The genus Fomitopsis (Polyporales, Basidiomycota) reconsidered

Abstract

Based on seven- and three-gene datasets, we discuss four alternative approaches for a reclassification of Fomitopsidaceae (Polyporales, Basidiomycota). After taking into account morphological diversity in the family, we argue in favour of distinguishing three genera only, viz. Anthoporia, Antrodia and Fomitopsis. Fomitopsis becomes a large genus with 128 accepted species, containing almost all former Fomitopsis spp. and most species formerly placed in Antrodia, Daedalea and Laccocephalum. Genera Buglossoporus, Cartilosoma, Daedalea, Melanoporia, Neolentiporus, alongside twenty others, are treated as synonyms of Fomitopsis. This generic scheme allows for morphologically distinct genera in Fomitopsidaceae, unlike other schemes we considered. We provide arguments for retaining Fomitopsis and suppressing earlier (Daedalea, Caloporus) or simultaneously published generic names (Piptoporus) considered here as its synonyms. Taxonomy of nine species complexes in the genus is revised based on ITS, ITS + TEF1, ITS + TEF1 + RPB1 and ITS + TEF1 + RPB2 datasets. In total, 17 species are described as new to science, 26 older species are reinstated and 26 currently accepted species names are relegated to synonymy. A condensed identification key for all accepted species in the genus is provided. Taxonomic novelties: New species: Fomitopsis algumicola Grebenc & Spirin, F. caseosa Vlasák & Spirin, F. cupressicola Vlasák, J. Vlasák Jr. & Spirin, F. derelicta Vlasák & Spirin, F. dollingeri Vlasák & Spirin, F. fissa Vlasák & Spirin, F. lapidosa Miettinen & Spirin, F. lignicolor Vlasák & Spirin, F. maculosa Miettinen & Spirin, F. pannucea Runnel & Spirin, F. perhiemata Viner & Spirin, F. purpurea Spirin & Ryvarden, F. retorrida Spirin & Kotiranta, F. solaris Rivoire, A.M. Ainsworth & Vlasák, F. tristis Miettinen & Spirin, F. tunicata Miettinen & Spirin, F. visenda Miettinen & Spirin. New combinations: Fomitopsis aculeata (Cooke) Spirin & Miettinen, F. aethalodes (Mont.) Spirin, F. alaskana (D.V. Baxter) Spirin & Vlasák, F. albidoides (A. David & Dequatre) Bernicchia & Vlasák, F. amygdalina (Berk. & Ravenel) Spirin & Vlasák, F. angusta (Spirin & Vlasák) Spirin & Vlasák, F. atypa (Lév.) Spirin & Vlasák, F. caespitosa (Murrill) Spirin & Miettinen, F. calcitrosa (Spirin & Miettinen) Spirin & Miettinen, F. circularis (B.K. Cui & Hai J. Li) Spirin, F. concentrica (G. Cunn.) M.D. Barrett, F. cyclopis (Miettinen & Spirin) Miettinen & Spirin, F. dickinsii (Berk. ex Cooke) Spirin, F. elevata (Corner) Spirin & Miettinen, F. eucalypti (Kalchbr.) Spirin, F. ferrea (Cooke) Spirin & Viner, F. flavimontis (Vlasák & Spirin) Vlasák & Spirin, F. foedata (Berk.) Spirin & Miettinen, F. gilvidula (Bres.) Spirin & Miettinen, F. glabricystidia (Ipulet & Ryvarden) Miettinen & Ryvarden, F. globispora (Ryvarden & Aime) Spirin, F. hartmannii (Cooke) M.D. Barrett & Spirin, F. hyalina (Spirin, Miettinen & Kotir.) Spirin & Miettinen, F. hypoxantha (Bres.) Spirin & Miettinen, F. incana (Lév.) Spirin & V. Malysheva, F. infirma (Renvall & Niemelä) Miettinen & Niemelä, F. juniperina (Murrill) Spirin & Vlasák, F. kuzyana (Pilát ex Pilát) Spirin & Vlasák, F. leioderma (Mont.) Spirin & Vlasak, F. leucaena (Y.C. Dai & Niemelä) Spirin & Miettinen, F. luzonensis (Murrill) Spirin & Miettinen, F. maculatissima (Lloyd) Spirin, F. madronae (Vlasák & Ryvarden) Vlasák & Ryvarden, F. malicola (Berk. & M.A. Curtis) Spirin, F. marchionica (Mont.) Spirin & Miettinen, F. marianii (Bres.) Spirin, Vlasák & Cartabia, F. mellita (Niemelä & Penttilä) Niemelä & Miettinen, F. microcarpa (B.K. Cui & Shun Liu) Spirin, F. micropora (B.K. Cui & Shun Liu) Spirin, F. modesta (Kuntze ex Fr.) Vlasák & Spirin, F. monomitica (Yuan Y. Chen) Spirin & Viner, F. morganii (Lloyd) Spirin & Vlasák, F. moritziana (Lév.) Spirin & Miettinen, F. neotropica (D.L. Lindner, Ryvarden & T.J. Baroni) Vlasák, F. nigra (Berk.) Spirin & Miettinen, F. nivosella (Murrill) Spirin & Vlasák, F. oboensis (Decock, Amalfi & Ryvarden) Spirin, F. oleracea (R.W. Davidson & Lombard) Spirin & Vlasák, F. philippinensis (Murrill) Spirin & Vlasák, F. primaeva (Renvall & Niemelä) Miettinen & Niemelä, F. psilodermea (Berk. & Mont.) Spirin & Vlasák, F. pulverulenta (Rivoire) Rivoire, F. pulvina (Pers.) Spirin & Vlasák, F. pulvinascens (Pilát ex Pilát) Niemelä & Miettinen, F. quercina (L.) Spirin & Miettinen, F. ramentacea (Berk. & Broome) Spirin & Vlasák, F. renehenticii (Rivoire, Trichies & Vlasák) Rivoire & Vlasák, F. roseofusca (Romell) Spirin & Vlasák, F. sagraeana (Mont.) Vlasák & Spirin, F. sandaliae (Bernicchia & Ryvarden) Bernicchia & Vlasák, F. sclerotina (Rodway) M.D. Barrett & Spirin, F. serialiformis (Kout & Vlasák) Vlasák, F. serialis (Fr.) Spirin & Runnel, F. serrata (Vlasák & Spirin) Vlasák & Spirin, F. squamosella (Bernicchia & Ryvarden) Bernicchia & Ryvarden, F. stereoides (Fr.) Spirin, F. subectypa (Murrill) Spirin & Vlasák, F. substratosa (Malençon) Spirin & Miettinen, F. tropica (B.K. Cui) Spirin, F. tumulosa (Cooke) M.D. Barrett & Spirin, F. tuvensis (Spirin, Vlasák & Kotir.) Spirin & Vlasák, F. uralensis (Pilát) Spirin & Miettinen, F. ussuriensis (Bondartsev & Ljub.) Spirin & Miettinen, F. variiformis (Peck) Vlasák & Spirin, F. yunnanensis (M.L. Han & Q. An) Spirin, Daedaleopsis candicans (P. Karst.) Spirin, Megasporoporia eutelea (Har. & Pat.) Spirin & Viner, Neofomitella hemitephra (Berk.) M.D. Barrett, Pseudophaeolus soloniensis (Dubois) Spirin & Rivoire, P. trichrous (Berk. & M.A. Curtis) Vlasák & Spirin. New synonyms: Antrodia bondartsevae Spirin, A. huangshanensis Y.C. Dai & B.K. Cui, A. taxa T.T. Chang & W.N. Chou, A. wangii Y.C. Dai & H.S. Yuan, Antrodiella subnigra Oba, Mossebo & Ryvarden, Antrodiopsis Audet, Boletus quercinus Schrad., Brunneoporus Audet, Buglossoporus Kotl. & Pouzar, Buglossoporus eucalypticola M.L. Han, B.K. Cui & Y.C. Dai, Caloporus P. Karst., Cartilosoma Kotlaba & Pouzar, Coriolus clemensiae Murrill, C. cuneatiformis Murrill, C. hollickii Murrill, C. parthenius Hariot & Pat., C. rubritinctus Murrill, Daedalea Pers., Daedalea allantoidea M.L. Han, B.K. Cui & Y.C. Dai, D. americana M.L. Han, Vlasák & B.K. Cui, D. radiata B.K. Cui & Hai J. Li, D. rajchenbergiana Kossmann & Drechsler-Santos, D. sinensis Lloyd, Daedalella B.K. Cui & Shun Liu, Dentiporus Audet, Flavidoporia Audet, Fomes subferreus Murrill, Fomitopsis cana B.K. Cui, Hai J. Li & M.L. Han, F. caribensis B.K. Cui & Shun Liu, F. cystidiata B.K. Cui & M.L. Han, F. ginkgonis B.K. Cui & Shun Liu, F. iberica Melo & Ryvarden, F. incarnata K.M. Kim, J.S. Lee & H.S. Jung, F. subfeei B.K. Cui & M.L. Han, F. subtropica B.K. Cui & Hai J. Li, Fragifomes B.K. Cui, M.L. Han & Y.C. Dai, Leptoporus epileucinus Pilát, Melanoporia Murrill, Neoantrodia Audet, Neolentiporus Rajchenb., Nigroporus macroporus Ryvarden & Iturr., Niveoporofomes B.K. Cui, M.L. Han & Y.C. Dai, Pilatoporus Kotl. & Pouzar, Piptoporus P. Karst., Polyporus aurora Ces., P. durescens Overh. ex J. Lowe, P. griseodurus Lloyd, Poria incarnata Pers., Pseudoantrodia B.K. Cui, Y.Y. Chen & Shun Liu, Pseudofomitopsis B.K. Cui & Shun Liu, Ranadivia Zmitr., Rhizoporia Audet, Rhodofomes Kotl. & Pouzar, Rhodofomitopsis B.K. Cui, M.L. Han & Y.C. Dai, Rhodofomitopsis pseudofeei B.K. Cui & Shun Liu, R. roseomagna Nogueira-Melo, A.M.S. Soares & Gibertoni, Rubellofomes B.K. Cui, M.L. Han & Y.C. Dai, Subantrodia Audet, Trametes fulvirubida Corner, T. lignea Murrill, T. lusor Corner, T. pseudodochmia Corner, T. subalutacea Bourdot & Galzin, T. supermodesta Ryvarden & Iturr., T. tuberculata Bres., Tyromyces multipapillatus Corner, T. ochraceivinosus Corner, T. palmarum Murrill, T. singularis Corner, T. squamosellus Núñez & Ryvarden, Ungulidaedalea B.K. Cui, M.L. Han & Y.C. Dai. Lectotypes: Hexagonia sulcata Berk., Polyporus castaneae Bourdot & Galzin, Poria incarnata Pers., Trametes subalutacea Bourdot & Galzin, Ungulina substratosa Malençon. Neotypes: Agaricus soloniensis Dubois, Boletus pulvinus Pers. Citation: Spirin V, Runnel K, Vlasák J, Viner I, Barrett MD, Ryvarden L, Bernicchia A, Rivoire B, Ainsworth AM, Grebenc T, Cartabia M, Niemelä T, Larsson K-H, Miettinen O (2024). The genus Fomitopsis (Polyporales, Basidiomycota) reconsidered. Studies in Mycology 107: 149-249. doi: 10.3114/sim.2024.107.03.

