Гриб Ганодерма, классификация (на английском языке)

Non abbreviated authority: (Curtis) P. Karsten
Bibliography: Karsten, P.A., 1881, Revue mycologique, Toulouse 3(9): 17 (more info)
Year of publication: 1881
Status of the name: combination; legitimate
Sanctioned by: Fries, SM1:353 (Polyporus lucidus (Curtis) Fr. 1821)
MycoBank's opinion: currently used
Classification: Fungi, Basidiomycota, Agaricomycotina, Agaricomycetes, Polyporales, Ganodermataceae, Ganoderma
Basionym: Boletus rugosus Jacq. 1774 [UNA; MB473292]
Obligate synonym(s): Grifola lucida (Curtis) Gray 1821 [LEG; MB362180]
Boletus lucidus Curtis 1781 [LEG; MB269837]
Polyporus lucidus (Curtis) Fr. 1821 [LEG; MB449530]
Fomes lucidus (Curtis) Cooke 1885 [LEG; MB159723]
Placodes lucidus (Curtis) Quél. 1888 [LEG; MB472359]
Phaeoporus lucidus (Curtis) J. Schröt. 1888 [LEG; MB470439]
Morphic status: Teleomorph (anamorph connection(s) unknown or not recorded)
Taxonomic synonym(s):
Ordered by epithet name Boletus castaneus Weber 1787 [ILL; MB473713]
Boletus supinus var. castaneus (Weber) J.F. Gmel. 1792 [LEG; MB498207]
Boletus crustatus J.J. Planer 1788 [LEG; MB468261]
Boletus dimidiatus Thunb. 1784 [LEG; MB468264]
Boletus flabelliformis Leyss. 1761 [LEG; MB255032]
Boletus ramulosum var. flabelliformis (Leyss.) J.F. Gmel. 1792 [LEG; MB498190]
Boletus laccatus Timm 1788 [LEG; MB468291]
Polyporus laccatus (Timm) Pers. 1825 [LEG; MB156470]
Agaricus lignosus Lam. 1783 [LEG; MB468048]
Ganoderma nitens Lázaro Ibiza 1916 [ILL; MB451357]
Boletus obliquatus Bull. 1781 [LEG; MB194313]
Ganoderma ostreatum Lázaro Ibiza 1916 [LEG; MB141406]
Agaricus pseudoboletus Jacq. 1773 [LEG; MB468090]
Ganoderma pseudoboletus (Jacq.) Murrill 1902 [LEG; MB469335]
Agarico-igniarium trulla Paulet 1793 [LEG; MB467960]
Boletus verniceus Brot. 1804 [LEG; MB468341]
Boletus vernicosus Bergeret 1783 [LEG; MB468342]
Descriptions:
Culture description (Bazzalo, M.E.; Wright, J.E. 1982, Mycotaxon 16(1): 300)

Cultural features Figs. 75-79, 81-82, 88-90, 107.
Strains: BAFC n°112 = ARGENTINA: Catamarca, Dique de Collagasta leg. Laterra, 14.1.1981. BAFC n°815 ibid.
Code Number: 2. 3. 8. 10. 37. 39. 45. 53. 54. 55.
Macroscopic characters: growth slow, covering Petri dishes in 5 weeks. Mycelium mat not dense, adhering to agar, arranged in concentric bands the first week, alternatively white and ochraceous (Pl. 11 K 1). After the 4th week it becomes totally ochraceous yellowish with more or less darker concentric bands (Fig. 107). Texture completely farinaceous, margin subfelty. Reverse:discoloration brown. Margen regular, border smooth. Odour sweetish. Oxidase reaction: with tannic acid= ++++ with growht; with gallic acid= ++++ without growth; with gum guaic = +.
Microscopic features: marginal mycelium formed only by generative, clamped hyphae, with septa restricted to clamps, scarcely branched, 2,5 µm diam. (Figs. 88-90); during the 2nd week fibrous hyphae appear, slender, hyaline, thick-walled, clampless, heavily branched, 1-2 µm diam. (Figs. 77-78). Ochraceous yellowish mycelium formed by: i) generative hyhae as above; ii) clamped, thick-walled hyphae, golden, unbranched, 2-7 µm diam., of uniform thickness or like rosary beads (Figs. 79, 81-82); iii) stag-horn clamped hyphae, thick-walled , with numerous branches, generally dichotomic, projecting in several planes, 1-3 µm diam., or with short branches laterally disposed in a single plane at regular intervals, 2-3 µm diam.; iv) globose, thin-walled golden yellow cuticular cells which are very abundant, tightly packed forming a "pseudoparenchyma", 5-20 µm (Figs. 75-76); v) fiber hyphae as in the margin, scarce. Submerged mycelium similar to aerial.
These cultures are characterized by a strongyellowish colour that extends to all the mycelial layer, whereas in other species it is limited to a few spots and is not so intense. The cultures resemble those of G. subamboinense var. laevisporum but differ in the lack of chlamydospores.
For a long time the G. lucidum-complex has been a difficult problem for taxonomists. It has been a tradition to consider all the stipitate forms as G. lucidum s. str. and the sessile olies as G. resinaceum (= G. sessile Murr.). This distinction has an absolute lack of anatomical foundation. When one deals with a large number of specimens, clear distinctions appear with regard to spore ornamentation, whatever the macroscopic configuration may be. This had already been observed by Haddow (1931) and was only reconfirmed 35 years later by Steyaert (1967a). Haddow termed the spores of G. lucidum as of the "rugose" type. Unfortunately the holotype of this species has not been found, although Steyaert (1972) states there is a coloured illustration of such specimen in Flora Londinensis (1781), which was collected at Peckham, S of London. This would constitute the present type of the species. However, such an illustration does not reveal the spore features. Attempts to find a neotype at Peckham have failed. Karsten, the founder of the genus, left at H a specimen with the same type of "rugose" spores (Steyaert, 1972), which could be selected as neotype and thus arrive at a satisfactory "modus vivendi" to distinguish among both species.




