adnate, but in some old fructifications loosely attached.Corticioid and grandinioid (with granulose hymenium) forms dominate in the investigated area, whereas odontioid and raduloid ones are less frequently met with (in North America the situation seems to be reverse - see Miller and Boyle, 1943). The data on some forms with rather deviating hymenial configuration are given in Table 2.426Table 2. Substrates and spore variability in some deviating growth forms of H. setigerum  (Russian material studied)Growth form of basidiomataLENo.SubstrateSpore size and shapeLength x Spore width (pm ) quotientOdontioid« semiglobose 206586 Alnus incana 7.5-12.0x2.8-4.5 2.4« with dendroid 209772 Alnus incana 7.5-12.5x3-4 2.8hyphae in teeth « effused with a 206847 Betula  sp. 7.0-11.2x2.8-3.5 2.6striking central wart Raduloid 209704 Alnus incana 7.5-12.5x3.2-4.5 2.8<< with bolster-like 202106 Alnus incana 7.5-10x2.5-4.3 2.5sterile marginConcerning the thickness of fruiting bodies we made surprisingobservations. We found specimens with conspicuous cushion-like basidiomata, up to 7-10 mm thick. Such findings seem to have been observed earlier, because Fries (1838, 1874) mentions that basidiomata in Kneiffia setigera  may be very thick, «... in sta tu  perfectissimo hemisphaericas...» and also «Specim ina maxima Pomi magnitudie...»  (dimensions were not indicated). Regardless that there are no principal limits for basidiomata growth in fungi, such thickened fruitbodies are certainly rare in nature.The specimen with thickened basidiomata from Belarus (MSK 4431b, Fig. 1) was found on a fallen decayed trunk of Sorbus aucuparia, and thickened basidiomata grown jointly with a typical, thin one (MSK 4431a). They are resupinate, cushion-like, 2.5-5 mm thick in average, up to 7 mm thick, 5-7 cm in extent, pale coloured, with rather loose, but elastic (when fresh) consistency. Hymenial surface seems not to be quite mature, uneven, lacunose-tuberculate with minute ridges formed by hyphal fascicles or even and partly powdery due to crystalline masses. The specimen collected in St. Petersburg growing on the stump of Atnus incana  (LE 206586, Fig. 2) reach a thickness of 10 mm. Its cross-section is clearly semiglobose, whereas the hymenophore is oblong, with thin, sterile margin (Fig. 2 A). Surface, like the Belarusian specimen, is in most parts uneven, lacunose, but so far as basidioma is mature, the odontioid processes are more expressed. In living state aculei in the specimen reached 4 mm.Thus, we have a rare growth form being revealed from time by time by independent investigators (see Fries, 1838; Scheremeteva, 1908). The possible reason stimulating such forms could be breakings of the normal growth conditions (change of substrate orientation), or simply interpreted as an ageby E.O. Yurchenko.1Ж428state. Specimens with unusually thickened fruitbodies recorded by the authors were mainly collected at the end of the vegetation period (October - November).Figure 2. Thickened basidiomata of Hyphoderma setigerum  (LE 206586), collected in St. Petersburg (05.10.1997, Piskaryovskii forest park) from decayed stum p of Alnus incana. A. odontioid basidioma with sterile edging, B, basidioma in cross-sections (photo I V. Zmitrovich).Hyphae (Fig. 1 B). Hyphae are distinct, 2 - 5.5 (6) pm wide, thin- to somewhat thick-walled, fibulate, with characteristic hyphodermoid ramifications, vertically arranged in subhymenium and more horizontal in subicular part; smooth or encrusted, hyaline or subhyaline, even, rarely with ampullate (MSK 4976) or irregular swellings up to 8.5 - 10 pm in diam; sometimes with adventitious septa. Walls up to 0.8 - 1 pm thick, although sometimes reaching 2 pm (MSK 4983). As a rule in young or weakly developed basidiomata all the hyphae are thin-walled. Subiculum loose or tightly packed.S ep to cy stid ia  (Fig. 3 A - C, F - H, J - N). The most peculiar and characteristic element of high diagnostic value. They are pronounced, rare in young basidiomata, as a rule well developed and abundant in mature ones, associated mainly with apices of warts, teeth or other protuberances. 