V. F. Malysheva, I. V. Zmitrovich
Testing the Trametes hirsuta complex
Introduction. The Trametes hirsuta complex comprises two weakly differentiated species, Trametes hirsuta and T. pubescens, and one problematic species, T. velutina. The differences between three taxa were ambiguously estimated in the literature (Table 1):
Table 1
Morphological species delimitations in Trametes hirsuta complex in basic literature
Fries (1821) / Bondartsev (1953) / Ryvarden, Gilbertson (1994) / Niemelä (2000)Trametes hirsuta
Basidiocarp features and pileus cover / Whitish, with coriaceous suberose, strigose-zonate pileus / Pilei 1.5-5 x 3-10 x 0.3-1 cm, coriaceous, semicircular to decurrent, with concentrically sulcate and zonate strigose surface of yellowish, cineraceous, isabelline, rufescent or grayish-olivaceous colour / Basidiocarps annual, effused-reflexed to resupinate, applanate to thick, coriaceous, upper surface hirsute, gray, zonate or concentrically sulcate / Annual or persistent, thin and applanate, suberose, small of mediumsized; externatlly hispid, zonate, gray at the base, creamish at margin, with greenish tint due to algae
Hymenophore / Pores round with obtuse bruishing dissepiments / Pores round, with obtuse whitish, creamish, isabelline or grayish margins, 3-4 per mm / Pores with thick entire dissepiments, white to tan or cinereous, (1)3-4 per mm / Pores creamish to gray, round with even margins, 2-4 per mm
Hyphal structure / No data / Hyphae thin-walle or subsolid, (1.5) 2-5 (6) mm in diam. / Trimitic; generative hyphae 2.5-9 mm in diam., skeletal hyphae 3-9 mm in diam., binding hyphae 2-4 mm in diam. / Trimitic
Basidiospores / No data / Subcylindrical, slightly curved, with attenuated base, 6-8 x 2-3 mm / Cylindric, 6-9 x 2-2.5 mm / (4.6)5.2-6.5(7.8) x (1.8)2-2.4(2.6) mm
Trametes pubescens
Basidiocarp features and pileus cover / White, fleshy-suberose, pubescent-zonate pileus / Pilei 2-5 x 3-8 x 0.2-1 cm, coriaceous to almost suberose, semicircular to conchate, with weakly radially and concentrically sulcate pubescent surface of white, yellowish to ochraceous colour / Basidiocarps annual, sessile or effused-reflexed, up to 6 cm wide, dimidiate, often in imbricate clusters, thin coriaceous; upper surface tomentose to finely pubescent or almost glabrous, cream colour to warm buff, azonate or faintly zonate / Annual, applanate or of medium sizes, imbricate, externally finely pubescent, creamish (in some cases with weakly expressed fumose zones)
Hymenophore / Pores almost round, plane / Pores roundish to sinuous and angular, thin-walled with subacute margins, 2-4 per mm / Pores angular, 3-5 per mm, dissepiments becoming thin / Pores creamish-white or yellowish, 2-3 per mm
Hyphal structure / No data / Hyphae thin-walled or subsolid, 2.5-3.5(6.5) mm / Trimitic; generative hyphae 2-4 mm in diam., skeletal hyphae 5-10 mm in diam., binding hyphae1.5-4 mm in diam. / Trimitic
Basidiospores / No data / Cylindrical, asymmetrical to slightly curved, 5-8 x 2-2.5(3) mm / Cylindric, slightly curved, 5-7 x 2-2.5 mm / (5.4)6-7(7.3) x 2.1-2.6 mm
Trametes velutina
Basidiocarp features and pileus cover / Whitish, with suberose-coriaceous, finely pubescent, weakly zonate pileus / Considered as a synonym of Coriolus pubescens / Considered as a synonym of Trametes versicolor / Annual, thin and elastic, applanate or rosette-like, in some cases fan-shaped, externally creamish-white, finely pubescent and radial fibrils, with obscure hygrophanous zones
Hymenophore / Pores round, white / Pores creamish to yellowish, 3-4(5) per mm
Hyphal structure / No data / Trimitic
Basidiospores / No data / (4.8)5.3-7.6(7.9) x (1.7)1.9-2.6(2.7) mm
The type material on Boletus hirsutus Wulfen (1788), B. pubescens Schumach. (Schumacher, 1803) and B. velutinus J. J. Planer (None, Planer, 1788; Persoon, 1794) is absent, and validation ascends to Fries (1821), where three taxa were considered as a members of the genus Polyporus.
