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Research Article
Phylogeny and diversity of Bjerkandera (Polyporales, Basidiomycota), including four new species from South America and Asia
expand article infoChao-Ge Wang, Josef Vlasák§, Yu-Cheng Dai
‡ Beijing Forestry University, Beijing, China
§ Biology Centre, Czech Academy of Sciences, České Budějovice, Czech Republic
Open Access

Abstract

Four new species of Bjerkandera, viz. B. ecuadorensis, B. fulgida, B. minispora, and B. resupinata spp. nov., are described from tropical America and Asia. B. ecuadorensis is characterised by dark grey to black pore surface, a monomitic hyphal system, hyaline to yellowish-brown generative hyphae, and ellipsoid basidiospores measuring 3.9–4.5 × 2.7–3 μm. B. fulgida is distinguished from the other species in the genus by clay buff to pale brown and shiny pore surface. B. minispora is characterised by white tomentose pore mouth and small basidiospores measuring 3.1–4.2 × 2–2.8 μm. B. resupinata is characterised by resupinate basidiomata, pinkish buff to pale brownish pore surface, and ellipsoid to broadly ellipsoid basidiospores measuring 4.5–6 × 3.2–4.1 µm. All these new species grow on angiosperm trunks or rotten wood, and cause a white rot. The closely related taxa to four new species are discussed. An identification key to the ten accepted species of Bjerkandera is provided, and a phylogeny comprising all known Bjerkandera species is provided.

Keywords

Phylogeny, polypore, taxonomy, wood-decaying fungi

Introduction

The genus Bjerkandera P. Karst. (Polyporales, Basidiomycota), typified by B. adusta (Willd.) P. Karst., was established by Karsten (1879). It is traditionally characterised by annual, effused-reflexed to pileate basidiomata, the presence of a dark resinous layer between context and tubes, grey to black pore surface, which contrasts with the pale cream context, a monomitic hyphal system with abundant clamps on generative hyphae, oblong ellipsoid to ellipsoid, hyaline, thin-walled basidiospores, and a white-rotting ecology (Murrill 1907; Ryvarden and Gilbertson 1993; Núñez and Ryvarden 2001; Westphalen et al. 2015; Cui et al. 2019). Based on the above morphological studies, Bjerkandera and Gloeoporus Mont. share some important characteristics, and both genera are confused about definition (Pilát 1937; Corner 1989; Motato-Vásquez et al. 2020). Also, Tyromyces P. Karst. is defined by white, annual, resupinate to pileate basidiomata, a mono-dimitic hyphal system with clamped generative hyphae, and a white-rotting ecology, so Tyromyces and Bjerkandera overlap in many characteristics (Ryvarden 1991; Pouzar 1966; Núñez and Ryvarden 2001). Due to these similarities, Pouzar (1966) considered Bjerkandera as subgenus of Tyromyces (Kotlaba and Pouzar 1964). Nevertheless, recent phylogenetic analyses have showed that Bjerkandera, Gloeoporus, and Tyromyces belong to different clades in the families Phanerochaetaceae Jülich (Syn. Bjerkanderaceae Jülich 1981), Irpicaceae Spirin & Zmitr., and Incrustoporiaceae Jülich, respectively (Binder et al. 2013; Floudas and Hibbett 2015; Justo et al. 2017; Jung et al. 2018; Viktor and Bálint 2018; Cui et al. 2019; Motato-Vásquez et al. 2020). Morphologically, Tyromyces differs in pale tubes not darkening upon drying, and Gloeoporus usually has a continuum hymenium (covering the dissepiments) and gelatinous tubes (Ryvarden and Gilbertson 1993), while species of Bjerkandera have distinct sterile dissepiments, and corky to hard corky tubes.

Bjerkandera is a common polypore genus that grows mostly on dead angiosperm wood and has a wide distribution around the world. Two species, Bjerkandera adusta (Willd.) P. Karst. and B. fumosa (Pers.) P. Karst., are well recognised in the northern hemisphere (Gilbertson and Ryvarden 1986; Jung et al. 2014; Ryvarden and Melo 2017; Cui et al. 2019). Tyromyces atroalbus (Rick) Rajchenb. was combined into Bjerkandera based on morphology and molecular phylogenetic analysis (Westphalen et al. 2015). Recently, two new species, Bjerkandera albocinerea Motato-Vásq., Robledo & Gugliotta and B. centroamericana Kout, Westphalen & Tomšovský, were described from the neotropics based on morphological characters and molecular data (Westphalen et al. 2015; Motato-Vásquez et al. 2020). Additionally, B. mikrofumosa Ryvarden was described from Venezuela without molecular data (Ryvarden 2016), but DNA sequences from this species were generated by Motato-Vásquez et al. (2020).

During a study on polypores collected from China, Ecuador, and Thailand, four unknown species of Bjerkandera were distinguished by both morphological and molecular data. They are described and illustrated in this study. In this study, nuclear ribosomal RNA genes were used to determine the phylogenetic position of the new species. Furthermore, an identification key to all the accepted species in the genus is provided.

Materials and methods

Morphological studies

The studied specimens are deposited in the herbaria of the Institute of Microbiology, Beijing Forestry University (BJFC), and the private herbarium of Josef Vlasák (JV), which will later be deposited at the National Museum Prague of Czech Republic (PRM). Morphological descriptions are based on field notes and herbarium specimens. Microscopic analyses follow Miettinen et al. (2018). In the description: KOH = 5% potassium hydroxide, IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, CB = Cotton Blue, CB+ = cyanophilous in Cotton Blue, CB– = acyanophilous in Cotton Blue, L = arithmetic average of all spore length, W = arithmetic average of all spore width, Q = L/W ratios, and n = number of spores/measured from given number of specimens. Colour terms are cited from Anonymous (1969) and Petersen (1996).

