Research Article |
Corresponding author: Karl-Henrik Larsson ( k.h.larsson@nhm.uio.no ) Academic editor: R. Henrik Nilsson
© 2018 Ilya Viner, Viacheslav Spirin, Lucie Zíbarová, Karl-Henrik Larsson.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Viner I, Spirin V, Zíbarová L, Larsson K-H (2018) Additions to the taxonomy of Lagarobasidium and Xylodon (Hymenochaetales, Basidiomycota). MycoKeys 41: 65-90. https://doi.org/10.3897/mycokeys.41.28987
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Lagarobasidium is a small genus of wood-decaying basidiomycetes in the order Hymenochaetales. Molecular phylogenetic analyses have either supported Lagarobasidium as a distinct taxon or indicated that it should be subsumed under Xylodon, a genus that covers the majority of species formerly placed in Hyphodontia. We used sequences from the ITS and nuclear LSU regions to infer the phylogenetic position of the type species L. detriticum. Analyses confirm Lagarobasidium as a synonym of Xylodon. Molecular and morphological information show that the traditional concept of L. detriticum covers at least two species, Xylodon detriticus from Europe and X. pruinosus with known distribution in Europe and North America. Three species currently placed in Lagarobasidium are transferred to Xylodon, viz. X. magnificus, X. pumilius and X. rickii. Three new Xylodon species are described and illustrated, X. ussuriensis and X. crystalliger from East Asia and X. attenuatus from the Pacific Northwest America. The identity of X. nongravis, described from Sri Lanka, is discussed.
Agaricomycetes , Hyphodontia , ITS, LSU, phylogeny
The genus Lagarobasidium was introduced by
Controversies over the taxonomic position of Peniophora detritica emerged during the last decades. In modern morphology-based systems, it was first attributed to Hyphodontia J. Erikss., mainly due to hyphal characters and the shape of basidia (
In the present study, we revise the Lagarobasidium detriticum complex based on morphological and molecular methods. We propose to consider Lagarobasidium as a later synonym of Xylodon and to restore Odontia pruinosa as an independent species. In addition, we describe three new Xylodon species and make five new combinations.
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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Hastodontia hastata (Litsch.) Hjortstam & Ryvarden | Larsson 14646 | MH638232 | MH638232 | this study |
Lyomyces allantosporus Riebesehl, Yurchenko & E. Langer | FR-0249548, Holotype | KY800397 | KY795963 |
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Lyomyces crustosus (Pers.) P. Karst. | Larsson 11731 | DQ873614 | DQ873614 |
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Lyomyces erastii (Saaren. & Kotir.) Hjortstam & Ryvarden | MA-Fungi 34,336 | JX857800 |
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Lyomyces griseliniae (G. Cunn.) Riebesehl & E. Langer | Larsson 12971 | DQ873651 |
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Lyomyces mascarensis Riebesehl, Yurchenko & E. Langer | KAS-GEL4833, Holotype | KY800399 | KY795964 |
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Lyomyces microfasciculatus (Yurchenko & Sheng H. Wu) Riebesehl & E. Langer | TNM F24757, Holotype | JN129976 |
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Lyomyces organensis Yurchenko & Riebesehl | MSK7247, Holotype | KY800403 | KY795967 |
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Lyomyces orientalis Riebesehl, Yurchenko & E. Langer | KAS-GEL3400 | DQ340326 | DQ340353 |
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Lyomyces pruni (Lasch) Riebesehl & E. Langer | Ryberg 021018 | DQ873624 | DQ873625 |
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Lyomyces sambuci (Pers.) P. Karst. | KAS-GEL2414 | KY800398 |
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KAS-JR7 | KY800402 | KY795966 |
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Lyomyces vietnamensis (Yurchenko & Sheng H. Wu) Riebesehl & E. Langer | TNM F973, Holotype | JX175044 |
