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Research Article
Three new species of Favolaschia (Mycenaceae, Agaricales) from South China
expand article infoQiu-Yue Zhang‡, Hong-Gao Liu§, Wan-Ying Li‡, Xin Zhang‡, Yu-Cheng Dai‡, Fang Wu‡, Lu-Sen Bian‡|
‡ Beijing Forestry University, Beijing, China
§ Zhaotong University, Yunnan, China
| Chinese Academy of Forestry, Beijing, China

Corresponding author: Yu-Cheng Dai ( yuchengdai@bjfu.edu.cn )

Corresponding author: Fang Wu ( fangwubjfu2014@bjfu.edu.cn )

Corresponding author: Lu-Sen Bian ( apsenvictory@126.com )

Academic editor: Samantha C. Karunarathna
© 2024 Qiu-Yue Zhang, Hong-Gao Liu, Wan-Ying Li, Xin Zhang, Yu-Cheng Dai, Fang Wu, Lu-Sen Bian.
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: Zhang Q-Y, Liu H-G, Li W-Y, Zhang X, Dai Y-C, Wu F, Bian L-S (2024) Three new species of Favolaschia (Mycenaceae, Agaricales) from South China. In: Wijayawardene N, Karunarathna S, Fan X-L, Li Q-R (Eds) Taxonomy and secondary metabolites of wood-associated fungi. MycoKeys 104: 71-89. https://doi.org/10.3897/mycokeys.104.117310
Open Access

Abstract

The genus Favolaschia within the family Mycenaceae is characterised by the gelatinous basidiomata with poroid hymenophore and most species inhabit monocotyledonous plants. In this study, many samples covering a wide geographic range in China were examined morphologically and phylogenetically using concatenated ITS1-5.8S-ITS2-nLSU sequence data. Three new species clustering in Favolaschia sect. Anechinus, namely Favolaschia imbricata, F. miscanthi and F. sinarundinariae, are described. Favolaschia imbricata is characterised by imbricate basidiomata with pale grey to greyish colour when fresh and broadly ellipsoid basidiospores measuring 7–9 × 5–6.8 µm; F. miscanthi is characterised by satin white basidiomata when fresh, broadly ellipsoid basidiospores measuring 7.5–10 × 5.5–7 µm and inhabit rotten Miscanthus; F. sinarundinariae is characterised by greyish-white basidiomata when fresh, dark grey near the base upon drying, broadly ellipsoid to subglobose basidiospores measuring 7–9 × 5–7 µm and inhabit dead Sinarundinaria. The differences amongst the new species and their morphologically similar and phylogenetically related species are discussed. In addition, an updated key to 19 species of Favolaschia found in China is provided.

Key words

New taxa, phylogeny, taxonomy, wood-decaying fungi

Introduction

The genus Favolaschia (Pat.) Pat., typified by F. gaillardia (Pat.) Pat. (Patouillard 1887), belongs to the Mycenaceae Overeem (Agaricales) (Moncalvo et al. 2002; Bodensteiner et al. 2004). It is characterised by gelatinous basidiomata with poroid hymenophore, a monomitic hyphal system, a gelatinous hyphal structure, the presence of gloeocystidia and acanthocystida which are terminal, mostly swollen tips of hyphae covered by outgrowths (Clémençon 1977) and amyloid basidiospores (Singer 1945, 1974; Gillen et al. 2012; Capelari et al. 2013; Magnago et al. 2013; Zhang and Dai 2021; Zhang et al. 2023). Favolaschia species have a worldwide distribution and prefer humid climates with the highest species diversity in subtropical to tropical zones (Singer 1974; Gillen et al. 2012; Magnago et al. 2013). The basidiomata are found mostly on decaying parts of monocotyledonous plants, as well as ferns and cause white rot (Singer 1974; Gillen et al. 2012).