The Friesian concept of Polyporus velutinus was changed between 1821 and 1832. In 1821, Fries used this taxon in sense of Planer (None, Planer, 1788), i.e. approaching the concept to those of P. hirsutus. However, in ‘Index’ (Fries, 1832), the author refers to diagnosis of P. velutina in Elenchus (Fries, 1828), where description of fungus gravitates rather to Personian concept (Persoon, 1794), that is approaching to P. pubescens.
The second step in complication of species concept in the conglomerate was description of Polyporus fibula by Fries (1874). This description sufficiently corresponded to P. hirsutus with exception to finely velutinate (not strigose) pileus cover of the fungus.
The absence of type material and scarce macroscopic descriptions of taxa defends a certain variability of their interpretations during a long time. The attribution of strigose specimens to P. hirsutus became a standard, whereas the rest variability concept was rotated due to unstable interpretation of P. velutina.
Overholts (1953) gave a new concept of P. velutinus, which include this taxon in equally instable taxonomical field of Trametes ochracea (Pers.) Gilb. & Ryvarden.
In the modern manuals, the most of authors have reduced P. velutina to synonymy of T. pubescens (Donk, 1974; Ryvarden & Gilbertson, 1994; Bondartseva, 1998), however, Niemelä (2000) has kept Trametes velutina as a single taxon.
The purpose of our investigation is molecular approach to distinguishing of basal divergency in this species complex.
Material and methods.
Microscopical study of basidiomata was carried out as described by Gilbertson & Ryvarden (1986). Freehand sections and squash mounts of basidiomata were examined in 5% KOH and 2% Cotton Blue. Spore conglomerations were searched in basidiocarps’ tomentum. Spore measurements contained 30 spores per specimen. The following abbreviations were used: L – spore length, W – spore width, Q* – quotient of the mean spore length and mean spore width (L/W ratio). Specimens are preserved in the herbarium of Komarov Botanical Institute (LE).
Results
Discussion.
Two clades of T. hirsuta-complex (including spore statistics)
Morphological series.
Species descriptions.
Trametes hirsuta (Wulfen) Lloyd, Mycol. Writ. 7: 1319. 1924.
≡ Boletus hirsutus Wulfen in Jacquin, Collnea bot. 2: 149. 1788.
= B. velutinus J.J. Planer, Ind. Pl. erfurt. Fung. add.: 26. 1788.
= B. wulfenii Humb., Fl. Friberg. Spec.: 96. 1793.
= B. nigromarginatus Schwein., Schr. naturf. Ges. Leipzig 1: 98. 1822.
= Polyporus galbanatus Berk., Ann. Mag. nat. Hist. (Ser. 1) 10: 377. 1843 (‘1842’).
= P. vellereus Berk., J. Bot. 1: 455. 1842.
= P. cinerescens Lév., Ann. Sci. Nat. (Bot., sér. 3) 2: 184. 1844.
= P. cinereus Lév., Ann. Sci. Nat. (Bot., sér. 3) 5: 140. 1846.
= P. gourliei Berk., Flora Tasmaniae. 2: 253. 1860.
= Coriolus velutinus P. Karst., Trudy Troitsk. Otd. imp. russk. geogr. obsc. 8: 61. 1906.
= Polyporus fagicola Velen., České Houby 4–5: 654. 1922.
= Trametes porioides Lázaro Ibiza, Polip. Fl. Esp.: 372. 1917.
Basidiocarps annual or two-three seasonal, sessile or slightly decurrent (in several cases dorsally attached and subresupinate), applanate, semicircular or reniform in outline, with slightly inflated base, 1.3-5 x 3-12 x 0.2-1.2 cm. Abhymenial surface concentrically sulcate, whitish, cream, grayish, or ochraceous-tan, in some cases combines several colorations; often greenish or green due to aerophytic algae; tomentum up to 5 mm thick, varies from soft-floccose to strigose, less expressed in fissures and near the margin. Margin initially subacute and slightly inrolled, later blunt and straight, sterile; initially white, than creamish with tan or gray pruina near the pores. Hymenophore as a single tube layer up to 6 mm thick; surface creamish with tan or gray pruina, more or less even; pores round, with indulited dissepiments, rather thick-walled, (2)3-4 per mm.