Molecular studies and phylogenetic analysis

A CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain DNA from dried specimens, and to perform the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications (Shen et al. 2019; Sun et al. 2020). Two DNA gene fragments – internal transcribed spacer (ITS) and large subunit nuclear ribosomal RNA gene (nLSU) – were amplified using the primer pairs ITS5/ITS4 and LR0R/LR7 (White et al. 1990; Hopple and Vilgalys 1999) (http://www.biology.duke.edu/fungi/mycolab/primers.htm). The PCR procedures for ITS and nLSU followed Zhao et al. (2013) in the phylogenetic analyses. DNA sequencing was performed at Beijing Genomics Institute and the newly-generated sequences were deposited in GenBank (Sayers et al. 2021). Sequences generated for this study were aligned with additional sequences downloaded from GenBank using BioEdit (Hall 1999) and ClustalX (Thompson et al. 1997). The final ITS and nLSU datasets were subsequently aligned using MAFFT v.7 under the E-INS-i strategy with no cost for opening gaps and equal cost for transformations (command line: mafft –genafpair –maxiterate 1000) (Katoh and Standley 2013) and visualised in BioEdit (Hall 1999).

In this study, nuclear ribosomal RNA genes were used to determine the phylogenetic position of the new species. The sequence alignment was deposited at TreeBase (submission ID 27872). Sequences of Tyromyces chioneus (Fr.) P. Karst, obtained from GenBank, was used as outgroup (Westphalen et al. 2015).

The phylogenetic analyses followed the approach of Han et al. (2016) and Zhu et al. (2019). Maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI) analyses were conducted for the datasets of ITS and nLSU sequences. The best-fit evolutionary model was selected by hierarchical likelihood ratio tests (hLRT) and Akaike Information Criterion (AIC) in MrModeltest 2.2 (Nylander 2004) after scoring 24 models of evolution in PAUP* version 4.0b10 (Swofford 2002).

The MP topology and bootstrap values (MP-BS) obtained from 1000 replicates were computed in PAUP* version 4.0b10 (Swofford 2002). All characters were equally weighted, and gaps were treated as missing. Trees were inferred using the heuristic search option with TBR branch swapping and 1000 random sequence additions. Max-trees were set to 5,000 branches of zero length were collapsed, and all parsimonious trees were saved. Descriptive tree statistics tree length (TL), composite consistency index (CI), retention index (RI), rescaled consistency index (RC), and homoplasy index (HI) were calculated for each maximum parsimonious tree (MPT) generated. Sequences were also analysed using Maximum Likelihood (ML) with RAxML-HPC2 through the CIPRES Science Gateway (www.phylo.org; Miller et al. 2009). Branch support (BT) for ML analysis was determined by 1000 bootstrap replicates.

Bayesian phylogenetic inference and Bayesian posterior probabilities (BPP) were computed with MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). Four Markov chains were run for 3,500,000 generations until the split deviation frequency value was less than 0.01 and trees were sampled every 100 generations. The first 25% of the sampled trees were discarded as burn-in and the remaining ones were used to reconstruct a majority rule consensus and calculate Bayesian posterior probabilities (BPP) of the clades.

Branches that received bootstrap support for maximum parsimony (≥ 75% MP-BT), maximum likelihood (≥75% (ML-BS)), and Bayesian posterior probabilities (≥ 0.95BPP) were considered as significantly supported.

Results

Phylogeny

The combined ITS and nLSU dataset contained sequences from 75 specimens, comprising a total of 40 species (Table 1). The dataset had an aligned length of 2158 characters, of which 1410 (65%) characters are constant, 208 (0.1%) are variable and parsimony-uninformative and 540 (25%) are parsimony informative. Maximum parsimony analysis yielded eleven equally-parsimonious tree (TL = 2701, CI = 0.439, RI = 0.751, RC = 0.330, HI = 0.561), and a strict consensus tree of these trees is shown in Fig. 1. The best model-fit applied in the Bayesian analysis was GTR+I+G, lset nst = 6, rates = invgamma, and prset statefreqpr = dirichlet (1, 1, 1, 1). Bayesian analysis resulted in the nearly congruent topology with an average standard deviation of split frequencies = 0.006804 to MP and ML analysis, and thus only the MP tree was provided.

Table 1.

Information on the sequences used in this study. New sequences are shown in bold.