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Palifer verecundus (G. Cunn.) Stalpers & P.K. Buchanan | Larsson 12261 | DQ873642 |
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Xylodon apacheriensis (Gilb. & Canf.) Hjortstam & Ryvarden | Canfield 180, Holotype | KY081800 |
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Xylodon asperus (Fr.) Hjortstam & Ryvarden | H6013167 | UDB031926 | Unpublished | |
KG Nilsson s. n. | DQ873606 | DQ873607 |
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UC2023169 | KP814365 |
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Xylodon astrocystidiatus (Yurchenko & Sheng H. Wu) Riebesehl, Yurchenko & E. Langer | Wu 9211-71 | JN129972 | JN129973 |
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Xylodon attenuatus Spirin & Viner | Spirin 8775, Holotype | MH324476 | this study | |
Xylodon borealis (Kotir. & Saaren.) Hjortstam & Ryvarden | Spirin 9416 | MH317760 | MH638259 | this study |
TU115575 | UDB016473 | Unpublished | ||
UC2022850 | KP814307 |
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KUN2352 | MH307753 | MH638263 | this study | |
TU115495 | UDB016350 | Unpublished | ||
TU124171 | UDB028164 | Unpublished | ||
Xylodon bubalinus (Min Wang, Yuan Y. Chen & B.K. Cui) C.C. Chen & Sheng H. Wu | Cui 12887 | KY290982 |
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Xylodon chinensis (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu | Wu 1307-42 | KX857802 |
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Wu 1407-105, Holotype | KX857804 |
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Xylodon crystalliger Viner | KUN2312, Holotype | MH324477 | this study | |
Xylodon detriticus (Bourdot) Viner & Spirin | Zíbarová 30.10.17 | MH320793 | MH651372 | this study |
Zíbarová 26.05.17 | MH320794 | MH638264 | this study | |
Xylodon flaviporus (Berk. & M.A. Curtis ex Cooke) Riebesehl & E. Langer | ICMP13836 | AF145585 |
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Xylodon hastifer (Hjortstam & Ryvarden) Hjortstam & Ryvarden | Ryvarden 19767, Holotype | KY081801 |
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Xylodon heterocystidiatus (H.X. Xiong, Y.C. Dai & Sheng H. Wu) Riebesehl, Yurchenko & E. Langer | Wu 9209-27 | JX175045 |
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Xylodon lenis Hjortstam & Ryvarden | Wu 0808-32 | JX175043 | KX857820 |
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Wu 890714-3, Holotype | KY081802 |
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Xylodon mollissimus (L.W. Zhou) C.C. Chen & Sheng H. Wu | LWZ20160318-3, Holotype | KY007517 |
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Xylodon nespori (Bres.) Hjortstam & Ryvarden | B Nordén 030915 | DQ873622 |
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GEL3158 | DQ340310 | DQ340346 |
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GEL3290 | DQ340309 | Unpublished | ||
GEL3302 | DQ340308 | Unpublished | ||
GEL3309 | DQ340307 | DQ340345 |
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Xylodon niemelaei (Sheng H. Wu) Hjortstam & Ryvarden | GC 1508-146 | KX857798 |
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GEL4998 | EU583422 | DQ340348 |
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Wu 1010-62 | KX857799 |
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Xylodon nongravis (Lloyd) Spirin & Viner | CHWC1506-2 | KX857800 |
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Dai 11686 | KT989968 |
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GC1412-22 | KX857801 |
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Spirin 5763 | MH324469 | MH656724 | this study | |
Xylodon nothofagi (G. Cunn.) Hjortstam & Ryvarden | PDD:91630 | GQ411524 |
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Xylodon ovisporus (Corner) Riebesehl & E. Langer | ICMP13837 | AF145587 |
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KUC20130725-29 | KJ668513 | KJ668365 |
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Wu 0809-76 | KX857803 |
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Xylodon paradoxus (Schrad.) Chevall. | FCUG 1517 | AF145572 |