Culture description (Nobles, M.K. 1948, Canad. J. Res., C 26: 336)

Ganoderma lucidum (Leyss. ex Fries) Karst.
Cultural characters: (PI. V, Fig. 3; Pl. VI, Figs. 9 to 13).
Growth characters. Growth rapid, plates covered in two weeks. Advancing zone even, raised aerial mycelium extending to limit of growth. Mat white and remaining so or becoming "wood brown" (7.0YR5.7/4.0) (two weeks), with color subsequently masked by overgrowth of whitish 'bloom', at first slightly raised, cottony, then appressed, felty to pellicular with farinaceous surface, traversed by deep wrinkles or grooves. Reverse unchanged for two or three weeks, then with scattered patches of "honey yellow" (2.0Y6.7/6.2) to "buckthorn brown" (8.0YR4.8/6.5), finally bleached, with wrinkles prominent, producing a veined appearance. No odor. On gallic acid agar diffusion zone weak to strong, no growth or diameter up to 4.0 cm.; on tannic acid agar diffusion zone moderately strong to strong, diameter 2.0-4.0 cm.
Hyphal characters. Advancing zone: hyphae hyaline, nodose-septate, 2.2-4.5(-6.0) µm diameter. Aerial mycelium (thin tough skin that peels from agar): (a) hyphae as in advancing zone, with frequent branches and numerous small projections; (b) fiber hyphae very numerous, with walls thick and refractive, lumina narrow or apparently lacking, except in main hyphae, frequently branched, the ends long, slender, curving and interwoven, 1.0-3.0 µm diameter; (c) cuticular cells thin-walled, produced by inflation of nodose-septate hyphae, at first with contents staining in phloxine, then empty, closely compacted and interwoven with fiber hyphae and staghorn hyphae to form pseudoparenchymatous layer, which may remain hyaline or become brown; (d) chlamydospores very numerous, walls slightly thickened, terminal and intercalary, broadly ovoid to elongate, 12.0-21.0 x 7.5-10.5 ìm; (e) staghorn branched hyphae with minute branches in all planes very numerous in 10222, not observed in other isolates. Submerged mycelium: (a) nodose-septate hyphae and (b) chlamydospores as described above.
Type of rot: soft spongy white rot of broad-leaved trees (see below).
Descriptions of cultural characters: Davidson, Campbell, and Blaisdell (64) (under Polyporus lucidus), Davidson, Campbell, and Vaughn (67) (under P. lucidus), Humphrey and Siggers (92).
As a result of a comparison of European collections of Polyporus lucidus, which occurs commonly on broad-leaved trees and more rarely on coniferous trees in Europe, with American collections of Ganoderma tsugae from coniferous trees, usually Tsuga spp., Atkinson (2) concluded that they should be brought together in one species, Ganoderma pseudoboletum. Haddow (79) made a critical study of European and American collections of G. lucidum and American collections of G. tsugae and concluded that there were no constant morphological differences and that host specificity was not a valid basis for separation of the species. Therefore he agreed with Atkinson in placing the two under one species, for which he used the accepted name, Ganoderma lucidum. Boyce (35), Lowe (99), and others have followed this practice, but Overholts (119) has retained G. (Polyporus) tsugae in specific rank. In the present study the three cultures of G. lucidum isolated from fruit bodies on broad-leaved trees were distinguishable from the four cultures of G. tsugae from Tsuga spp. on the basis of clamydospore production, which was prolific in all the cultures of G. lucidum and completely lacking in those of G. tsugae. Because of this difference, the species have been treated separately, but this does not constitute an argument for their separation in taxonomic studies, since the number of isolates in each species is too small to provide convincing evidence. Interfertility tests are required to establish conclusively whether one or two species are involved.
Ganoderma lucidum is unique among the Ganoderma species studied in having chlamydospores and therefore its key pattern does not coincide with that for any of the other species.