Species Specimen number Countries GenBank accession numbers
ITS LSU
Aurantiporus croceus Miettinen-16483 Malaysia KY948745 KY948901
Bjerkandera adusta Dai 14516 China MW507097 MW520204
adusta Dai 15665 China MW507098 MW520205
B. adusta Dai 15495 China MW507099
B. adusta SFC20120409-08 Rep. Korea KJ704814 KJ704829
B. adusta SFC20111029-15 Rep. Korea KJ704813 KJ704828
B. adusta Dai 13201 France MW507100 MW520206
B. adusta Dai 12640 Finland MW507101
B. albocinerea MV 346 Brazil MH025421 MH025421
B. albocinerea RP 317 Brazil MH025420
B. albocinerea Dai 16411 USA MW507102 MW520207
B. atroalba MW 425 Brazil KT305930 KT305930
B. atroalba MV 158 Brazil KT305932 KT305932
B. atroalba Dai 17457 Brazil MW507103 MW520208
B. centroamericana JK0610/A13 Mexico KT305934 KT305934
B. centroamericana JK0610/A7 Mexico KT305933 KT305933
B. centroamericana JV1704/97 Costa Rica MW507104
B. ecuadorensis JV1906/C16-J Ecuador MW507105
B. fulgida Dai 16107 China MW507106 MW520209
B. fulgida Dai 12284 China MW507107
B. fulgida Dai 13597 China MW507108 MW520210
B. fumosa SFC20121009-04 Rep. Korea KJ704824 KJ704839
B. fumosa Dai 21100 China MW507109 MW520211
B. fumosa Dai 21087 China MW507110
B. fumosa Cui 10747 China MW507111 MW520212
B. fumosa Dai 12674B Finland MW507112 MW520213
B. fumosa N37 Latvia FJ903376
B. fumosa Homble 1900 Norway KF698740 KF698751
B. mikrofumosa MV 353 Brazil MH025416 MH025416
B. mikrofumosa MV 363 Brazil MH023526 MH023526
B. mikrofumosa JV1707/10J-1 Costa Rica MW507113
B. mikrofumosa JV1707/10J-2 Costa Rica MW507114
B. minispora Dai 15234 China MW507115 MW520214
B. minispora Cui 5376 China MW507116 MW520215
B. resupinata Dai 16642 Thailand MW507117 MW520216
B. resupinata Cui 8017 China KU509526
Byssomerulius corium KHL 8593 AY463389 AY586640
Ceriporia viridans KHL 8765 AF347109 AF347109
Ceriporiopsis alboaurantia Cui 4136 China KF845955 KF845948
C. alboaurantia Cui 2877 China KF845954 KF845947
C. aneirina TAA 181186 Estonia FJ496683 FJ496704
C. aneirina H 6002107 Finland FJ496682 FJ496705
carnegieae RLG-7277-T USA KY948792 KY948854
C. carnegieae JV1209/45 USA KX081134
C. carnegieae JV0407/27-J USA MW507122
C. fimbriata Dai 11672 China KJ698633 KJ698637
C. fimbriata Cui 1671 China KJ698634 KJ698638
C. gilvescens BRNM 710166 Czech FJ496684 FJ496720
C. gilvescens BRNM 709970 Czech EU546104 FJ496721
C. pseudogilvescens BRNM 686416 Slovakia FJ496679 FJ496703
C. pseudogilvescens TAA 168233 Estonia FJ496673 FJ496702
Ceriporiopsis sp. JV1512/13-J Costa Rica MW507118
Gloeoporus taxicola SK 0075 Sweden JX109847 JX109847
G. pannocinctus FP 135015 USA MG572755 MG572739
G. thelephoroides BZ 2896 Belize MG572757 MG572741
Hapalopilus nidulans FD-512 USA KP135419
Hydnophlebia chrysorhiza FD-282 USA KP135338 KP135217
Hyphodermella corrugata KHL 3663 Norway EU118630 EU118630
Irpex lacteus DO 421951208 Sweden JX109852 JX109852
Merulius tremellosus FD-323 USA KP135231
Mycoacia fuscoatra KHL 13275 Estonia JN649352 JN649352
M. nothofagi KHL 13750 France GU480000 GU480000
Phanerochaete chrysosporium BKM-F-1767 HQ188436 GQ470643
P. sordida KHL 12054 Norway EU118653 EU118653
Phlebia nitidula GB 020830 Sweden EU118655 EU118655
P. radiata AFTOL 484 AY854087 AF287885
Phlebiopsis gigantea FP-70857-Sp USA KP135390 KP135272
Porostereum spadiceum KUC 2013051 Rep. Korea KJ668473 KJ668325
Terana caerulea FP 10473 USA KP134980 KP135276
Trametopsis cervina TJV 93216 T USA JN165020 JN164796
Tyromyces chioneus Miettinen 7487 Finland HQ659244 HQ659244
T. fissilis Dai 18182 China MW507119 MW520217
T. fissilis Dai 19583 China MW507120 MW520218
T. fissilis BRNM 699803 Czech HQ728292 HQ729002
T. fissilis Dai 19589 China MW507121
Figure 1. 

Phylogeny of Bjerkandera and related species generated by maximum parsimony analysis, based on combined ITS and nLSU sequences. Bootstrap support for maximum parsimony (MP), maximum likelihood (ML), and Bayesian posterior probabilities: (BPP) ≥ 50% (MP-BT), 50% (ML-BS) and 0.90 (BPP) are given in relation to the branches.

In our phylogeny (Fig. 1), the genus Bjerkandera was supported as a monophyletic clade, which was consistent with previous studies on monophyly nature of Bjerkandera (Westphalen et al. 2015; Motato-Vásquez et al. 2020). Bjerkandera ecuadorensis, B. fulgida, B. minispora, and B. resupinata were nested within the Bjerkandera forming four distinct lineages (95/98/1.00, 90/87/1.00, 100/100/1.00, and 60/85/0.90 respectively).

Taxonomy

Bjerkandera ecuadorensis Y.C. Dai, Chao G. Wang & Vlasák, sp. nov.

MycoBank No: 538578
Figs 2, 3

Diagnosis

Bjerkandera ecuadorensis is characterised by grey to dark-brown pore surface, tiny pores (7–9 per mm), and ellipsoid basidiospores measuring 3.9–4.5 × 2.7–3 μm.