A total of 119 records of Favolaschia are listed in Index Fungorum (http://www.indexfungorum.org) and around 60 species are accepted (Johnston et al. 2006; Gillen et al. 2012; Magnago et al. 2013). The name “Favolaschia” was first introduced by Patouillard (1887) as a section of Laschia Fr and treated at a generic level later (Patouillard and Lagerheim 1892). The taxonomic history of the genus has been systematically compiled by Singer (1974) and Johnston et al. (2006). Singer (1974) divided Favolaschia into two sections, based on morphological studies, namely section Favolaschia Singer and section Anechinus Singer. Amongst them, taxa of F. sect. Favolaschia is characterised by having mostly orange, yellow, red, lateritious or pink pileus and the presence of acanthocytes in pileipellis. Inversely, taxa of F. sect. Anechinus are characterised by lacking acanthocystidia or replaced by diverticulate hyphae. Later, Johnston et al. (2006), Gillen et al. (2012) and Magnago et al. (2013) confirmed the two sections by phylogenetic analyses.

Recently, the systematic study of Favolaschia in China has gradually increased. Tolgor et al. (2021) investigated the phylogenetic relationships of Favolaschia and its related genera in Mycenaceae and reported four species distributed in China. Zhang and Dai (2021) investigated phylogenetic analysis of the genus Favolaschia, based on a large sample and revealed four new species in the F. calocera complex. Until Zhang et al. (2023) re-summarised the species diversity, phylogenetic relationships, divergence time and potential geographic distribution using a large number of samples covering a wide geographic range in China. To date, 16 species of Favolaschia have been reported from China, based on phylogenetic analyses, namely, F. brevibasidiata Q.Y. Zhang and Y.C. Dai, F. brevistipitata Q.Y. Zhang and Y.C. Dai, F. longistipitata Q.Y. Zhang and Y.C. Dai, F. minutissima Q.Y. Zhang and Y.C. Dai, F. bannaensis Q.Y. Zhang & Y.C. Dai, F. crassipora Q.Y. Zhang & Y.C. Dai, F. flabelliformis Q.Y. Zhang & Y.C. Dai, F. manipularis (Berk.) Teng, F. peziziformis (Berk. and M.A. Curtis) Kuntze, F. pustulosa (Jungh.) Kuntze, F. rigida Q.Y. Zhang & Y.C. Dai, F. semicircularis Q.Y. Zhang & Y.C. Dai, F. subpustulosa Q.Y. Zhang & Y.C. Dai, F. tenuissima Q.Y. Zhang & Y.C. Dai, F. tephroleuca Q.Y. Zhang & Y.C. Dai and F. tonkinensis (Pat.) Kuntze; the first four species belong to the F. calocera complex of the F. sect. Favolaschia; while the others belong to the F. sect. Anechinus (Zhang and Dai 2021; Guo et al. 2022; Ma et al. 2022; Wu et al. 2022a; Dong et al. 2023; Zhang et al. 2023).

During a study on wood-rotting fungi from south China, several samples belonging to Favolaschia were collected and three unknown species were morphologically distinguished. To confirm the affinity of the taxa, phylogenetic analysis was performed, based on a combined sequence dataset of ITS1-5.8S-ITS2-nLSU.

Materials and methods

Morphological studies

The specimens were collected from Guangxi Autonomous Region, Guangdong and Sichuan Provinces in south China and the specific locations were marked in Fig. 1. They were deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC), Beijing, China. Macro-morphological descriptions were based on field notes and dried specimens. Microscopic features were examined and described in 5% KOH (potassium hydroxide) and 2% phloxine B (C20H2Br4Cl4Na2O5) with a magnification of up to 1,000× using a Nikon Eclipse 80i microscope and phase contrast illumination. Colour terms followed Kornerup and Wanscher (1978) and Petersen (1996). A Nikon Digital Sight DS-L3 camera was used to photograph microscopic structures. Other reagents, including Cotton Blue and Melzer’s reagents were used to observe the micromorphology structures following Zhang et al. (2023). To show the variation in spore sizes, 5% of measurements were excluded from each end of the range and shown in parentheses. Thirty basidiospores from each specimen were measured. The following abbreviations are used: IKI = Melzer’s reagent, IKI+ = amyloid; CB = Cotton Blue, CB– = acyanophilous in Cotton Blue; L = arithmetic average of basidiospores length, W = arithmetic average of basidiospores width, Q = L/W ratios, (n = x/y) = the number of spores (x) measured from a given number of specimens (y).

