Corresponding authors: Karl-Henrik Larsson (
Academic editor: R. Henrik Nilsson
Viner I, Spirin V, Zíbarová L, Larsson K-H (2018) Additions to the taxonomy of
The genus
Controversies over the taxonomic position of
In the present study, we revise the
Type material and specimens from herbaria H, S, O, GB, BPI, TAAM and BAFC were studied. Herbarium abbreviations are given according to Index Herbariorum (Thiers). Microscopic methods are described in
For DNA extraction we used either the standard CTAB protocol (
DNA sequences were edited in Geneious (Biomatters Ltd, Auckland, New Zealand) or in Sequencher 5.2.4 (Gene Codes Co., Ann Arbor, MI, USA) and deposited in GenBank (Table
Specimens and GenBank and UNITE accession numbers for DNA sequences used in this study.
Species | Specimen voucher | GenBank or UNITE accession numbers for ITS | GenBank or UNITE accession numbers for LSU | Reference |
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Larsson 14646 |
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this study | |
FR-0249548, Holotype |
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Larsson 11731 |
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MA- |
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Larsson 12971 |
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KAS-GEL4833, Holotype |
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TNM F24757, Holotype |
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MSK7247, Holotype |
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KAS-GEL3400 |
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Ryberg 021018 |
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KAS-GEL2414 |
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KAS-JR7 |
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TNM F973, Holotype |
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Larsson 12261 |
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Canfield 180, Holotype |
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H6013167 |
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Unpublished | ||
KG Nilsson s. n. |
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UC2023169 |
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Wu 9211-71 |
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Spirin 8775, Holotype |
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this study | ||
Spirin 9416 |
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this study | |
TU115575 |
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Unpublished | ||
UC2022850 |
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KUN2352 |
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this study | |
TU115495 |
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Unpublished | ||
TU124171 |
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Unpublished | ||
Cui 12887 |
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Wu 1307-42 |
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Wu 1407-105, Holotype |
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KUN2312, Holotype |
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this study | ||
Zíbarová 30.10.17 |
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this study | |
Zíbarová 26.05.17 |
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this study | |
ICMP13836 |
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Ryvarden 19767, Holotype |
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Wu 9209-27 |
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Wu 0808-32 |
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Wu 890714-3, Holotype |
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LWZ20160318-3, Holotype |
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B Nordén 030915 |
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GEL3158 |
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GEL3290 |
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Unpublished | ||
GEL3302 |
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Unpublished | ||
GEL3309 |
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GC 1508-146 |
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GEL4998 |
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Wu 1010-62 |
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CHWC1506-2 |
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Dai 11686 |
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GC1412-22 |
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Spirin 5763 |
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this study | |
PDD:91630 |
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ICMP13837 |
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KUC20130725-29 |
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Wu 0809-76 |
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FCUG 1517 |
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FCUG 2425 |
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Miettinen 7978 |
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Larsson 14653 |
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Unpublished | ||
Spirin 2877 |
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this study | ||
UC2023108 |
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Dai 10768, Holotype |
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Kotiranta 27060 |
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this study | ||
Larsson 11076 |
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Miettinen 15050,1 |
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Spirin 8565 |
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this study | ||
Spirin 8840 |
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this study | ||
Dai 12631 |
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ICMP13833 |
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Spirin 7664, Holotype |
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GC 1512-1 |
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Wu 1109-178, Holotype |
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Dai 12354 |
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CFMR:DLL2011-081 |
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Ryberg 021031 |
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UC2022842 |
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UC2023109 |
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UC2023147 |
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UC2023148 |
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GEL2690 |
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Larsson 7085 |
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Wu 1508-2 |
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Wu 9806-105, Holotype |
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KUN1989, Holotype |
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this study |
We inferred phylogenetic trees with maximum likelihood (
Maximum-likelihood (
Maximum parsimony (
For both datasets, the Bayesian inference returned trees with two main clades (Figures
Phylogenetic relationships of
Phylogenetic relationships of
In the ITS-only tree, three terminal branches represent new species that are described below.