Type

Ecuador, Pichincha Province, volcan Pasochoa, 3300 m, VI. 2019, J. Vlasák Jr. JV 1906/C16-J (holotype in PRM, isotypes in JV and BJFC032992).

Etymology

Ecuadorensis (Lat.): referring to the species being found in Ecuador.

Basidiomata

Annual, pileate, soft corky, without odor or taste when fresh, becoming corky when dry, projecting up to 4 cm, 5 cm wide and 1.3 mm thick at base. Pileal surface pinkish-buff to buff, glabrous, faintly zonate, margin blunt. Pore surface grey to dark-brown, becoming almost black when touched or bruised; sterile margin distinct, up to 2 mm wide; pores round to angular, 7–9 per mm; dissepiments thin, entire. Context buff-yellow, slightly fibrous to corky, up to 1 mm thick. Tubes concolorous with the pore surface and darker than context, corky, up to 0.3 mm long, and with a distinct dark line between tubes and context.

Figure 2. 

Pileal surface and pore surface of Bjerkandera ecuadorensis (holotype, JV 1906/C16-J). Scale bars: 4 mm.

Hyphal structure

Hyphal system monomitic; generative hyphae with clamp connections, smooth, hyaline to yellowish-brown, CB+, IKI–; tissues becoming dark in KOH.

Context

Generative hyphae thick-walled with a wide lumen, occasionally branched, densely compacted, and more or less regularly arranged to loosely interwoven, up to 3.8–6 μm in diam.

Figure 3. 

Microscopic structures of Bjerkandera ecuadorensis (holotype, JV 1906/C16-J) a basidiospores b basidia and basidioles c hyphae from trama d hyphae from context.

Tubes

Generative hyphae thin- to slightly thick-walled, rarely branched, subparallel along the tubes to loosely interwoven, 2.5–3.8 μm in diam. Cystidia and cystidioles absent. Basidia clavate to barrel-shaped, with four sterigmata and a basal clamp connection, 13–14.5 × 4.5–5.5 µm; basidioles of similar shape to basidia, but smaller.

Basidiospores

Ellipsoid, hyaline, thin-walled, smooth, often with one or more guttules, CB–, IKI–, (3.8–)3.9–4.5 × 2.7–3 µm, L = 4.09 μm, W = 2.86 μm, Q = 1.43 (n = 30/1).

Remarks

Bjerkandera ecuadorensis is characterised by grey to dark-brown pore surface, small pores (7–9 per mm), hyaline to yellowish-brown generative hyphae, and ellipsoid basidiospores measuring 3.9–4.5 × 2.7–3 μm. Morphologically, Bjerkandera ecuadorensis is similar to B. minispora in having pinkish-buff to buff pileal surface and round to angular pores (6–9 per mm), but the latter has buff-yellow pore surface and smaller basidiospores (3.1–4.2 × 2–2.8 μm). Bjerkandera adusta resembles B. ecuadorensis by having grey to dark-brown pore surface, distinct sterile margin, but the former has short-cylindric to subellipsoid and bigger basidiospores (4.5–6 × 2.5–3.5 μm, Ryvarden and Melo 2017).

Bjerkandera fulgida Y.C. Dai & Chao G. Wang, sp. nov.

MycoBank No: 838579
Figs 4, 5

Diagnosis

Bjerkandera fulgida is characterised by the clay buff to pale brown and shiny pore surface, and ellipsoid to broadly ellipsoid basidiospores measuring 3.9–4.5 × 2.8–3.3 μm.

Type

China. Hainan Province, Lingshui County, Diaoluoshan Forest Park, 18°42'N, 109°49'E, rotten angiosperm wood, 13.XI.2015, Y.C. Dai 16107 (holotype BJFC020200).

Etymology

Fulgida (Lat.): referring to the species having the shiny pore surface.

Basidiomata

Annual, effused-reflexed, soft corky, without odor or taste when fresh, becoming corky upon drying, resupinating up to 5.5 cm long, 3 cm wide and 1.3 mm thick, with a pileal projection up to 0.6 cm, 2.3 cm wide and 1.3 mm thick at base. Pileal surface pinkish buff to clay-buff, glabrous and faintly zonate when dry; margin acute. Pore surface clay-buff to pale brown, bruised part becoming dark brown to black when dry, shiny; sterile margin up to 2 mm wide; pores round or sometimes angular, 6–8 per mm; dissepiments thin, entire. Context pale cream, slightly fibrous to corky, up to 0.5 mm thick. Tubes concolorous with the pore surface, darker than context, corky, up to 0.8 mm long, with a distinct dark line between tubes and context.

Figure 4. 

Bjerkandera fulgida (holotype, Y.C. Dai 16107) A basidiomata B poroid surface detail C a dark line between tubes and context. Scale bars: 1 cm (A); 1 mm (B, C).

Hyphal structure

Hyphal system monomitic; generative hyphae with clamp connections, smooth, hyaline to yellowish, CB+, IKI–; tissues becoming dark in KOH.

Context

Hyphae thick-walled with a wide lumen, occasionally branched, loosely interwoven, 3–5 μm in diam.

Tubes

Hyphae thin- to slightly thick-walled, frequently branched, agglutinated and loosely interwoven, 2.5–3.5 μm in diam. Cystidia and cystidioles absent. Basidia clavate to more or less pyriform, with four sterigmata and a basal clamp connection, 10–12 × 4–5.5 μm; basidioles of similar shape to basidia, but smaller. Crystals present among hymenium.

Figure 5. 

Microscopic structures of Bjerkandera fulgida (holotype, Y.C. Dai 16107) a basidiospores b basidia and basidioles c hyphae from trama d hyphae from context.