Figure 1. 

The geographical locations of Favolaschia species distributed in China (the base map of China was obtained from the Map Technology Review Center, Department of Natural Resources, http://bzdt.ch.mnr.gov.cn/).

DNA extraction and sequencing

A cetyl trimethylammonium bromide (CTAB) rapid plant genome extraction kit (Aidlab Biotechnologies, Co., Ltd., Beijing, China) was used to extract DNA (Wu et al. 2020, 2022b). The ITS1-5.8S-ITS2 region was amplified with the primer pair ITS5/ITS4 (White et al. 1990) using the following protocol: initial denaturation at 95 °C for 4 min, followed by 34 cycles at 94 °C for 40 s, 54 °C for 45 s and 72 °C for 1 min and final extension at 72 °C for 10 min. The nLSU region was amplified with the primer pair LR0R/LR7 (White et al. 1990) using the following protocol: initial denaturation at 94 °C for 1 min, followed by 34 cycles at 94 °C for 30 s, 50 °C for 1 min and 72 °C for 1.5 min and final extension at 72 °C for 10 min. The PCR products were purified and sequenced by the Beijing Genomics Institute (BGI), China with the same primers. The newly-generated sequences in this study have been deposited in GenBank and are listed in Table 1.

Table 1.
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Names, specimen numbers, locality and corresponding GenBank accession numbers of the taxa used in this study.