The results allow us to introduce new species and new combinations as follows.
USA. Washington: Jefferson Co., Hoh River, on
Attenuatus (lat., adj.) – exhausted, thin.
Basidiocarp effused, up to 5 cm in widest dimension. Sterile margin white, up to 1 mm wide. Hymenial surface cream-coloured, grandinioid to odontoid; projections rather regularly arranged, from 80 µm to 200 μm high, 70–90 μm broad at base, 6–8(–9) per mm. Hyphal structure monomitic, hyphae clamped, cyanophilous. Subicular hyphae densely interwoven, thin-walled, (2–)2.4–4.6 μm in diam. (n=60/6), often short-celled, the outline of these hyphae often irregular. Tramal hyphae subparallel, thin-walled, in subhymenium densely arranged, sometimes short-celled, 2.4–3.6 μm in diam. (n=62/6). Large stellate crystals 10–13.3 μm in diam. present in subiculum and trama. Cystidia originating from subhymenium, of two types: a) subcapitate or capitate cystidia, (12–)13.5–25.1(–37)×(2.7–)3.3–5(–5.5) μm (n=80/6), b) hyphoid cystidia, (14–)16–38.3(–40.8)×2.8–4.5 (n=51/6), sometimes with crystalline cap on the top; some cystidia with granular contents in CB. Basidia suburniform, 4-spored, (12.2–)14–22(–25)×(3–)3.3–4.6(–5) μm (n=61/2), slightly thick-walled at the base. Basidiospores thin-walled, ellipsoid, (3.7–)4.1–5.5(–6)×(3–)3.4–4.5(–4.9) μm (n=180/6), L=4.85, W=3.98, Q=1.22, slightly cyanophilous.
North-western USA (Washington), on angiosperm and gymnosperm wood (fallen decorticated logs).
RUSSIA. Primorie: Khasan Dist., Kedrovaya Pad Nat. Res., on angiosperm wood, 25 Jul 2016, I.Viner KUN 2312 (H) – ITS sequence, GenBank
Crystalliger (lat., adj.) – bearing crystals.
Basidiocarp effused, soft membranaceous, up to 6 cm in widest dimension. Sterile margin poorly defined, up to 0.3 mm wide. Hymenial surface white, minutely odontioid, i.e. covered by small peg-like hyphal projections up to 60–100 μm high, 60–75 μm broad at base, 10–15 per mm, with flattened fimbriate apices. Surface between projections porulose-reticulate. Hyphal structure monomitic, hyphae clamped, faintly cyanophilous. Subicular hyphae densely interwoven, often with thickened walls, 3.2–4.4 μm in diam. (n=20/2), smooth or sparsely encrusted. Tramal hyphae subparallel, thin- to clearly thick-walled, sparsely encrusted, subhymenial hyphae densely arranged, sometimes short-celled, 2.5–3.2 μm in diam. (n=20/2), sparsely encrusted. Hyphal ends at the top of projections often strongly encrusted. Cystidia of two types: a) sparsely encrusted hyphoid cystidia at the top of projections, 21.0–29.0×2.9–4.1(–4.4) μm (n=40/2), b) subcapitate or cylindrical cystidia, of subhymenial origin, rather variable in shape and size, (11.8–)14.1–25.0(–28.0)×(2.6–)2.9–4.6(–4.8) μm (n=40/2), often heavily encrusted and rarely with a stellate crystalline cap 3.5–4.5 μm in diam. Basidia suburniform, 4-spored, 13.4–18.4(–19.0)×4.2–4.7 μm (n=20/2), slightly thick-walled at the base. Basidiospores thin-walled, elliptical, occasionally with an oil-drop, (3.1–)4.2–5.1(–5.9)×(2.4–)3.3–4.2 μm (n=60/2), L=4.66, W=3.71, Q=1.26, slightly cyanophilous.