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FCUG 2425 | AF145571 |
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Miettinen 7978 | FN907912 | FN907912 |
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Xylodon pruinosus (Bres.) Spirin & Viner | Larsson 14653 | UDB024816 | Unpublished | |
Spirin 2877 | MH332700 | this study | ||
UC2023108 | KP814412 |
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Xylodon pseudotropicus (C.L. Zhao, B.K. Cui & Y.C. Dai) Riebesehl, Yurchenko & E. Langer | Dai 10768, Holotype | KF917543 |
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Xylodon quercinus (Pers.) Gray | Kotiranta 27060 | MH320792 | this study | |
Larsson 11076 | KT361633 | AY586678 |
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Miettinen 15050,1 | KT361632 |
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Spirin 8565 | MH316007 | this study | ||
Spirin 8840 | MH320791 | this study | ||
Xylodon raduloides (Pers.) Riebesehl & E. Langer | Dai 12631 | KT203307 | KT203328 |
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ICMP13833 | AF145580 |
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Xylodon ramicida Spirin & Miettinen | Spirin 7664, Holotype | KT361634 |
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Xylodon reticulatus (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu | GC 1512-1 | KX857808 |
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Wu 1109-178, Holotype | KX857805 |
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Xylodon rhizomorphus (C.L. Zhao, B.K. Cui & Y.C. Dai) Riebesehl, Yurchenko & E. Langer | Dai 12354 | KF917544 |
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Xylodon rimosissimus (Peck) Hjortstam & Ryvarden | CFMR:DLL2011-081 | KJ140600 |
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Ryberg 021031 | DQ873627 | DQ873628 |
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UC2022842 | KP814311 |
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UC2023109 | KP814414 |
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UC2023147 | KP814193 |
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UC2023148 | KP814194 |
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Xylodon spathulatus (Schrad.) Kuntze | GEL2690 | KY081803 |
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Larsson 7085 | KY081804 |
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Xylodon subtropicus (C.C. Chen & Sheng H. Wu) C.C. Chen & Sheng H. Wu | Wu 1508-2 | KX857806 |
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Wu 9806-105, Holotype | KX857807 |
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Xylodon ussuriensis Viner | KUN1989, Holotype | MH324468 | this study |
We inferred phylogenetic trees with maximum likelihood (ML), maximum parsimony (MP) and Bayesian Inference (BI) but provide only the last one since all trees show congruity of the phylogenetic signal. Substitution models were determined with the aid of TOPALi 2.5 (
Maximum-likelihood (ML) analysis was performed in RAxML 7.2.8 (
Maximum parsimony (MP) analysis was performed using MEGA 7 (
Xylodon attenuatus. USA. Washington: Clallam Co., La Push, Pseudotsuga menziesii, 8 Oct 2014, Spirin 8286a (H), Sol Duc, Tsuga heterophylla, 6 Oct 2014, Spirin 8133 (H); Jefferson Co., Hoh River, Acer macrophyllum, 20 Oct 2014, Spirin 8775* (H, holotype), Tsuga heterophylla, 20 Oct 2014, Spirin 8779 (H); Pend Oreille Co., Gypsy Meadows, Picea engelmannii, 17 Oct 2014, Spirin 8694* (H). Canada. British Columbia: Fraser-Fort George Reg. Dist., Mt. Robson Provincial Park, Picea sp., 25 Jul 2015, Spirin 8900a (H).
X. borealis. Russia. Nizhny Novgorod Reg.: Lukoyanov Dist., Panzelka, Quercus robur (very rotten log), 17 Aug 2015, Spirin 9416* (H).
X. brevisetus. Russia. Moscow: Losiny Ostrov Nat. Park, log of Pinus sylvestris, 1 Oct 2016, A.Nechaev KUN2352* (H).
X. crystalliger. Russia. Primorie: Khasan Dist., Kedrovaya Pad Nat. Res., on angiosperm wood, 25 Jul 2016, I.Viner KUN 2312* (H, holotype); ibidem 29 Jul 2017, F.Bortnicov, KUN 3347 (H).