Species Specimen no. Locality ITS no. nLSU no.
Favolaschia andina KG0025 Panama HM246678 HM246679
F. aurantiaca FK2047 Brazil JX987670 –
F. aurantiaca KG0013 Panama – HM246676
F. auriscalpium Isolate 5 – KY649461 –
F. auriscalpium TH1018 Guyana DQ026241 –
F. austrocyatheae PDD75609 New Zealand NR132809 –
F. austrocyatheae PDD75609 New Zealand DQ026257 –
F. bannaensis Dai 22587 Yunnan, China ON870497 ON870473
F. bannaensis Dai 22589 Yunnan, China ON870498 ON870474
F. bannaensis Dai 22590 Yunnan, China ON870499 ON870475
F. brevibasidiata Cui 6573 Hainan, China MZ661794 –
F. brevibasidiata JM98186 Yunnan, China DQ026239 –
F. brevistipitata Dai 19780 Yunnan, China MZ661772 MZ661742
F. brevistipitata Dai 19855 Yunnan, China MZ661773 MZ661743
F. brevistipitata Dai 19856 Yunnan, China MZ661774 MZ661744
F. calocera PC99060 Madagascar DQ26252 –
F. calocera PC99497 Madagascar DQ026253 –
F. cinnabarina Isolate 4421 Brazil JX987669 –
F. cinnabarina RVPR82 – – AF261416
F. claudopus Dai 18656 Australia MZ661775 MZ661735
F. claudopus Dai 18663 Australia MZ661776 MZ661734
F. claudopus SR346 Kenya DQ026237 –
F. claudopus PDD74554 New Zealand DQ026251 –
F. claudopus PDD75323 New Zealand DQ026248 –
F. claudopus PDD75686 New Zealand DQ026249 –
F. claudopus DUKE2952 New Zealand DQ026238 –
F. crassipora Dai 19769 Yunnan, China ON870500 ON870476
F. crassipora Dai 19871 Yunnan, China ON870501 ON870477
F. cyatheae PDD75316 New Zealand NR132808 –
F. cyatheae PDD75316 New Zealand DQ026256 –
F. dealbata KG0015 Panama – HM246677
F. flabelliformis Dai 20010 Yunnan, China ON870502 ON870478
F. flabelliformis Dai 20016 Yunnan, China ON870503 –
F. flabelliformis Dai 24354 Yunnan, China OR271272 OR260441
F. heliconiae KG0026 Panama – HM246680
F. imbricata Dai 24702 Guangdong, China OR334876 –
F. imbricata Dai 24703 Guangdong, China OR334877 OR334881
F. imbricata Dai 24704 Guangdong, China OR334878 OR334882
F. longistipitata Dai 13221 Yunnan, China MZ661777 –
F. longistipitata Dai 13226 Yunnan, China MZ661778 –
F. longistipitata Cui 11128 Yunnan, China MZ661779 –
F. longistipitata Dai 17597 Yunnan, China MZ661780 –
F. longistipitata Dai 17598 Yunnan, China MZ661781 –
F. longistipitata Dai 17601 Yunnan, China MZ661782 –
F. longistipitata Dai 19799 Yunnan, China MZ661784 MZ661739
F. longistipitata Dai 19893 Yunnan, China MZ661785 MZ661740
F. longistipitata Dai 20019 Yunnan, China MZ661786 MZ661741
F. longistipitata Dai 20328 Yunnan, China MZ661787 –
F. longistipitata Dai 20341 Yunnan, China MZ661788 –
F. longistipitata Dai 20355 Yunnan, China MZ661789 –
F. luteoaurantiaca Isolate 4475 – JX987667 –
F. luteoaurantiaca SP445750 Brazil NR132874 –
F. macropora KG0027 Panama NR132845 HM246682
F. manipularis Dai 20612 Yunnan, China MZ801776 MZ914395
F. manipularis Dai 20653 Yunnan, China MZ801777 –
F. minutissima JM98372 Thailand DQ026240 –
F. minutissima Dai 10753 Hainan, China MZ661790 –
F. minutissima Dai 20085 Hainan, China MZ661791 MZ661736
F. minutissima Dai 20086 Hainan, China MZ661792 MZ661737
F. minutissima Dai 20088 Hainan, China MZ661793 MZ661738
F. miscanthi Dai 24652 Guangxi, China OR334879 OR334883
F. miscanthi Dai 24653 Guangxi, China OR334880 OR334884
F. peziziformis ICMP1575 Japan DQ026255 –
F. peziziformis PDD67440 New Zealand – AY572008
F. pustulosa PDD75686 New Zealand DQ026254 –
F. pustulosa Dai 19758 Yunnan, China MT292325 MT293226
F. rigida Dai 18566A Guangxi, China ON870504 ON870479
F. rigida Dai 20764 Yunnan, China ON870505 ON870480
F. rigida Dai 22614 Fujian, China ON870506 –
F. semicircularis Dai 19725 Guangdong, China ON870507 ON870481
F. semicircularis Dai 19923 Yunnan, China ON870508 ON870482
F. semicircularis Dai 19936 Yunnan, China ON870509 ON870483
F. semicircularis Dai 19939 Yunnan, China ON870510 –
F. semicircularis Dai 19980 Yunnan, China ON870511 ON870484
F. semicircularis Dai 19981 Yunnan, China ON870512 ON870485
F. semicircularis Dai 22290 Zhejiang, China ON870513 ON870486
F. semicircularis Dai 22298 Zhejiang, China ON870515 ON870488
F. semicircularis Dai 22302 Zhejiang, China ON870516 –
F. semicircularis Dai 23702 Hunan, China OR271273 OR260442
F. semicircularis Dai 24689 Guangxi, China OR271274 OR260443
F. semicircularis Dai 24748 Jiangxi, China OR271275 OR260444
F. semicircularis Dai 24687 Guangxi, China OR271276 OR260445
F. semicircularis Dai 22383 Fujian, China ON870517 –
F. sinarundinariae Dai 26115 Sichuan, China OR575908 OR575906
F. sinarundinariae Dai 26116 Sichuan, China OR575909 OR575907
F. sinarundinariae Dai 26123 Sichuan, China OR855969 OR855975
F. sinarundinariae Dai 26129 Sichuan, China OR855970 –
F. sinarundinariae Dai 26130 Sichuan, China OR855971 –
F. sinarundinariae Dai 26131 Sichuan, China OR855972 –
F. sinarundinariae Dai 26134 Sichuan, China OR855973 OR855976
F. sinarundinariae Dai 26140 Sichuan, China OR855974 OR855975
F. sp. 1 DUKE2708 Australia DQ026234 –
F. sp. 1 DUKE2876 Australia DQ026235 –
F. sp. 1 DUKE3195 Papua New Guinea DQ026236 –
F. sp. 2 Isolate 4550 Panama JX987668 –
F. sprucei TH6418 Guyana DQ026246 –
F. subpustulosa Dai 20719 Yunnan, China ON870518 ON870489
F. tenuissima Dai 22072 Hainan, China ON870520 ON870491
F. tenuissima Dai 22071 Hainan, China ON870519 ON870490
F. tephroleuca Dai 22282 Yunnan, China ON870521 ON870492
F. tephroleuca Dai 22288 Chongqing, China ON870522 ON870493
F. tonkinensis Dai 21955 Hainan, China ON870523 ON870494
F. tonkinensis Dai 21956 Hainan, China ON870524 ON870495
F. tonkinensis Dai 21964 Hainan, China ON870525 –
F. tonkinensis Dai 21965 Hainan, China ON870526 ON870496
F. tonkinensis Dai 21966 Hainan, China ON870527 –
F. tonkinensis Dai 19704 Guangdong, China OR271277 OR260446
F. tonkinensis BCC 18686 – MN093316 MN093317
F. tonkinensis JM98229 Yunnan, China DQ026247 –
F. varariotecta DUKE3893 Puerto Rico DQ026243 –
F. varariotecta DUKE4038 Puerto Rico DQ026244 –
F. xtbgensis HKAS 121667 Yunnan, China OL413048 OL413044
F. xtbgensis HKAS 121975 Yunnan, China OL413036 OL413035
Mycena seminau ACL136 Malaysia KF537250 KJ206952
Mycena seminau ACL308 Malaysia KF537252 KJ206964