East Asia (Russian Far East), on decayed angiosperm logs.
The peg-like hymenial projections and cystidia with stellate caps are characteristic for
Basidiocarps effused, up to 5 cm in widest dimension. No differentiated margin. Hymenial surface white, smooth or warted, farinaceous. Hyphal structure monomitic, hyphae clamped, faintly cyanophilous, thin-walled. Subicular hyphae interwoven and frequently branched, (2.2–)3.0–5.9 μm in diam. (n=61/6). Tramal hyphae subparallel, subhymenial hyphae short-celled, (1.5–)1.9–3.5 μm in diam. (n=61/6). Large, rhomboid or stellate crystals abundant in trama and subiculum, 8–10.5 μm in diam. Cystidia of two types: a) large, thin-walled cystidia of subicular or tramal origin, cylindrical or clavate, rarely slightly thick-walled (wall not exceeding 1 μm thick), (30.0–)58.9–110.0(–115.0)×4.1–8.5(–9.6) μm (n=120/6), occasionally bearing 1–2 clamped septa, b) rare astrocystidia of subhymenial origin, with a stellate crystalline cap 10–23×2–3.1 μm, in some specimens difficult to find. Basidia suburniform, 4-spored, (12.2–)13.1–20.0×(3.1–)3.4–5.0 μm (n=61/6), thin-walled. Basidiospores clearly thick-walled, elliptical to broadly elliptical, usually with an oil-drop, (3.3–)4.3–5.7(–6.1)×3.2–4.1(–4.5) μm (n=190/6), L=4.92, W=3.69, Q=1.34, cyanophilous.
Cystidial elements of
. Europe (Czech Republic, France, Italy), on herbaceous remnants, once collected from pine bark at the same spot where it was found on fern remains.
Basidiospores of two
Basidiocarp of
Cystidial elements and basidia of
Spore measurements of five
Species / specimen | L' | L | W' | W | Q' | Q | n |
---|---|---|---|---|---|---|---|
|
(3.7) 4.1–5.5 (6) | 4.85 | (3) 3.4–4.5 (4.9) | 3.98 | (0.98) 1.06–1.38 (1.46) | 1.22 | 180 |
Holotype | (4.3) 4.4–5.7 (5.8) | 4.86 | (3) 3.5–4.3 (4.7) | 3.84 | (1.1) 1.2–1.4 (1.5) | 1.27 | 30 |
Spirin 8133 | (4.4) 4.54–5.3 (5.5) | 5.01 | (3.2) 3.8–4.6 (4.7) | 4.14 | (1.06) 1.1–1.33 (1.38) | 1.21 | 30 |
Spirin 8286 | (4.1) 4.14–5.74 (6) | 4.98 | (3.1) 3.84–4.5 (4.5) | 4.11 | (1.02) 1.09–1.34 (1.36) | 1.21 | 30 |
Spirin 8779 | (4) 4–5.2 (5.4) | 4.67 | (3) 3.2–4.3 (4.4) | 3.82 | (0.98) 1.04–1.38 (1.43) | 1.23 | 30 |
Spirin 8900a | (3.7) 3.95–5.25 (5.6) | 4.56 | (3.4) 3.4–4.35 (4.9) | 3.94 | (1.02) 1.02–1.29 (1.37) | 1.16 | 30 |
Spirin 8964 | (4.5) 4.6–5.6 (5.7) | 5.02 | (3.5) 3.6–4.3 (4.8) | 4.04 | (1.1) 1.1–1.4 (1.4) | 1.25 | 30 |
|
(3.1) 4.2–5.1 (5.9) | 4.66 | (2.4) 3.3–4.2 (4.3) | 3.71 | (1) 1.1–1.4 (1.6) | 1.26 | 60 |