Hyphal system dimitic with skeleto-binding hyphae. Generative hyphae 2.5-5(9) mm in diam., fibulate, moderately branched, with rare septation; prevails in marginal zones of basidiocarp and in tubes; often protruded the hymenium as agglutinated fascicles (pegs). Skeleto-binding hyphae 2-9 mm in diam., with straight dendroid axes 3-5 mm in diam. (diameter falls to dendrite periphery) and staghorn-like appendages 2-3 mm in diam. Sclerified hyphae prevail in basidiocarp; in abhymenial surface these hyphae form gross fascicles composing the tomentum.
Leptocystidia fusoid, 10-21 x 3-5.5 mm.
Basidia clavate with inflated epibasidium, 12-21 x 4-6.5 mm, 4-spored, with a basal clamp.
Basidiospores cylindrical, slightly curved, thin-walled, inamyloid, acyanophilous or weakly cyanophilous.
Inhabits dead tissues of angiosperms and occasionally gymnosperms (Abies, Cupressus, Juniperus, Larix, Picea, Pinus, Thuja, Taxus). Produces a white rot.
Trametes pubescens(Schumach.) Pilát in Kavina & Pilát, Atl. Champ. Eur. 3: 268. 1939.
≡ Boletus pubescens Schumach., Enum. Pl. 2: 384. 1803.
= B. velutinus Pers., Ann. Bot. 11: 29. 1794.
= Polyporus sullivantii Mont., Annls Sci. Nat., Bot. (Sér. 2) 18. 1842.
= P. molliusculus Berk., J. Bot. 6: 320. 1847.
= Hansenia imitata P. Karst., Medd. Soc. Fauna Flora Fennica. 13: 161. 1886.
= Polyporus subluteus Ellis & Everh., Am. Nat. 31: 339. 1897.
= Coriolus applanatus P. Karst., Finl. Basidsvamp. 46: 3. 1904.
= Trametes merisma Peck, Bull. N.Y. St. Mus. 139: 31. 1910.
= T. quercina Lloyd, Mycol. Writ. 7: 1114. 1922.
Basidiocarps annual, sessile or slightly decurrent (usually in rosette-like or imbricate clusters), conchate-applanate, semicircular or reniform in outline, with pointed or depressed base, 1.5-6 x 2-8.5 x 0.2-1 cm. Abhymenial surface even or slightly concentrically rolled, near the base often scrupose and ridged, white, yellowish, cream, watery-grayish, or tan, with hygrophanous zones when fresh; finely tomentose to subnude, in some cases with greenish tint; tomentum up to 3.5 mm thick, soft-floccose or usually even, velvety, less expressed near the margin. Margin subacute and slightly inrolled, later straight, sterile; white to yellowish. Hymenophore as a single tube layer up to 5 mm thick; surface creamish or yellowish, uneven in maturity; pores anisodiametric, thin-walled, in some cases with acute margins, (2)3-4 per mm.
Hyphal system dimitic with skeleto-binding hyphae. Generative hyphae 2-6(10) mm in diam., fibulate, moderately branched, with rare septation; prevails in marginal zones of basidiocarp and in tubes; often protruded the hymenium as agglutinated fascicles (pegs). Skeleto-binding hyphae 2.5-10 mm in diam., with straight dendroid axes 2.5-6 (10) mm in diam. (diameter falls to dendrite periphery) and staghorn-like appendages 2-3.5 mm in diam. Sclerified hyphae prevail in basidiocarp; in abhymenial surface these hyphae form porous, rarely friable fasciculate tomentum.
Leptocystidia fusoid, 12-19 x 3-5 mm.
Basidia clavate with inflated epibasidium, 13-21 x 4.5-6.5 mm, 4-spored, with a basal clamp.
Basidiospores cylindrical, slightly curved, thin-walled, inamyloid, acyanophilous or weakly cyanophilous.
Inhabits dead tissues of angiosperms in temperate zones. Produces a white rot.
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