Basidiospores

Ellipsoid to broadly ellipsoid, hyaline, thin-walled, smooth, CB–, IKI–, (3.8–)3.9–4.5 × (2.6–)2.8–3.3(–3.4) µm, L = 4.21 μm, W = 3.02 μm, Q = 1.37–1.43 (n = 90/3).

Additional specimens (paratypes) examined

China. Yunnan Province, Jinghong, Sanchahe Nature Reserve, 22°09'N, 100°51'E, fallen angiosperm trunk, 24. VI. 2011, Y.C. Dai 12284 (BJFC010566); Xishuangbanna Tropical Botanical Garden, fallen angiosperm trunk, 21°55'N, 101°15'E, 21.X.2013, Y.C. Dai 13597 (BJFC015059).

Remarks

Bjerkandera fulgida is characterised by the resupinate to effused-reflexed basidiomata, clay buff to pale brown and shiny pore surface, and ellipsoid to broadly ellipsoid basidiospores measuring 3.9–4.5 × 2.8–3.3 μm. Phylogenetically, Bjerkandera resupinata nests in a sister clade to B. fulgida (Fig. 1), also having morphological similarities, as the pore surface coloration and presence of branched hyphae in the tubes. However, B. resupinata differs in having resupinate basidiomata, larger pores (4–6 per mm), and basidiospores measuring 4.5–6 × 3.2–4.1 μm.

Bjerkandera minispora Y.C. Dai & Chao G. Wang, sp. nov.

MycoBank No: 838580
Figs 6, 7

Diagnosis

The tiny pores (6–9 per mm), and ellipsoid small basidiospores measuring 3.1–4.2 × 2–2.8 μm set this species apart from others in Bjerkandera.

Type

China. Hainan Province, Wuzhishan County, Wuzhishan Nature Reserve, 18°54'N, 109°42'E, fallen angiosperm trunk, 31. V. 2015, Y.C. Dai 15234 (holotype BJFC019345).

Etymology

Minispora (Lat.): referring to the species having small basidiospores.

Basidiomata

Annual, pileate, solitary or imbricate, soft corky, without odor or taste when fresh, becoming corky when dry. Pilei flabelliform, projecting up to 4 cm, 5 cm wide and 3 mm thick at base. Pileal surface pinkish-buff to buff, becoming dark when touched, velutinate to glabrous, azonate; margin a bit acute. Pore surface buff-yellow, ash-grey to pale brown when dry, touched or bruised parts becoming almost black; sterile margin distinct, up to 1.5 mm wide; pores tiny, round to angular, 6–9 per mm; pores mouth sometimes with white tomentum; dissepiments thin, entire to lacerate. Context cream to pinkish-buff, corky, up to 2 mm thick. Tubes concolorous with the pore surface, darker than context, corky, up to 1 mm long, with a distinct dark line between tubes and context.

Figure 6. 

Bjerkandera minispora (holotype, Y.C. Dai 15234) A basidiomata B poroid surface detail C a dark line between tubes and context. Scale bars: 1 cm (A); 1 mm (B, C).

Hyphal structure

Hyphal system monomitic; generative hyphae with clamp connections, smooth, hyaline to pale yellow, CB+, IKI–; tissues becoming dark in KOH.

Context

Generative hyphae thick-walled with a wide lumen, moderately branched, loosely interwoven, 3.5–6 μm in diam.

Figure 7. 

Microscopic structures of Bjerkandera minispora (holotype, Y.C. Dai 15234) a basidiospores b basidia and basidioles c hyphae from trama d hyphae from context.

Tubes

Generative hyphae thin-walled, frequently branched, agglutinated and loosely interwoven, 2.5–3.5 μm in diam. Cystidia and cystidioles absent. Basidia clavate, sometimes with an intermediate constriction, with four sterigmata and a basal clamp connection, 9.5–11.5 × 4–5 μm; basidioles of similar shape to basidia, but smaller.

Basidiospores

Oblong-ellipsoid to ellipsoid, hyaline, thin-walled, smooth, often with one or more guttules, CB–, IKI–, (3–)3.1–4.2(–4.8) × 2–2.8(–3) µm, L = 3.64 μm, W = 2.4 μm, Q = 1.49–1.54 (n = 60/2).

Additional specimen (paratype) examined

China. Hainan Province, Wuzhishan County, Wuzhishan Nature Reserve, 18°54'N, 109°42'E, fallen angiosperm trunk, 24. XI. 2007, B.K. Cui 5376 (BJFC003417).

Remarks

The buff-yellow pore surface, darkening when touched or bruised, the small pores (6–9 per mm) sometimes with white tomentum, and the ellipsoid small basidiospores (3.1–4.2 × 2–2.8 μm) set this species apart from others in Bjerkandera. Bjerkandera albocinerea resembles B. minispora by oblong-ellipsoid to ellipsoid basidiospores, but the former has sordid white fresh pileal surface, and dark brownish grey pore surface (Motato-Vásquez et al. 2020). Bjerkandera ecuadorensis is similar to B. minispora in having pinkish-buff to buff pileal surface and round to angular pores (6–9 per mm), but the former has grey to dark-brown pore surface and bigger basidiospores measuring 3.9–4.5 × 2.7–3 μm.

Bjerkandera resupinata Y.C. Dai & Chao G. Wang, sp. nov.

MycoBank No: 838581
Figs 8, 9

Diagnosis

Differs from other species of Bjerkandera by resupinate basidiomata.