Holotype | (3.1) 4.2–5.1 (5.9) | 4.63 | (2.4) 3.1–3.8 (3.9) | 3.5 | (1.2) 1.2–1.5 (1.6) | 1.32 | 30 |
Bortnicov KUN 3347 | (4.2) 4.2–5.3 (5.5) | 4.69 | (3.3) 3.6–4.2 (4.3) | 3.91 | (1) 1.1–1.4 (1.4) | 1.2 | 30 |
|
(3.3) 4.3–5.7 (6.1) | 4.92 | (3.1) 3.2–4.1 (4.5) | 3.69 | (0.7) 1.1–1.6 (1.8) | 1.34 | 190 |
Lectotype | (4.2) 4.3–6 (6.1) | 5.07 | (3.1) 3.2–4 (4.1) | 3.59 | (1.2) 1.2–1.6 (1.7) | 1.42 | 39 |
Larsson 5496 | (3.3) 4.2–5.5 (6) | 4.87 | (3.1) 3.2–4.1 (4.5) | 3.61 | (0.7) 1.1–1.6 (1.8) | 1.36 | 30 |
Larsson 5622 | (4) 4.2–5.1 (5.5) | 4.6 | (3.3) 3.4–3.9 (4) | 3.63 | (1.1) 1.1–1.4 (1.5) | 1.27 | 30 |
Larsson 5627 | (4) 4.2–5 (5.6) | 4.69 | (3.3) 3.3–4.1 (4.2) | 3.73 | (1.1) 1.2–1.4 (1.4) | 1.26 | 31 |
Zibarova 26.V.2017 | (4.4) 4.7–5.8 (5.9) | 5.26 | (3.2) 3.3–4.2 (4.3) | 3.83 | (1.1) 1.2–1.6 (1.7) | 1.38 | 30 |
Zibarova 30.X.2017 | (4.2) 4.2–5.7 (5.9) | 4.99 | (3.2) 3.3–4.1 (4.2) | 3.78 | (1.1) 1.1–1.5 (1.7) | 1.32 | 30 |
|
(4) 4.5–5.9 (7) | 5.09 | (3.3) 3.7–4.8 (5.7) | 4.12 | (0.8) 1.1–1.4 (1.5) | 1.24 | 192 |
Holotype of |
(4.6) 4.7–6 (7) | 5.26 | (3.5) 3.8–5 (5.3) | 4.32 | (1) 1.1–1.4 (1.4) | 1.22 | 31 |
Holotype of |
(4) 4.1–5.7 (5.9) | 4.95 | (3.5) 3.6–4.5 (4.6) | 4.03 | (1.1) 1.1–1.4 (1.4) | 1.23 | 40 |
Spirin 2877 | (4.5) 4.7–6.1 (6.3) | 5.28 | (3.5) 3.8–5 (5.2) | 4.21 | (1) 1.1–1.4 (1.5) | 1.26 | 30 |
Spirin 9350 | (4.4) 4.7–5.7 (6.2) | 5.21 | (3.5) 3.8–4.8 (5.7) | 4.17 | (0.8) 1.1–1.4 (1.5) | 1.26 | 31 |
Spirin 9581 | (4.2) 4.2–5.8 (6.1) | 4.99 | (3.3) 3.6–4.4 (4.6) | 3.98 | (1) 1.1–1.4 (1.4) | 1.25 | 30 |
Spirin 9994 | (4.2) 4.6–5.1 (5.3) | 4.89 | (3.5) 3.6–4.5 (4.6) | 4.04 | (1.1) 1.1–1.3 (1.4) | 1.21 | 30 |
Holotype of |
(4) 4.3–5.5 (5.6) | 4.92 | (3.1) 3.1–4 (4.2) | 3.68 | (1.1) 1.1–1.6 (1.7) | 1.35 | 30 |
|
(4.8) 5.1–6 (6.2) | 5.48 | (3.7) 3.8–4.6 (4.8) | 4.21 | (1.2) 1.2–1.4 (1.5) | 1.3 | 92 |
Holotype | (4.9) 5.1–5.9 (6.2) | 5.48 | (3.7) 3.8–4.6 (4.8) | 4.22 | (1.2) 1.2–1.4 (1.4) | 1.3 | 32 |
Viner KUN 2103 | (4.8) 5–6.1 (6.2) | 5.6 | (3.8) 3.8–4.7 (4.7) | 4.24 | (1.2) 1.2–1.4 (1.5) | 1.32 | 30 |
Viner KUN 2186 | (5) 5–5.7 (5.8) | 5.37 | (3.8) 4–4.5 (4.6) | 4.18 | (1.2) 1.2–1.4 (1.5) | 1.28 | 30 |
Measurements of cystidial elements of
Species / specimen | L' | L | W' | W | n |
---|---|---|---|---|---|
|
(30) 58.9–110 (115) | 85 | (4) 4.1–8.5 (9.6) | 6.3 | 120 |
Lectotype | (67) 69.9–96.7 (110) | 83.8 | (4) 4–9.1 (9.2) | 6.5 | 20 |