X. detriticus. Czech Republic. Karlovarský kraj: Sokolov, Antonín mine spoil, on Phragmites australis, 26 May 2017, L.Zíbarová (H*); Liberecký kraj: Liberec, Uhelná, on Calamagrostis epigejos, 30 Oct 2017, L.Zíbarová (H*). France. Auvergne: Allier, St. Priest, on fern, 1 Sep 1909, H.Bourdot 7226 (S F204453, lectotype of Peniophora detritica). Italy. Lazio: Circeo Nat. Park, on Pinus pinea bark, 23 Oct 1984, K.H.Larsson 5496 (GB); ibidem, on fallen leaves, 24 Oct 1984, K.H.Larsson 5622 (GB); ibidem, on ferns, 24 Oct 1984, K.H.Larsson 5627 (GB).
X. magnificus. Argentina. Tierra del Fuego: Ushuaia, Estancia Moat, on Drimys winteri, 21 Mar 1998, A.Greslebin 1387 (GB, paratype duplicate).
X. nongravis. Russia. Khabarovsk Reg.: Khabarovsk Dist., Ulun, on Salix schwerinii, 25 Aug 2012, V.Spirin 5615 (H); ibidem, on Corylus mandshurica, 28 Aug 2012, V.Spirin 5763* (H); Primorie Reg.: Krasnoarmeiskii Dist., Melnichnoe, on Corylus mandshurica, 21–23 Aug 2013, V.Spirin 6218, 6260, 6281 (H). Sri Lanka. Peradeniya, on rotten branch, T.Petch (BPI US0305211, holotype of Polyporus nongravis).
X. pruinosus. Estonia. Ida-Virumaa: Kohtla-Järve, Pärnassaare, on Betula pubescens, 1 Oct 1958, E.Parmasto (TAAM, holotype of Hyphodontia nikolajevae). Finland. Helsinki: Veräjämäki, on Salix caprea, 4 Sep 2011, O.Miettinen 14651.4 (H). Germany. Nordrhein-Westfalen, on Betula sp., W.Brinkmann (S F204462, isolectotype of Odontia pruinosa). Norway. Akershus: Frogn, decaying deciduous wood, 3 Oct 2010, K.H.Larsson 14653* (O). Russia. Nizhny Novgorod Reg.: Bogorodsk Dist., Krastelikha, on Quercus robur, 11 Aug 2009, V.Spirin 2877* (H); Lukoyanov Dist., Panzelka, on Populus tremula, 19 Aug 2015, V.Spirin 9581 (H); Razino, on Quercus robur, 16 Aug 2015, V.Spirin 9350 (H); Srednii, on Tilia cordata, 18 Aug 2006, V.Spirin 2601 (H); Pavlovo Dist., Chudinovo, on Populus tremula, 3 Oct 2015, V.Spirin 9994 (H); Sverdlovsk Reg.: Nizhnisereginskii Dist., Olenii Ruchii Nat. Park, on Populus tremula, 19–20 Aug 2002, H.Kotiranta 19684b, 19687, 19715a (H). USA. New York: Franklin County, Paul Smith’s, on Populus tremuloides, 12 Sep 1965, R.L.Gilbertson 5481 (GB, isotype of Hyphodontia magnacystidiata).
X. pumilius. Argentina. Chubut: Río Senguer, Lago La Plata, on Nothofagus pumilio, 26–28 Mar 1996, A.Greslebin 701 (GB, paratype duplicate).
X. quercinus. Canada. Alberta: Yellowhead Co., William A. Switzer Prov. Park, on Populus tremuloides, 24 Jul 2015, V.Spirin 8840* (H). Finland. Uusimaa: Helsinki, Veräjänmäki, on angiosperm wood, 12 Apr 2008, O.Miettinen 12409* (H). Russia. Chukotka: Anadyr, on Alnus fruticosa, 19 Sep 2009, H.Kotiranta 27060* (H). USA. Washington: Pend Oreille Co., Slate Creek, on Corylus cornuta, 15 Oct 2014. V.Spirin 8565* (H).