Type

Thailand. Chiang Rai, Doi Mae Salong, rotten angiosperm trunk, 22. VII. 2016, Y.C. Dai 16642 (holotype BJFC022752).

Etymology

Resupinata (Lat.): referring to the species having resupinate basidiomata.

Basidiomata

Annual, resupinate, adnate, soft corky, without odor or taste when fresh, becoming corky when dry, up to 6 cm long, 2 cm wide, 0.5 mm thick at base. Pore surface pinkish buff to pale brownish when dry, becoming dark grey in bruised parts; sterile margin distinct, thinning out, somewhat incised, up to 3 mm wide; pores round to angular, 4–6 per mm; dissepiments thin, entire to lacerated. Subiculum pale cream, slightly fibrous to corky, up to 0.2 mm thick. Tubes concolorous with the pore surface, darker than the subiculum, corky, up to 0.3 mm long, with a distinct dark line between tubes and subiculum.

Figure 8. 

Bjerkandera resupinata (holotype, Y.C. Dai 16642) A basidiomata B poroid surface detail C a dark line between tubes and subiculum. Scale bars: 1 cm (A); 1 mm (B, C).

Hyphal structure

Hyphal system monomitic; generative hyphae with clamp connections, smooth, hyaline to yellowish, CB+, IKI–; tissues becoming dark in KOH.

Subiculum

Generative hyphae thick-walled with a wide lumen, rarely branched, loosely interwoven, 4–5 μm in diam.

Tubes

Generative hyphae thin- to slightly thick-walled, frequently branched, loosely interwoven, 2.7–3.8 μm in diam. Cystidia and cystidioles absent. Basidia clavate, with four sterigmata and a basal clamp connection, 14–16 × 5–6.5 μm; basidioles in shape similar to basidia, but smaller.

Figure 9. 

Microscopic structures of Bjerkandera resupinata (holotype, Y.C. Dai 16642) a basidiospores b basidia and basidioles c hyphae from trama d hyphae from subiculum.

Basidiospores

Ellipsoid to broadly ellipsoid, hyaline, thin-walled, smooth, CB–, IKI–, 4.5–6(–6.2) × 3.2–4.1(–4.2) µm, L = 5.23 μm, W = 3.71 μm, Q = 1.40–1.42 (n = 60/2).

Additional specimen (paratype) examined

China. Yunnan Province, Tengchong County, Gaoligong Mts., fallen angiosperm branch, 24. X. 2009, B.K. Cui 8017 (BJFC006506).

Remarks

Bjerkandera resupinata is characterised by resupinate basidiomata, pinkish buff to pale brownish pore surface, clavate basidia, and ellipsoid to broadly ellipsoid basidiospores measuring 4.5–6 × 3.2–4.1 µm. Ceriporiopsis umbrinescens (Murrill) Ryvarden and Bjerkandera resupinata have resupinate basidiomata, pale buff to brownish pore surface, similar sterile margin, a monomitic hyphal structure, and almost the same size of basidiospores, but C. umbrinescens has bigger pores (2–4 per mm), unchanged pore surface when touched, and a dark line absent between tubes and subiculum (Murrill 1920; Domański 1963; Núñez and Ryvarden 2001; Zhao et al. 2015).

Table 2.

Morphological comparison of the currently accepted species in Bjerkandera.

Basidiomata type Pilei colour Pore shape and number of pores Poroid surface Basidiospores size (μm) Basidiospores shape Reference
B. adusta Pileate, effused-reflexed to resupinate Cream to buff, then greyish to greyish-blue Round to angular, 6–7/mm Grey to black 4.5–6 × 2.5–3.5 Short-cylindrical to subellipsoid Ryvarden and Melo 2017
B. albocinerea Pileate to effused-reflexed Sordid white to pale cream Round, 8–11/mm Dark brown grey to almost black when bruised 3.5–4.5 × 2–2.6 Oblong-ellipsoid to ellipsoid Motato-Vásquez et al. 2020
B. atroalba Pileate to effused-reflexed White to cream, then grey Round or more commonly angular, 2–5/mm White to cream, then becoming dark 4–5 × 3–4 Narrowly ellipsoid to broadly ellipsoid Westphalen et al. 2015
B. centroamericana Pileate to effused-reflexed White to cream, then brownish Angular, 7–11/mm Sordid white, then brown to black in bruised parts 4–5 × 3–4.5 Broadly ellipsoid to subglobose Westphalen et al. 2015
B. ecuadorensis Pileate Pinkish-buff to buff Round to angular, 7–9 /mm Grey to dark-brown, then almost black in bruised parts 3.9–4.5 × 2.7–3 Ellipsoid Present study
B. fulgida Effused-reflexed Pinkish buff to clay-buff Round or sometimes angular, 6–8/mm Clay-buff to pale brown, then dark brown in bruised parts 3.9–4.5 × 2.8–3.3 Ellipsoid to broadly ellipsoid Present study
B. fumosa Pileate to effused-reflexed Buff to woody coloured Round to angular 2–5/mm Buff to isabelline 5.5–7 × 2.5–3.5 Short cylindrical Ryvarden and Melo 2017
B. mikrofumosa Effused-reflexed Pale golden-brown Angular, 7–9/mm Pale to smoky brown, then dark grey in bruised parts 3.5–4.8 × 2.3–3 Ellipsoid Motato-Vásquez et al. 2020
B. minispora Pileate Pinkish-buff to buff Round to angular, 6–9/mm Buff-yellow, ash-grey to pale brown, then almost black in bruised parts 3.1–4.2 × 2–2.8 Oblong-ellipsoid to ellipsoid Present study
B. resupinata Resupinate Round to angular, 4–6/mm Pinkish buff to pale brownish, then dark grey in bruised parts 4.5–6 × 3.2–4.1 Ellipsoid to broadly ellipsoid Present study