Notes: New sequences are in bold; “–” represents missing data.

Phylogenetic analyses

The dataset of concatenated ITS1-5.8S-ITS2-nLSU sequences of the Favolaschia was analysed. Sequences of Mycena seminau A.L.C. Chew & Desjardin were used as outgroups (Chew et al. 2014). The newly-generated sequences in this study and additional sequences retrieved from Zhang et al. (2023) were partitioned to ITS1, 5.8S, ITS2, nrLSU and then aligned separately using MAFFT v.74 (http://mafft.cbrc.jp/alignment/server/, Katoh et al. 2017) with the G-INS-I iterative refinement algorithm and optimised manually in BioEdit v.7.0.5.3 (Hall 1999). The separate alignments were then concatenated using PhyloSuite v.1.2.2 (Zhang et al. 2020).

Maximum Likelihood (ML) analyses and Bayesian Inference (BI) were carried out by using RAxML v.8.2.10 (Stamatakis 2014) and MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003), respectively. In ML analysis, statistical support values were obtained by using rapid bootstrapping with 1000 replicates, with default settings for other parameters. For BI, the best-fit partitioning scheme and substitution model were determined by using ModelFinder (Kalyaanamoorthy et al. 2017) via the “greedy” algorithm, branch lengths estimated as “linked” and AICc. Four Markov chain Monte Carlo chains (one cold) were constructed for 5,000,000 generations, with sampling every 1000 generations. Convergence was assessed as the standard deviation of split frequencies < 0.01. The first quarter of the trees, which represented the burn-in phase of the analyses, were discarded and the remaining trees were used to calculate posterior probabilities (BPP) in the majority rule consensus tree.