Keywords: brown-rot fungi; new taxa; phylogeny; polypores; taxonomy.

Fig 1.

Phylogenetic placement of the Daedalea – Fomitopsis clade (box) within the “antrodia clade” (including all taxa shown other than outgroups) based on Maximum Likelihood of the ITS + LSU + RPB1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 2.

Phylogenetic placement of the Daedalea – Fomitopsis clade (box) within the Fomitopsidaceae and allied taxa (including all taxa shown other than outgroups) based on Maximum Likelihood of the ITS + LSU + SSU + mtSSU + RPB1 + RPB2 + TEF1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig. 3

Phylogenetic relationships of species in the Daedalea – Fomitopsis clade based on Maximum Likelihood of the ITS + LSU + RPB1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The species represented with ITS + LSU only are marked with an asterisk (*). The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin. Names of pink-coloured perennial species are given in red, those of monomitic species are in blue.

Fig 4.

Three generic concepts for delimiting genera in the Daedalea – Fomitopsis clade. A. Microgeneric approach. B. Mesogeneric approach. C. Macrogeneric approach (accepted in the present study). For details about topology, see Fig. 2.

Fig 5.

Phylogenetic relationships of species in the Fomitopsis (Daedalea) quercina clade based on Maximum Likelihood of the ITS dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 6.

Phylogenetic relationships of species in Pilatoporus (Fomitopsis palustris) group based on Maximum Likelihood of the ITS dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 7.

Phylogenetic relationships of species in the Fomitopsis marianii complex based on Maximum Likelihood of the ITS + TEF1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 8.

Phylogenetic relationships of species in the Fomitopsis meliae complex based on Maximum Likelihood of the ITS + TEF1 dataset. Numbers on nodes represent bootstrap values > 70 % and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 9.

Phylogenetic relationships of species in the Pilatoporus (Fomitopsis palustris) group based on Maximum Likelihood of the ITS + TEF1 + RPB1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin. The tree is midpoint-rooted.

Fig 10.

Phylogenetic relationships of species in the Fomitopsis pinicola complex based on Maximum Likelihood of the ITS + TEF1 +RPB2 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 11.

Phylogenetic relationships of species in the Fomitopsis feei clade based on Maximum Likelihood of the ITS dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 12.

Phylogenetic relationships of species in the Fomitopsis rosea clade based on Maximum Likelihood of the ITS dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 13.

Phylogenetic relationships of species in the Fomitopsis (Antrodia) ramentacea clade based on Maximum Likelihood of the TEF1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 14.

Phylogenetic relationships of species in the Fomitopsis (Antrodia) ramentacea clade based on Maximum Likelihood of the ITS + TEF1 dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin.

Fig 15.