Additional specimens examined

Bjerkandera adusta : China. Heilongjiang Province, Heihe, Shengshan Nature Reserve, Populus, 26. VIII. 2014, Y. C. Dai 14516 (BJFC017794); Yunnan Province, Yongde County, Daxueshan Nature Reserve, rotten Angiosperm stump, 27. VIII. 2015, Y. C. Dai 15665 (BJFC019769); Gansu Province, Tianshui, Fangmatan Forest Park, fallen branch of Populus, 08. VIII. 2015, Y. C. Dai 15495 (BJFC019600). France. Lyons, Abies, 24. XI. 2012, Y. C. Dai 13201 (BJFC014065). Finland, Helsinki, Tamisto Nature Reserve, Betula, 4. XI. 2011, Y. C. Dai 12640 (BJFC012222). B. albocinerea: USA. CT, CAES Valley Lab, Dead log, 13. XII. 2015, Y. C. Dai 16411 (BJFC020499). B. fumosa: China. Chongqing, Jinfoshan Forest Park, dead angiosperm tree, 1. XI. 2019, Y. C. Dai 21100 (BJFC032759); Beijing, Chinese Academy of Sciences, living tree of Diospyros, Y. C. Dai 21087 (BJFC032746); Sichuan Province, Xiaojin County, Jiajin Mts., Hippophae, 17. X. 2012, B. K. Cui 10747 (BJFC013669). Finland, Helsinki, Tamisto Nature Reserve, Populus, 6. XI. 2011, Y. C. Dai 12474B (BJFC012257). B. mikrofumosa: Costa Rica. Monteverde, J. Vlasák Jr. JV 1707/10J-1; JV 1707/10J-2. B. centroamericana: Costa Rica, Carara Nature Reserve, J. Vlasák JV 1704/97. B. atroalba: Brazil. Recife, Charles Darwin Ecological Reserve, on angiosperm stump, Y. C. Dai 17457 (BJFC024988). Ceriporiopsis carnegieae: USA, Virgin Islands, St. John, on hard wood, J. Vlasák Jr. JV 0409/27-J. Ceriporiopsis sp.: Costa Rica. Arenal Mts., J. Vlasák Jr. JV 1512/13-J.

Key to the species of Bjerkandera

1 Basidiomata resupinate B. resupinata
Basidiomata effused-reflexed to pileate 2
2 Pores < 5 per mm 3
Pores > 5 per mm 4
3 Pileal surface white to cream; basidiospores broadly ellipsoid B. atroalba
Pileal surface buff to woody-coloured; basidiospores short cylindrical B. fumosa
4 Pileal surface white to cream when fresh 5
Pileal surface buff to grey when fresh 6
5 Basidiospores subglobose to broadly ellipsoid B. centroamericana
Basidiospores oblong-ellipsoid to ellipsoid B. albocinerea
6 Crystals present among hymenium 7
Crystals absent among hymenium 8
7 Pileal margin dark brown when dry B. mikrofumosa
Pileal margin buff when dry B. fulgida
8 Basidiospores > 4.5 μm in length B. adusta
Basidiospores < 4.5 μm in length 9
9 Basidiospores 3.1–4.2 × 2–2.8 μm, Q = 1.49–1.53 B. minispora
Basidiospores 3.9–4.5 × 2.7–3 μm, Q = 1.43 B. ecuadorensis

Discussion

Our phylogeny recovered Bjerkandera as a monophyletic genus, with ten species including the four new species – Bjerkandera ecuadorensis, B. fulgida, B. minispora, and B. resupinata – nested in the Bjerkandera clade (Fig. 1).

Bjerkandera ecuadorensis, B. minispora, B. adusta, B. albocinerea, and B. fumosa are phylogenetically related (Fig. 1). B. adusta, B. albocinerea, and B. fumosa form a group which is consistent with previous studies (Westphalen et al. 2015; Motato-Vásquez et al. 2020). The specimen we studied Dai 16411 from CT, USA and Bjerkandera albocinerea share cream to buff-yellow pileal surface when dry, dark brownish grey to black pore surface, round pores (8–11 per mm), and oblong-ellipsoid to ellipsoid basidiospores (3.5–4.5 × 2–2.5 μm). Also, there are two base pairs differences between them, which amounts to < 1% nucleotide differences in the ITS regions. So both specimens represent the same species. The type of Bjerkandera albocinerea and other specimens were collected from Brazil, but the specimen Dai 16411 from CT, USA, B. albocinerea has a wide distribution in America. Morphologically, Bjerkandera albocinerea is different from other four species by its white fresh pileal surface (Motato-Vásquez et al. 2020), and B. minispora can be distinguished from B. ecuadorensis, B. adusta and B. fumosa by smaller basidiospores (3.1–4.2 × 2–2.8 μm in B. minispora, 3.9–4.5 × 2.7–3 μm in B. ecuadorensis, and 4.5–6 × 2.5–3.5 μm in B. adusta, 5.5–7 × 2.5–3.5 μm in B. fumosa, Ryvarden and Melo 2017). Bjerkandera fumosa has the thicker context usually more than 6 mm, while the other two less than 6 mm (Ryvarden and Melo 2017). Bjerkandera adusta has short-cylindric to subellipsoid and bigger basidiospores (4.5–6 × 2.5–3.5 μm, Ryvarden and Melo 2017), which can differ from B. ecuadorensis. Also, there are 21 base pairs differences between Bjerkandera ecuadorensis and B. minispora, which amounts to > 2% nucleotide differences in the ITS regions.