55 - 185 (280) x 5.5 - 12 (14) pm, immersed or protruding to 55 - 170 pm. walls up to 2 - 2.5 pm thick, sometimes seemingly layered (MSK 4131) Typical cystidiaFigure 3. Cystidia (A-N) and basidioles (O) of Hyphoderma setigerum. A, cylindrical with wrinkled tip (MSK 4986). B, capitate (MSK 3872). C, clampless capitate, with spout-like tip (MSK 3872). D and E, capitate aseptate (D - MSK 4985, E - MSK 4988). F and G, with widened apical part (F - MSK 4131, G - MSK 4843). H, clampless, narrowed at the tip (MSK 4866). I, fusiform smooth (MSK 4866). J, with adventitious septa (MSK 3870). K, distorted (MSK 4843). L, branched (MSK 3870). M, branched at clamp (MSK 4984). N, with many short-celled segments (MSK 4991). O, clavate basidioles (MSK 4988).430are cylindrical and unbranched with truncate apices and clamped septa, but variation occur, namely: 1) with clampless septa, 2) aseptate, 3) with capitate thin-walled tip, 4) apically widened or swollen (MSK 3872), 5) with inflations in other parts, 6) tapering to the apex, 7) with many short-celled segments, 8) branched (MSK 3870).The content is hyaline to yellowish or slightly greenish, refractive or not, homogeneous or granular.The encrustation if present varies from poor to abundant, consists of minute or big double-pyramidal crystals (0.3 - 7 pm long. Fig. 3 F, G). Cystidial apices as a rule smooth or less encrusted.Basidia and basidioles. Basidia in the fungus are oi typical hyphodermoid appearance, subclavate, with characteristic median constriction and 4 long sterigmata, 25 - 35 (45) x 5 - 6 (7) pm (sterigm ata reaching 9.5 pm length), arranged in a palisade or sometimes a more loose structure. Hyaline to yellowish ovoid or short-clavate structures 7 - 9 pm wide and ca 20 pm long present in some specimens (Fig. 3 O), which probably are young stages of either basidia or cystidia.Spores (Figs. 1 F, 4). Spore shape varies from ovoid to ellipsoid and cylindrical, adaxially flattened and often slightly curved. Apiculus short to prominent, sometimes not visible. Surface smooth, in one specimen (MSK 4989) with adhered granules.Figure 4. Spores of Hyphoderma setigerum. A, MSK 4870. B, MSK 4987. C, MSK 4989. D, MSK 4733. E, MSK 4700.431Spore wall from very to moderately thin, forming distinct outline. Content hyaline or subhyaline (seemingly greenish), refractive or not, as a rule granular.Data on spore sizes are presented in Table 1 (mean length and width, standard deviation, and mean length/w idth ratio). For each specimen 30 spores were measured. Spores sizes varies in limits 5.5 - 14 x 2.0 - 5.5 pm with the marginal mean values for our specimens 7.3 - 11.2 x 2.7 - 3.8 jjm. Spore quotient varies considerably: from 1.2 to 5.0, its mean values are from 2.2 to 3.2. The generalized mean spore size for Belarusian material is 8.85 x 3.25 pm, and spore quotient is 2.76. We observed certain dependence of spore length and width from the type of substrate: fallen or erect branches and trunks. The basidiomata growing on vertical trunks and still-attached branches have mostly bigger spores than basidiomata on fallen wood (Fig. 5). Mean spore sizes are 9.4 x 3.4 for basidiomata in exposed position versus 8.7 x 3.2 for basidiomata in close contact with the ground.80 Ll7 . 3- 8.5  8 . 6 - 9.9  10.0 - 11.2Mean spore length, pm2 . 7- 3.0  3 . 1- 3.5  3 . 6- 3.9Mean spore width, pmFigure 5. Number of specimens of Hyphoderma setigerum  with mean spore length and width belonged to one of three classes, with basidiomata growing on still-attached branches and vertical trunks (white rectangles), and on fallen branches and trunks (gray rectangles).Any dependence of spore size and shape from both the host species and the type of host tissue were not found. An exception is rather short and wide spores of specimens from Pinus sylvestris (length/w idth = 2.2 and 2.3).A deviating form is represented by MSK 4978 (Fig. 6) with small ovoid or ellipsoid spores (7.7 x 3.5 pm in average), very few septocystidia, and well visible basal layer up to 90 pm thick. It is remarkable, that some spores of this specimen are subglobose (length/w idth = 1.2 - 1.3), subhymenium is richly encrusted, cystidia are weakly projecting and sparse, the hymenial palisade is very dense.Other structures. Rounded or slightly angular bodies 3.7 - 18.5 x 3.5 - 11 pm with some oily hyaline to yellowish contents (Fig, 6 B) were observed in a number of specimens. They are dispersed or aggregated in subiculum, mainly near substrate, resembling a parasitic fungus. In one thickened specimen (cited432above) was found subhyaline yeast-like structures 3 - 9.5 (15) x 3 - 5.5 pm with contents and refraction resemble those of basidiospores (Fig. 1 