Phylogenetic trees were visualised by using FigTree version 1.4.4 (Rambaut 2018). Branches that received bootstrap supports for ML (≥ 75%) and BPP (≥ 0.95) were considered as significantly supported. The best topologies from ML analyses are shown in this study and the final alignments and the retrieved topologies were deposited in TreeBASE (http://treebase.org/treebase-web/home.html), under accession ID: 30973.

Results

Phylogenetic analyses

In this study, the combined ITS1-5.8S-ITS2-nLSU dataset included sequences from 113 specimens, representing 34 species of Favolaschia and one species of Mycena (Pers.) Roussel as the outgroup (Table 1, Fig. 2). ModelFinder suggested models were HKY+F+G4 for ITS1+ITS2, K2P+G4 for 5.8s and K2P+G4 for nLSU, for the Bayesian analysis. The BI analysis resulted in a concordant topology with an average standard deviation of split frequencies of 0.008584. The ML and BI analyses resulted in nearly identical topologies and only the ML tree is presented with the bootstrap supports for ML and BPP not less than 50% and 0.90, respectively.

Figure 2. 

Maximum Likelihood (ML) tree illustrating the phylogeny of Favolaschia, based on a combined ITS1-5.8S-ITS2-nLSU-mtSSU-nuSSU dataset. Branches are labelled with parsimony bootstrap values (ML) higher than 50% and Bayesian Posterior Probabilities (BPPs) more than 0.90. (Group A: Favolaschia sect. Favolaschia; Group B: Favolaschia sect. Anechinus)

The phylogeny is similar to those of Johnston et al. (2006), Magnago et al. (2013) and Zhang et al. (2023). All Favolaschia samples are clustered within two groups (A and B), which correspond to the sections proposed by Singer (1974): Favolaschia sect. Favolaschia (group A) and F. sect. Anechinus (group B), except F. manipularis. In this study, three new lineages with high support (100/1.00, 100/1.00 and 71/1.00, respectively) nest in F. sect. Anechinus (group B). Amongst them, five specimens from Guangdong and Guangxi formed two lineages and clustered into a clade with strong support (100/1.00), namely F. imbricata and F. miscanthi. Eight specimens from Sichuan formed a support lineage, namely F. sinarundinariae, sister to Favolaschia tephroleuca. Furthermore, similarity searches with the ITS barcoding from the three new species were performed using BLAST (http://www.ncbi.nlm.nih.gov/BLAST/) from NCBI and showed less than 95% identity, respectively, in the genus Favolaschia, which demonstrated the obvious nucleotide differences of the three species with other species in the genus.

Taxonomy

Favolaschia imbricata Q.Y. Zhang, L.S. Bian, F. Wu & Y.C. Dai, sp. nov.

MycoBank No: 851807
Figs 3A, 4

Type

China, Guangdong Province, Guangzhou, Baiyun District, Maofengshan Forest Park, dead bamboo, 15 Apr 2023, Dai 24702 (BJFC042256, holotype).

Etymology

“imbricata” (Lat.): refers to the species having imbricate basidio­mata.

Macrostructures

Basidiomata annual, gregarious, gelatinous when fresh and dry. Pilei 2–5 × 1.5–3 mm, conchoid, semicircular to subcircular; pileal surface pale grey to greyish-white (B1) when fresh, dark grey or mouse-grey (1D1–1F1) upon drying, convex with a reticulate pattern matching the pores below, faintly pruinose when dry; margin incurved, entire; context thin. Hymenophore paler than pileal surface, greyish-white (B1) when fresh and ash-grey (1B1–1C1) when dry, poroid, about 22–55 pores per basidiomata; mature pores 0.2–0.6 mm diam., polygonal, larger near the base and smaller near the edge, the marginal pores often incomplete; tubes up to 0.3 mm long. Stipe absent.

Figure 3. 