X. rickii. Brazil. Rio Grande do Sul: S. Salvador, 5 Apr 1944, J.Rick 20847 (O, isotype of Odontia polycystidifera).
X. ussuriensis. Russia. Primorie: Khasan Dist., Kedrovaya Pad Nat. Res., angiosperm wood, 24 Jul 2016, I.Viner KUN 1989* (H, holotype of Xylodon ussuriensis), I.Viner KUN 2103, 2186.
For both datasets, the Bayesian inference returned trees with two main clades (Figures
Phylogenetic relationships of Xylodon inferred from ITS and LSU sequences using Bayesian analysis. A 50% majority rule consensus phylogram. Bayesian posterior probabilities, ML bootstrap and MP bootstrap values are shown on nodes; branch lengths reflect estimated number of changes per site.
In the ITS-only tree, three terminal branches represent new species that are described below. Xylodon attenuatus occurs as a sister taxon to X. rimosissimus; X. crystalliger forms a subclade with X. astrocystidiatus, X. paradoxus and X. heterocystidiatus; and X. ussuriensis is the sister taxon to X. detriticus and X. pruinosus (Figure
The results allow us to introduce new species and new combinations as follows.
USA. Washington: Jefferson Co., Hoh River, on Acer macrophyllum, 20 Oct 2014, V.Spirin 8775 (H) – ITS sequence, GenBank MH324476.
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).
Xylodon attenuatus bears morphological similarity to X. borealis, although densely arranged hyphae, star-like crystals and a regular presence of cystidia with granular contents make it easily recognisable. The crystalline caps on hyphoid cystidia are other characteristics useful for the identification of X. attenuatus.
RUSSIA. Primorie: Khasan Dist., Kedrovaya Pad Nat. Res., on angiosperm wood, 25 Jul 2016, I.Viner KUN 2312 (H) – ITS sequence, GenBank MH324477.
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 X. crystalliger and make it reminiscent of Xylodon astrocystidiatus (Yurchenko & Sheng H. Wu) Riebesehl, Yurchenko & Langer. The latter species is known from Taiwan and differs from X. crystalliger by having longer basidiospores and presence of constricted and bladder-like hymenial cystidia.
Peniophora detritica Bourdot, Revue Scientifique du Bourbonnais et du Centre de la France 23: 13. 1910. ≡ Lagarobasidium detriticum (Bourdot) Jülich, Persoonia 10: 334. 1979. Type. France. Auvergne: Allier, St. Priest, fern, 1.IX.1909 Bourdot 7226 (lectotype S! [F204453], designated by
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.
. Europe (Czech Republic, France, Italy), on herbaceous remnants, once collected from pine bark at the same spot where it was found on fern remains.
Species / specimen | L' | L | W' | W | Q' | Q | n |
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Xylodon attenuatus | (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 |
Xylodon crystalliger | (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 |
Xylodon detriticus | (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 |
Xylodon pruinosus | (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 Hyphodontia nikolajevae | (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 Odontia pruinosa | (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 Hyphodontia magnacystidiata | (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 |
Xylodon ussuriensis | (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 Xylodon detriticum and X. pruimosus.
Species / specimen | L' | L | W' | W | n |
---|---|---|---|---|---|
Xylodon detriticus | (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 |
Xylodon pruinosus | (35) 44–84 (107) | 61.9 | (4) 4.9–10.9 (12.4) | 7.2 | 146 |
Holotype of Hyphodontia nikolajevae | (41) 43–95 (99) | 64 | (4) 5–12 (12) | 7.7 | 21 |
Isolectotype of Odontia pruinosa | (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 Hyphodontia magnacystidiata | (48) 51–95 (104) | 75.8 | (4.1) 6–12 (14) | 8.4 | 25 |
Xylodon detriticus grows on ferns and grasses, developing thin farinaceous basidiocarps. The species evidently has a more southern distribution than X. pruinosus. Earlier reports of X. detriticus from woody substrates should be treated with caution and may represent X. pruinosus or as yet undescribed taxa.
Hyphodontia magnifica Gresl. & Rajchenb., Mycologia 92: 1160. 2000.