Non-original description (Bazzalo, M.E.; Wright, J.E. 1982, Mycotaxon 16(1): 297)

GANODERMA LUCIDUM (Leys.:Fr.)P. Karst., Rev. Mycol. 3 (9): 17.1881.
= Boletus lucidus Leys. Fl. Halensis p. 300. 1783. = B. dimidiatus Thunb., Fl. p. 348, tab. 39. 1784 (Fide Imazeki). = B. laccatus Timm, F1. megalop. Prodr., p. 269. 1788 (Fide Imazeki).
= Polyporus lucidus Leys.:Fr., Syst. Mycol. I: 353. 1821. = P. laccatus Timm:Pers., Mycol. europ. 2: 54. 1825 (Fide Imazeki). = P. japonicus Fr., Epicr. p. 442. 1838 (Fide Imazeki) lucidus (Leys.:Fr.) Sacc., Syll. Fung. 6: 157. 1881. = Placodes lucidus (Curt.:Fr.)Quél., Fl. Mycol. p. 399. 1888 (Fide Domanski). = F. japonicus Fr. in Sacc., Syll. Fung. 6: 156. 1888 (Fide Imazeki). = Ganoderma pseudoboletus(Jacq.) Murr., Bull. Torrey bot. Cl. 29: 602. 1902 (Fide Imazeki). = G. laccatum Pat. in Bresadola, Icon. Mycol. 21, t. 1004. 1932, non G. laccatum Bourd. & Galz. 1928. Icon. Michael & Henning, Pilzfr. 2, t. 73. 1960 (Fide Domanski et al.).
Annual, sessile and dimidiate or, more frequently with a lateral stem and then usually reniform (Figs. 1,3). Pilei isolated, small to medium sized, 2-8 x 2-4,5 x 0,5-2 cm. Pileus surface radially rugose and concentrically sulcate, brilliantly laccate, light reddish brown (Pl. 6 L 12 of Maerz & Paul), to dark reddish brown or mahogany (Pl. 7 L 6). Margin sterile, generally thick and blunt, sometimes acute, white in actively .growing specimens, becoming yellowish and reddish brown inwards; in older specimens of the same colour as pileus surface, and then incurved. Stem lateral, vertical, cylindric, usually long, Blender, tortuous, 4-10 cm long,0,5-2 cm thick, reddish black to almost black, laccate, brilliant, somewhat thicker at the base. Cutis thin, brilliant black. Context almost as thick as the tube layer but thickening towards the base of the stem, ochraceous brown when young (Pl. 13 H 10) to dark brown when mat-ure (Pl. 14 L 12), corky. Dermis of the "hymenodermis" type, composed of thick, golden walled, claviform elements originating from the ends of skeletal hyphae, with narrow lumina and blunt ends, arranged in a palisade-like hymenium (Fig. 62), 5-10 mm thick, the total thickness of the dermis 14-42 mm. These elements are covered by a thick layer of a lacque-like substance that dissolves in a hot solution of 5% KOH. Hymenophore white to yellowish white when young, greyish white in mature specimens (Figs. 2, 4) with a tube layer up to 7 mm long, slightly lighter than context (Pl. 13 D 8). Pores small, round, somewhat irregular, 4-7 per mm, 6-200 mm diam. Dissepiments 17-116 ,mm diam. Hymenium not persistent, composed of scant globose to subglobose basidia, 9-19 x 7-14 µm. No other hymenial elements pres-
ent. Basidiospores subovoid with the apex truncate, perisporium hyaline, smooth and thin, and endosporium golden with relatively scant endosporic pillars, wide and long, reaching the perisporium and rumpling it so that it appears strongly "rugose" (Figs. 23-25, 61); 9-13 x 5-6,9 µm. Hyphal system trimitic with hyaline,thin-walled,clamped,septate generatives, 1-4 µm diam.,septa restricted to clamps, scantily branched, abundant at the growth margin of pileus and dissepiments, rare or absent in the context (Fig. 71). Skeletals "arboriform" (Teixeira, 1956), aseptate, clampless, very long, 3-6 pm diam., scantily branched, branches with limited growth at distal end, with thick golden walls, sometimes subsolid; they compose most of the context and dissepiments, originating immediately behind the growth margin from generative hyphae (Fig. 68). Binding hyphae of the "Bovista" type (Cunningham,1946a), aseptate, clampless, profuse ly branched, tortuous, of limited growth, generally thinner and ligliter than the skeletall, 1-3 µm diam., rather scant and only present in the context; they are intertwined with the latter, giving the context its firm cohesion (Figs. 6970).
Hosts: at the base of trunks and on roots of harwoods, rarely on conifers. Recorded also on Acer, Quercus, Castanea, Alnus, Populus, Fagus, Fraxinus and Pinus (Domanski, 1967). According to Domanski (loc. cit.) it grows saprophytically and only exceptionally attacks living trees. Boyce (1938) agrees with this. However, Pirone (1957) through experiments with Acer proved that it is an important parasite of hardwoods.
Distribution:apparently worldwide in temperate and tropical zones.It has been recorded for Europe, Asia, Philippines, Australia, Africa and North and South America (in the latter for Venezuela, Brazil, Uruguay and Argentina).