Dry basidiomata of Favolaschia species A Favolaschia imbricata B Favolaschia miscanthi C, D Favolaschia sinarundinariae. Scale bars: 1 mm.

Microstructures

Basidiospores (6.8–)7–9 × 5–6.8 µm, L = 7.92 μm, W = 5.84 μm, Q = 1.31–1.38 (n = 90/3), broadly ellipsoid to subglobose, hyaline, thin-walled, smooth, with some guttules, faintly IKI+, CB–. Basidia 24–30 × 7–13 μm, cylindrical or clavate with some guttules, 4–spored, sterigmata 3–6 μm long; basidioles similar in shape to the basidia, but slightly smaller. Gloeocystidia and acanthocystidia absent. Cheilocystidia 25–39 × 7–14 μm, present at dissepiment edge, cylindrical or clavate, with obtuse diverticulate projections at the apex, thin-walled. Pileipellis hyphae subparallel, smooth to diverticulate, thin-walled, 5–11 µm diam.; terminal cells cystidioid or pyriform, smooth, thin-walled. Tramal hyphae subparallel, widely spaced in a gelatinous matrix, some with dense contents, some collapsed, thin-walled, 2–5 μm diam. Clamp connections present.

Figure 4. 

Basidiomata and microscopic structures of Favolaschia imbricata A, B basidiomata (Dai 24702, holotype) C basidiospores D hymenium in trama E basidia and basidioles F tramal hyphae G cheilocystidia at dissepiment edge H hyphae of pileipellis. Scale bars: 5 mm (A, B); 10 µm (C–H).

Additional specimens examined

China, Guangdong Province, Guangzhou, Baiyun District, Maofengshan Forest Park, dead bamboo, 15 Apr 2023, Dai 24703 (BJFC042257, paratype), Dai 24704 (BJFC042258, paratype).

Distribution and ecology

Favolaschia imbricata is distributed in subtropical area of Guangdong Province, China; it grows on dead bamboo and causes a white rot.

Favolaschia miscanthi Q.Y. Zhang, L.S. Bian, F. Wu & Y.C. Dai, sp. nov.

MycoBank No: 851808
Figs 3B, 5

Type

China, Guangxi Autonomous Region, Hezhou, Dazhongshan Forest Park, rotten Miscanthus, 19 Apr 2023, Dai 24652 (BJFC042179, holotype).

Etymology

“miscanthi” (Lat.): refers to the species inhabiting rotten Miscanthus.

Macrostructures

Basidiomata annual, gregarious, gelatinous when fresh and dry. Pilei 3–6 × 1.5–4 mm, conchoid or semicircular; pileal surface satin white (1A1–2A1) when fresh and drying, convex, transparent with a reticulate pattern matching the pores below, glabrous; margin straight, crenulate; context thin, transparent. Hymenophore concolorous with pileal surface, poroid, up to 40 pores per basidiocarp; mature pores 0.5–1 mm diam., polygonal, larger near the base and smaller near the edge, the marginal pores often incomplete; tubes up to 0.4 mm long. Stipe absent.

Microstructures

Basidiospores 7.5–10 × 5.5–7(–7.5) µm, L = 8.90 μm, W = 6.28 μm, Q = 1.37–1.46 (n = 60/2), broadly ellipsoid to subglobose, hyaline, thin-walled, smooth, with some guttules, faintly IKI+, CB–. Basidia 32–45 × 7–10 μm, narrowly clavate, tapered towards the base, apex broadly rounded, 4–spored, sterigmata 2–8 μm long; basidioles similar in shape to the basidia, but slightly smaller. Gloeocystidia and acanthocystidia absent. Cheilocystidia 17–34 × 7–10 μm, present at dissepiment edge, broom-shaped or irregular with small diverticulate projections on the sides and at the apex, thin-walled. Pileipellis hyphae interwoven, smooth to diverticulate, thin-walled, 4–7 µm in diam.; terminal cells cylindrical or clavate, with obtuse diverticulate projections at the apex, orientated perpendicular to pileal surface. Tramal hyphae interwoven, widely spaced in a gelatinous matrix, some with dense contents, some collapsed, thin-walled, 2–6 μm in diam. Clamp connections present.