Bjerkandera fulgida grouped with B. resupinata in a joint subclade, and these two species are closely related to B. atroalba (Rick) Westph. & Tomšovský, B. centroamericana Kout, Westph. & Tomšovský, and B. mikrofumosa Ryvarden with strong support (99/98/1.00). Bjerkandera resupinata has resupinate basidiomata, big pores and basidiospores, which can be distinguished from B. fulgida indeed. Also, there are eight base pairs differences between them, which amounts to 2% nucleotide differences in the ITS regions. Bjerkandera atroalba, B. centroamericana and B. mikrofumosa have a neotropical distribution (Westphalen et al. 2015; Motato-Vásquez et al. 2020), while B. fulgida and B. resupinata from tropical China are proved to nest in the group according to our phylogenetic study. Morphologically, Bjerkandera resupinata is a resupinate species, while basidiomata are effused-reflexed to pileate in B. fulgida, B. atroalba, B. centroamericana, and B. mikrofumosa (Westphalen et al. 2015; Motato-Vásquez et al. 2020). Bjerkandera atroalba and B. centroamericana differ from B. fulgida by their white pilei when fresh, sordid white to cream pore surface, and the presence of cystidioles (Westphalen et al. 2015). B. mikrofumosa differs from B. fulgida by its pale golden-brown pileal surface and pale to smoky brown pore surface (Motato-Vásquez et al. 2020). In addition, we found Ceriporiopsis umbrinescens (Murrill) Ryvarden and B. resupinata have resupinate basidiomata, pale buff to brownish pore surface, similar sterile margin, a monomitic hyphal structure, and almost the same size of basidiospores, but C. umbrinescens has bigger pores (2–4 per mm) and unchanged pore surface when touched (Murrill 1920; Domański 1963; Núñez and Ryvarden 2001; Zhao et al. 2015).

In our phylogenetic analysis, Ceriporiopsis carnegieae (D.V. Baxter) Gilb. & Ryvarden is phylogenetically close to the genus Bjerkandera. Ceriporiopsis Domański is a polyphyletic genus, which is nested in the families Irpicaceae, Meruliaceae (the type species C. gilvescens (Bres.) Domański belongs to Meruliaceae), and Phanerochaetaceae (Justo et al. 2017). Meanwhile, Ceriporiopsis carnegieae resembles Bjerkandera by having a monomitic hyphal system, generative hyphae with abundant clamps, and oblong to short-cylindric basidiospores (Baxter 1941; Gilbertson and Ryvarden 1985). However, the former has basidiomata with sharp and pungent odor when fresh, unchanged pore surface when touched or bruised, and seem to lack any dark line between tubes and subiculum (Gilbertson and Ryvarden 1985). One specimen – JV1512–13J – from Costa Rica forms a sister group to the three sequences annotated as Ceriporiopsis carnegieae, and we treat this specimen as Ceriporiopsis sp. There is ongoing controversy regarding for the generic affiliation of C. carnegieae (Nobles 1965; Justo et al. 2017; Motato-Vásquez et al. 2020), because the black line is absent from Ceriporiopsis carnegieae. For the time being, we are reluctant to combine them in Bjerkandera although the two taxa are phylogenetically related. To solve this problem more specimens should be examined and analysed phylogenetically.

Beside the ten species of Bjerkandera in our phylogeny (Fig. 1), another three taxa – Bjerkandera terebrans (Berk. & M.A. Curtis) Murrill, B. subsimulans (Berk. et M. A. Curtis) Murrill and B. amorpha (Fr.) P. Karst. – were included in the genus. However, Bjerkandera terebrans was mentioned probably as a form of B. fumosa or variant of Osteina obducta (Berk.) Donk Because of its basidiomata with a stipe-like base (Murrill 1907; Zmitrovich et al. 2016). B. subsimulans has lobed and broadly sterile margin with a zone of appressed hairs, and angular irregular pores (1–3 per mm), which are in accord with the description of Abortiporus biennis (Bull.) Singer (Murrill 1907; Zmitrovich et al. 2016). B. amorpha has dimitic hyphal system and allantoid basidiospores that differ from Bjerkandera, so it is now Skeletocutis amorpha (Fr.) Kotl. & Pouzar (Kotlába and Pouzar 1958).

Tyromyces vivii Homble ex Ryvarden was described from Norway (Ryvarden et al. 2003), and later it was treated as a synonym of B. fumosa (Ryvarden and Melo 2017). The type material of T. vivii was analyzed, and it nested in B. fumosa (Fig. 1). We confirm this conclusion by molecular evidence.

Previously, the well-known Bjerkandera adusta and B. fumosa have been reported from the northern hemisphere and South America. However, the diversity of Bjerkandera was underestimated, B. centroamericana, B. mikrofumosa and B. albocinerea were recently described in the neotropics (Westphalen et al. 2015; Ryvarden 2016), and new species in our study have a distribution in the neotropics and tropical Asia. So, the genus has a wide distribution from boreal to tropical areas.

Acknowledgements

The research was supported by the National Natural Science Foundation of China (Project No. 31870007) and by the institutional support of the Academy Sciences of the Czech Republic RVO: 60077344. Special thanks are due to Josef Vlasák Jr. (USA) for forwarding specimens for our study.

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