Non-original description (Corner, E.J.H. 1983, Nova Hedwigia, Beih. 75)

G. lucidum (Fr.) Karst. Figures 4, 5, 19 The following details come from collections made in East Anglia. Spore 9.5-12 x 6-6.5 µm, but swelling in potash 10-13 x 6.8-8 µm, endospore 8-9.5 x 4.7-5.7 µm, in potash 9.3-10.5 x 5.3-6.5 µm, brown distinctly echinulate, 1-guttate. Basidia 24-33 x 11.5-14.5 µm; sterigmata 3-4, 4-4.5 µm. Cystidia -20 x 3-5 µm (-35 x 9.5-14 µm, Demelius), near the pore-mouths and in the immature hymenium, evanescent, ventricose with narrow apex, thin-walled, colourless. Hymenium continuous, not thickening.
Hyphae trimitic with skeleto-binding cells and Bovista-hyphae, not encrusted, not dextrinoid; skeleto-binding cells -2.500 µm long overall, terminal, the stalk 250-1.700 x 3.5-6(-8), µm, (-14 µm, Hansen), in places with ampulliform swellings -9 µm, with thick brown walls darening in potash but scarcely swelling, dividing over a short distance into 1-4(-5) branches -1.000 µm long, simple or once bifid, tapering into hyaline tips 1-2.5 µm wide; binding hyphae as Bovista-hyphae - 600 µm long, 1.5-3 µm, wide on origin and developing thick brown walls, 0-3 times bifid, tapering to filiform tips 0.5-1 µm wide, derived from short thick-walled intercalary generative cells; generative hyphae 1.5-5.5 µm wide, clamped, not or scarcely inflating, the growing tips at the margin of the pileus 2-3.5 wide; dissepiments with skeletals 3-4 µm wide, trimitic as the flesh (dimitic, Hansen), not agglutinated. Surface of pileus with a laccate crust -200 µm thick, consisting of three layers: - a, an outer amorphous red-brown resinaceous excretion 20-30 µm thick, mostly soluble in potash with a slight brown solution; b, a close palisade of agglutinated subclavate cells 40-75 x 6-10 µm (-15 µm, Hansen), with narrow interwoven stalks 3-4 µm, wide, arising from generative hyphae, developing thick brown walls (-5 µm, thick) and often linear lumen, the swollen ends of the cells forming a regular layer 30-40 µm thick immediately below the resinous excretion; c, a hypodermal layer c. 100 µm thick, consisting of agglutinated generative hyphae 3-4 µm, wide with slightly thickened bright yellow walls (the colour in the inner part of the walls and turning brown in potash, but in the cytoplasm before wall-thickening), also with the tapered ends of binding processes.
The crust develops from a delicate layer of thin-walled clavate cells 1120 x 5-7 µm, formed at the ends of generative hyphae, like an incipient hymenium. They excrete the resinous material which forms the outer layer, and they lengthen basipetally into the flexuous stalks. The tapering ends of binding processes are subsequent intrusions. The abundant Bovista-hyphae seem characteristic. Whether the dried spores always swell appreciably in potash, I am uncertain, but the possibility is a precaution that needs to be taken. It seems yet uncertain if typical G. lucidum occurs in the tropics. The most recent critical account of it is given by Jähn, Kotlaba and Pouzar (1980) and by Kotlaba and Pouzar (1981). In comparing it with their new species G. atkinsonii they emphasize the colour of the living pileus of G. lucidum as bright yellow to orange and cherry red when young, then bloodred, purplish red, to reddish brown, the colours darkening on drying. It seems that G. atkinsonii needs comparison with G. neojaponicum Imazeki (1939).
In my key to the Malesian species of laccate Ganoderma (p. 125) G. lucidum comes close to G. chalceum, but there is no record of lobulate palisade cells in G. lucidum. G. tropicum, with broader spores and many unbranched skeletal (acicular), should be compared.