Phylogenetic relationships of species in Fomitopsis spraguei group based on Maximum Likelihood of the ITS dataset. Numbers on nodes represent bootstrap values > 70% and Bayesian Inference posterior probabilities > 0.85. The scale bar indicates the number of expected substitutions per site. Two-letter codes in the parentheses denote the country of origin. The tree is midpoint-rooted.

Fig 16.

Anatomical structures (subicular and subhymenial hyphae, hymenial cells and basidiospores) of Fomitopsis spp. F. oleracea (Miettinen 17902), F. fissa (holotype), F. dollingeri (holotype), F. cellularis (Vlasák 1504/36J). Scale bar = 10 μm.

Fig 17.

Basidiocarps of Fomitopsis spp. A. Fomitopsis aculeata (Miettinen 8647). B. F. angusta (Spirin 10725). C. F. caespitosa (Miettinen 10227). D. F. cajanderi (Miettinen 22477). E. F. carnea (Miettinen 23610). F. F. caseosa (holotype). G. F. castanea (Spirin 5144). H. F. dickinsii (Niemelä 6435).

Fig 18.

Basidiospores of Fomitopsis spp. F. aculeata (Miettinen 8674); F. africana (Kout 1408/K9); F. algumicola (holotype); F. amygdalina (Vlasák 1707/9J); F. atypa (Ryvarden 17588); F. caespitosa (Miettinen 8737). Scale bar = 5 μm.

Fig 19.

Basidiospores of Fomitopsis spp. F. carnea (Miettinen 13120.1); F. caseosa (holotype); F. cupreorosea (Kout 0610/K4); F. cupressicola (holotype); F. derelicta (holotype); F. dochmia (Dunaev 7.II.2019). Scale bar = 5 μm.

Fig 20.

Basidiospores of Fomitopsis spp. F. dollingeri (holotype); F. elevata (Miettinen 20529); F. feei (Oinonen 60119006); F. ferrea (Dunaev 26.I.2019); F. fissa (holotype); F. foedata (Miettinen 11466). Scale bar = 5 μm.

Fig 21.

Basidiocarps of Fomitopsis spp. A. F. elevata (Miettinen 8692). B. F. foedata (Miettinen 11466). C. F. gilvidula (Miettinen 20535). D. F. lapidosa (holotype). E. F. leioderma (Vlasák 1908/82). F. F. luzonensis (Miettinen 14311). G. F. maculosa (holotype). H. F. marianii (Spirin 10575).

Fig 22.

Basidiospores of Fomitopsis spp. F. gilvidula (Miettinen 12905); F. incana (LE313649); F. lapidosa (holotype); F. leioderma (Vlasák 1908/82); F. lignea (lectotype of Fomes subferreus); F. lignicolor (holotype); F. luzonensis (Miettinen 14311). Scale bar = 5 μm.

Fig 23.

Basidiospores of Fomitopsis spp. F. maculosa (holotype); F. marchionica (Miettinen 11454); F. marianii (Spirin 10503); F. meliae (Dollinger 989); F. modesta (Vlasák 1504/52); F. moritziana (Miettinen 11662); F. nivosella (3 left – O F10833, 3 right – holotype). Scale bar = 5 μm.

Fig 24.

Basidiocarps of Fomitopsis spp. A. F. modesta (Vlasák 1808/87). B. F. moritziana (Miettinen 11662). C. F. ochracea (Miettinen 18568). D. F. ostreiformis (Miettinen 23532). E. F. perhiemata (holotype). F. F. pinicola (Niemelä 9424). G. F. pseudopetchii (Miettinen 14284). H. F. pulvina (Niemelä 9281).

Fig 25.

Basidiospores of Fomitopsis spp. F. ostreiformis (isolectotype); F. pannucea (holotype); F. perhiemata (holotype); F. pinicola (Kotiranta 26782); F. pseudopetchii (Miettinen 14284); F. psilodermea (Vlasák 1504/34). Scale bar = 5 μm.

Fig 26.

Basidiocarps of Fomitopsis spp. A. F. quercina (Niemelä 4030). B. F. roseofusca (Vlasák 1909/82). C. F. scalaris (Vlasák 1909/66). D. F. sulcata (Miettinen 11275). E. F. psilodermea (Vlasák 1504/34). F. F. tristis (holotype). G. F. tunicata (holotype). H. F. uralensis (Spirin 10946).

Fig 27.

Basidiospores of Fomitopsis spp. F. purpurea (holotype); F. quercina (Miettinen 12662); F. retorrida (holotype); F. roseofusca (Vlasák 1908/83J); F. sagraeana (O 14113); F. scalaris (Vlasák 1909/66). Scale bar = 5 μm.

Fig 28.

Basidiospores of Fomitopsis spp. F. solaris (holotype); F. subectypa (holotype); F. tristis (holotype); F. tunicata (holotype); F. uralensis (Spirin 5399); F. visenda (holotype). Scale bar = 5 μm.