Larsson 5496 | (30) 45.2–108.2 (112) | 81.2 | (4.1) 4.3–7 (7.2) | 5.7 | 20 |
Larsson 5622 | (30) 45–103 (110) | 82.7 | (4.1) 4.3–7.5 (8.5) | 5.7 | 20 |
Larsson 5627 | (56) 58.7–104.6 (110) | 79.1 | (4.4) 4.8–8.9 (9.6) | 6.4 | 20 |
Zibarova 26.V.2017 | (80) 83.8–103.3 (110) | 95.1 | (4) 5.4–8.1 (8.5) | 7.1 | 20 |
Zibarova 30.X.2017 | (67) 73.7–112.2 (115) | 87.7 | (4) 5–7.4 (7.5) | 6.3 | 20 |
|
(35) 44–84 (107) | 61.9 | (4) 4.9–10.9 (12.4) | 7.2 | 146 |
Holotype of |
(41) 43–95 (99) | 64 | (4) 5–12 (12) | 7.7 | 21 |
Isolectotype of |
(43) 45.9–80.4 (107) | 64 | (4.6) 5.3–10.6 (12.4) | 7.3 | 20 |
Spirin 2877 | (35) 42.6–80 (80) | 58.4 | (4) 4.8–7.9 (8) | 6.2 | 20 |
Spirin 9350 | (41) 44.8–83.2 (86) | 61.8 | (4.6) 4.7–10 (10.7) | 7.2 | 20 |
Spirin 9581 | (49) 51.8–84.1 (86) | 64.6 | (4.9) 5–9 (11) | 7.1 | 20 |
Spirin 9994 | (45) 45.8–75.3 (81) | 58.9 | (5.3) 5.6–10.2 (10.8) | 7.8 | 20 |
Isotype of |
(48) 51–95 (104) | 75.8 | (4.1) 6–12 (14) | 8.4 | 25 |
Argentina. Tierra del Fuego: Dpto. Ushuaia, Estancia Moat, on
For a detailed description and illustration, see
Sri Lanka. Peradeniya, on rotten branch, T.Petch (holotype BPI [305211]).
Basidiocarp of
Germany. Nordrhein-Westfalen, Lengerich, W.Brinkmann (lectotype L [L 0053271], designated by
=
=
Basidiocarps annual, resupinate, up to 5 cm in widest dimension. Margin poorly differentiated, pruinose. Hymenial surface greyish-white or pale cream-coloured, grandinioid to odontoid; projections rather regularly arranged, from 100 µm to 250 µm high, 80–100 μm broad at base, 6–8 per mm. Hyphal structure monomitic, hyphae clamped, faintly cyanophilous, thin-walled. Subicular hyphae interwoven and frequently branched, 2.2–4.7(–6.1) μm in diam. (n=60/6). Tramal hyphae subparallel, subhymenial hyphae short-celled, 2.0–3.5(–3.9) μm in diam. (n=60/6). Stellate crystals abundant in trama, subiculum and subhymenium, 4.4–8.3 μm in diam. Cystidia large, thin-walled, of subicular, tramal or subhymenial origin, clavate to spathuliform, often with an intercalary inflation, sometimes slightly thick-walled (wall not exceeding 1 μm thick), rarely forked, (35.0–)44.0–84.0(–107.0)×(4.0–)4.9–10.9(–12.4) μm (n=121/6), occasionally bearing 1–2 clamped septa. Basidia suburniform, 4-spored, (12.0–)14.0–20.8(–24.0)×3.4–4.2(–5.5) μm (n=60/6), thin-walled. Basidiospores clearly thick-walled, ellipsoid to broadly ellipsoid, usually with an oil-drop, (4.0–)4.5–5.9(–7.0)×(3.3–)3.7–4.8(–5.7) μm (n=192/6), L=5.09, W=4.12, Q=1.24, cyanophilous.