Non-original description (R.L. Steyaert 1975, CMI Descr. Pathog. Fungi Bact. 445)

Ganoderma lucidum (W. Curtis: Fr.) P. Karst., Rev. mycol. 3: 17, 1881.
= Boletus lucidus W. Curtis, Flora londinensis f. 4, pl. 224, 1781 (typus iconog.).
= Polyporus lucidus Fr.. Syst. Mycol. 1: 353, 1821.
= Polyporus laccatus (Tim.) Pers., Myc. Europ. 1: 54, 1825.
= Polyporus japonicus Fr., Epicr.: 442. 1836.
= Ganoderma valesiacum Boud., Bull. Soc. mycol. Fr. 11: 28, 1895.
= Ganoderma tsugae Murrill, Bull. Torrey bot. Cl. 29: 601, 1902.
= Ganoderma mongolicum Pilat, Annls mycol. 38: 78. 1940.
Basidiocarp (basidioma) annual, sessile to long mesopode or pleuropode depending on which substratum it is produced, either tree bole or buried wood, generally thick, up to 20 mm, sometimes more or less tumid when sessile, pileus subplane to very irregular, up to 160 mm across; stipe up to 100-120 mm long, 10-30 mm thick; upper surface and stipe shiny, lacquered, blackish-brown, irregularly humped; margin and pore surface white when growing, tawny with age. Cutis thin, a fraction of 1 mm, context from russet near the tube layer to white for the major part; context varies according to latitude from white (Scandinavia) to tawny (Portugal and central African mountains) with concentnc lines of various shades; tubes monolayered, up to 15 mm long, russet, concolorous with lower part of context. Cutis of hymenioderm type composed of anticline inflated extremities of hyaline context, hyphae swollen by melanoid substances leaving usually a central lumen; the melanoid substances are easily saponifed by KOH; inflations clavate, up to 50 µm long, 8-10 µm wide near apex. Skeletal Hyphae brown in the coloured parts of the context, nearly hyaline or hyaline close to cutis, 3-4 µm wide, sparsely ramified, subparallel to intertwining, specially in the less coloured parts of the context, no binding hyphae. Pores circular to irregular, 100-430 (av. 210) µm diam., with dissepiments usually relatively thin, 10-320 (av. 52) µm thick; distance between axes of pores ± 260 µm. Basidiospores ovoid, double-walled, epispore thin, ovoid, hyaline; endospore thick, ovoid, with usually a small part of the apex more or less Date brown or shades of brown, bearing relatively few long and thick echinules that support the epispore, sometimes fused into a short crest, in freshly discharged spores the apex of the epispore remains swollen and bulging, the tip of the apex soon collapses leaving an ovoid spore with a truncated tip, 8-13 (av. 10,6) x 5,5-9 (av. 6,9) µm; occasionally smooth, mono-walled, spores can be observed, brown, 8,5-10 (av. 9,3) x 6-7,5 (av. 6,5) µm; gasterospores are very rarely observed (only one basidium from Portugal) in the context, navicular, pale brown, with a very thick deposit on the walls, 16-34 (av. 22) x 9-14 (av. 10,3) µm.
Hosts: Angiosperms and Gymnosperms: Quercus sp., Quercus Ilex, Ligustrum vulgare, Carpinus betulus, Fagus
sylvatica, Corylus avellana, Populus alba, Prunus insititia, Picea sp., Taxus sp., Larix sp., Abies sp.
Disease: Butt rot and lethal root disease of many tree species.
Geographical distribution: Throughout the temperate zones of the northern hemisphere. From the Pacific shores
of the USA and Canada, through temperate Europe to the Pacific shores of Asia and Japan. Also in the mountains of Central Africa above the 1500 m level.
Physiological specialization: None reported.
Literature: Kavina & Pilat, Atlas des champignons de l'Europe III: 481-484, 1942; Steyaert, Persoonia 7: 93-95, 1971.