Argentina. Tierra del Fuego: Dpto. Ushuaia, Estancia Moat, on Drimys winteri, 21 Mar 1998, M. Rajchenberg 11370 (holotype: BAFC [50038], by original designation).
For a detailed description and illustration, see
Polyporus nongravis Lloyd, Mycol. Writings 6 (61): 891. 1919.
Sri Lanka. Peradeniya, on rotten branch, T.Petch (holotype BPI [305211]).
Odontia pruinosa Bres., Annales Mycologici 18 (1–3): 43. 1920. ≡ Lagarobasidium pruinosum (Bres.) Jülich, Persoonia 8: 84. 1974.
Germany. Nordrhein-Westfalen, Lengerich, W.Brinkmann (lectotype L [L 0053271], designated by
= Hyphodontia nikolajevae Parmasto, Conspectus Systematis Corticiacearum: 213. 1968. Type: Estonia. Ida-Virumaa, Kohtla-Järve, Pärnassaare, on Betula pubescens, 1 Oct 1958, E.Parmasto (holotype: TAAM [9683], by original designation).
= Hyphodontia magnacystidiata Lindsey & Gilb., Mycotaxon 5: 315. 1977. Type: USA. New York, Franklin County, Paul Smith’s, on Populus tremuloides, 12 Sep 1965, R.L.Gilbertson 5481 (holotype: BPI [266395], by original designation).
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.
Europe (Estonia, Finland, Germany, Norway, Russia – up to Ural Mts.), North America, on medium-decayed wood of angiosperms.
. The type specimen of Hyphodontia nikolajevae Parmasto reveals no essential differences from the type and other collections of X. pruinosus studied by us. On average, Xylodon pruinosus has wider basidiospores than X. detriticus (Table
Hyphodontia pumilia Gresl. & Rajchenb., Mycologia 92: 1162. 2000.
Argentina. Chubut. Dpto Languiñeo, Lago Engaño, on Nothofagus pumilio, 19 Apr 1996, A.Greslebin 650 (holotype BAFC [50031], by original designation).
For a detailed description and illustration, see
Hypochnicium rickii Hjortstam & Ryvarden, Mycotaxon 15: 271. 1982. ≡ Odontia polycystidifera Rick, Iheringia, Sér. Bot. 5: 163. 1959. Nom. inval. (Code Art. 40.1).
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 MH324468.
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 X. ussuriensis make it different from other Lagarobasidium-like species treated here. Subhymenial astrocystidia found in X. ussuriensis are also present in some specimens of X. detriticus although they are apparently rare in the latter species.
Our study confirms the results from
Blasting FM876212 against public sequence databases does not return any reliable results, which, if the sequence is correct, suggests that the species does not belong in Xylodon. Remaining species referred to Lagarobasidium and not already discussed include L. cymosum (D.P. Rogers & H.S. Jacks.) Jülich and L. subdetriticum (S.S. Rattan) J. Kaur & Dhingra. The former has been placed in Hypochnicium because of the thick-walled basidiospores but numerous subulate cystidia makes it a deviating element in that genus. Only access to sequence information can disclose its relationships. Lagarobasidium subdetriticum was originally described in Hyphodontia and should be retained in that genus also when the genus is taken in a restricted sense (
For the phylogenetic analyses of Hyphodontia sensu lato, only nuclear ribosomal genes have so far been applied. All published results confirm that Hyphodontia sensu lato is polyphyletic and that most species can be referred to one of three clusters, viz Hyphodontia sensu stricto, the Kneiffiella cluster and the Xylodon cluster (including Lyomyces). Within these clusters the relationships are not well resolved when the ribosomal genes are the sole source for genetic information. On such detailed level, analyses become highly sensitive to sampling and outgroup choice. It is clear that both a wider sampling and more markers must be included in analyses in order to establish a stable genus level classification for all species that have been referred to Hyphodontia in a wide sense.
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 Matyukhina. We also thank Eugeny Antonov and Fedor Bortnicov (Moscow) for their assistance during fieldwork and providing valuable fungal collections.