Cystidial elements of
Basidiocarp of
Europe (Estonia, Finland, Germany, Norway, Russia – up to Ural Mts.), North America, on medium-decayed wood of angiosperms.
. The type specimen of
Argentina. Chubut. Dpto Languiñeo, Lago Engaño, on
For a detailed description and illustration, see
Brazil. S. Salvador, 5 Apr 1944, Rick 20847 (holotype PACA, by original designation).
For a description, see
RUSSIA. Primorie: Khasan Dist., Kedrovaya Pad Nat. Res., on angiosperm wood, 24 Jul 2016, I.Viner KUN 1989* (H) – ITS sequence, GenBank
Ussuriensis (lat., adj.) – from the river Ussuri in Russian Far East and adjacent China.
Basidiocarps effused, up to 10 cm in longest dimension. Sterile margin white to pale ochraceous, floccose, up to 1 mm wide. Hymenial surface pale ochraceous, grandinioid to odontoid; projections rather regularly arranged, from 100 µm to 250 μm high, 90–110 μm broad at base, 6–8(–9) per mm. Hyphal structure monomitic, hyphae clamped, faintly cyanophilous, thin-walled. Subicular hyphae interwoven, (3.0–)3.4–6.2 μm in diam. (n=30/3). Tramal hyphae subparallel, subhymenial hyphae short-celled, 1.9–3.9 μm in diam. (n=30/3). Large rhomboid or stellate crystals rarely present in trama and subiculum, 10–19 μm in diam. Cystidia of three types: a) large, thin- or fairly thick-walled (wall up to 2.8 μm thick) cystidia of subicular, tramal or subhymenial origin, cylindrical, spathuliform, almost capitate or with one intercalary inflation at the upper part, (64.0–)71.0–188.9(–220.0)×(5.0–)5.7–9.4(–11.9) μm (n=30/3), often apically encrusted by large rhomboid crystals, b) astrocystidia of subhymenial origin, bearing a stellate crystalline cap 15–17×4.5–4.8 μm, sometimes rare, c) cystidia of subhymenial origin, thin-walled, varying from fusoid to cylindrical or submoniliform, rarely forked, 40.0–84.0(–92.0)×5.0–9.0(–11.4) μm (n=30/3). Basidia suburniform, 4-spored, 14.7–22.8(–24.0)×3.4–4.9 μm (n=30/3), thin-walled. Basidiospores clearly thick-walled, ellipsoid to broadly ellipsoid, usually with an oil-drop, (4.8–)5.1–6.0×3.8–4.6 μm (n=92/3), L=5.48, W=4.21, Q=1.30, cyanophilous.
East Asia (Russian Far East – Primorie), on decayed angiosperm wood; seemingly not rare in secondary oak-dominated forest.
The distinctly thick-walled tubular cystidia of
Our study confirms the results from
Blasting
For the phylogenetic analyses of
Curators of herbaria S, GB, BPI, TAAM and BAFC sent us types and other herbarium specimens used in the present study. The first author is grateful to the Kedrovaya Pad Nature Reserve staff, in particular, to Gleb Sedash and Dina