E). They seem to be budding, in some places forming large assemblages. The functional value of both types of structures remains unknown for the authors.Figure 6. Hyph.od.erma setigerum  (MSK 4978). A, vertical section of basidioma with basal layer. B, rounded bodies from subiculum. C, spores.Considering the here added information on basidiome morphology the elaborated diagnosis of the species may be presented as follows.Hyphoderma setigerum (Fr. : Fr.) Donk, Fungus 27: 15, 1957. Thelephora setigera  Fr. : Fr., 1828. - K neiffia setigera (Fr. : Fr.) Fr., 1838. - Peniophora setigera  (Fr. : Fr.) Hohn. et Litsch., 1906. - Odontia setigera  (Fr.: Fr.) L.W. Mill., 1934; Hydnum cristulatum  Fr : Fr., 1821; Thelephora aspera Pers., 1822 non Fr.; Hyphoderma spiculosum  Wallr., 1833; Corticium myxosporum  P. Karst., 1882.; Corticium latitans P. Karst., 1888; Odontia acerina Peck, 1901; Peniophora subtestacea Litsch., 1928.Basidiomata adnate, effused or orbicular with central wart, firstly arachnoid, then thickening, up to 1 5 - 2 0  cm in extent and 10 mm thick, ceraceous to floccose, often cracking, white to ochraceous. Margin sterile, arachnoid to bolster-like, white, then darkening. Hymenium smooth (farinaceous under lens) or toothed. Teeth 4 mm or less in length, conical to dendroid, blunt, sometimes crowded, hispid at the apex. Hyphae 2 - 6 (10) pm in diam, clamped, with age moderately thick-walled, smooth or with scattered encrustation, loosely arranged. Cystidia 30 - 280 x 5.5 - 14 pm, cylindrical, sometimes tapering or branched, as a rule septate with clamp connections, projecting on 10 - 170 pm mostly near the apex of hymenial warts, at first naked, becoming encrusted with coarse crystals. Basidia 25 - 45 x 5 - 7 pm, subclavate, constricted, with 4 sterigmata (reached to 9.5 jjm in length), hyaline; sometimes together with ovoid or short-clavate basidioles 15 - 25 x 7 - 9 pm. Spores 5.5 - 14 x 2.0 - 5.5 pm, mostly 7 - 1 1 x 3 - 4  pm, ovoid to cylindrical, slightly curved or at least clearly depressed, smooth, hyaline, as a rule granular and guttulate.On bark or wood of alder and many other deciduous, rarely coniferous trees, dead lichen thalli, old pyrenomycete stromata, plant debris and burnt pieces. Associated with a white rot. Grows from June to November. Common and widely distributed in North-Central Europe. Circumglobal species.AcknowledgementWe are grateful to Dr. N. Hallenberg, Department of Systematic Botany, University of Geteborg, Sweden, for critical review of the manuscript.Literature citedDomanski, S. 1988. Mate flora grzybow. T. 1. Basidiomycetes (Podstawczaki). Aphyllophorales (Bezblaszkowce). Cz^s'c 5. Corticiaceae; Acanthobasidium- Irpicodon. Warszawa-Krakow. 427 pp.Eriksson, J. and L. Ryvarden. 1975. The Corticiaceae of North E urope/ with drawings by J. Eriksson. Vol. 3; Coronicium-Hyphoderma. Oslo. P. 287- 546.Fries, E. 1836-1838. Epicrisis Systematis Mycologici seu Synopsis Hymenomycetum. Uppsala. 610 pp.Fries, E. 1874. Hymenomycetes Europaei sive epicriseos systematis mycologici. Uppsala. 755 pp.Hansen, L. and H. Knudsen (eds.). 1997, Nordic Macromycetes. Vol. 3: heterobasidioid, aphyllophoroid and gastromycetoid Basidiomycetes. Copenhahen. 445 pp.Julich, W. 1984. Die Nichtblatterpilze, Gallertpilze und Bauchpilze. Aphyllophorales, Heterobasidiomycetes , Gastromycetes. In; Gams, H. Kleine Kryptogamenflora. Bd. IIb /1 . Basidiomyceten. 1. Stuttgart-N.Y. 626 PP-433434Miller, L.W. and J.S. Boyle. 1943. The Hydnaceae of Iowa. Univ. Iowa Stud.Nat. Hist. 18 (2): 1-89.Nikolayeva, T.L. 1961. Hydnaceous fungi. (Flora plantarum cryptogamarum URSS, vol. VI. Fungi 2). Moscow-Leningrad. 433 pp. (In Russian) Parmasto, E. 1969. The main questions of systematics of the ordo Aphyllophorales. Mikologiya i fitopatologiya 3: 322-330. (In Russian) Sheremeteva, E.P. 1908. An illustrated handbook of fungi of Middle Russia. I. Hymenomycetineae/ compiled on the base of P. Henning’s «Hymenomycetineae» by prof. F.V. Buchholz. Part 1: Hypochnaceae, Thelephoraceae, Clavariaceae, Hydnaceae, Polyporaceae. Riga. 145 pp. (In Russian)

Variability of Hyphoderma setigerum (Corticiaceae S.L., Basidiomycetes) in Belarus and northwest Russia

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Variability of Hyphoderma setigerum (Corticiaceae S.L., Basidiomycetes) in Belarus and northwest Russia

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