Figure 5. 

Basidiomata and microscopic structures of Favolaschia miscanthi A, B basidiomata (A Dai 24652, holotype B Dai 24653) C basidiospores D hymenium in trama E basidia and basidioles F tramal hyphae G cheilocystidia at dissepiment edge H hyphae of pileipellis I terminal cells of pileipellis. Scale bars: 5 mm (A, B); 10 µm (C–I).

Additional specimen examined

China, Guangxi Autonomous Region, Hezhou, Dazhongshan Forest Park, rotten Miscanthus, 19 Apr 2023, Dai 24653 (BJFC042180, paratype).

Distribution and ecology

Favolaschia miscanthi is distributed in the subtropical area of the Guangxi Autonomous Region, China; it grows on rotten Miscanthus and causes white rot.

Favolaschia sinarundinariae Q.Y. Zhang, L.S. Bian, F. Wu & Y.C. Dai, sp. nov.

MycoBank No: 851809
Figs 3C, D, 6

Type

China, Sichuan Province, Bazhong, Nanjiang County, Micangshan National Forest Park, 25 Sep 2023, dead Sinarundinaria, Dai 26123 (BJFC043673, holotype).

Etymology

“sinarundinariae” (Lat.): refers to the species inhabiting dead Sinarundinaria.

Macrostructures

Basidiomata annual, gregarious, gelatinous when fresh and dry. Pilei 4–7 × 2–5 mm, conchoid to elliptic; pileal surface greyish-white (1B1) when fresh, dark grey (1F1) near the base upon drying, convex, transparent with a reticulate pattern matching the pores below, faintly pruinose when dry; margin straight, crenulate; context thin, opaque. Hymenophore concolorous with pileal surface, poroid, up to 80 pores per basidiomata; mature pores 0.4–1 mm diam., polygonal, the size varies irregularly, with some larger near the base or larger near the edges, the marginal pores often incomplete; tubes up to 0.5 mm long. Stipe absent.

Microstructures

Basidiospores 7–9 × 5–7(–7.5) µm, L = 7.87 μm, W = 6.08 μm, Q = 1.27–1.31 (n = 60/2), broadly ellipsoid to subglobose, hyaline, thin-walled, smooth, with some guttules, faintly IKI+, CB–. Basidia 40–50 × 7–9 μm, narrowly clavate, tapered towards the base, apex broadly rounded, 2 or 4–spored, sterigmata 5–10 μm long; basidioles similar in shape to the basidia, but slightly smaller. Gloeocystidia and acanthocystidia absent. Cheilocystidia 15–32 × 7–10 μm, present at dissepiment edge, antler-shaped or irregular with small diverticulate projections on the sides and at the apex, thin-walled. Pileipellis hyphae interwoven, with frequent diverticulate, thin-walled, 2–4 µm in diam., swollen to 8 µm on terminal; terminal cells irregular, with obtuse diverticulate projections at the apex, orientated perpendicular to pileal surface. Tramal hyphae interwoven, widely spaced in a gelatinous matrix, some with dense contents, some collapsed, thin-walled, 1.5–3 μm in diam. Clamp connections present.

Figure 6. 

Basidiomata and microscopic structures of Favolaschia sinarundinariae A, B basidiomata (A Dai 26123, holotype B Dai 26116) C basidiospores D hymenium in trama E basidia and basidioles F tramal hyphae G cheilocystidia at dissepiment edge H hyphae and terminal cells of pileipellis. Scale bars: 1 cm (A, B); 10 µm (C–H).

Additional specimens examined

China, Sichuan Province, Bazhong, Nanjiang County, Micangshan National Forest Park, 25 Sep 2023, dead Sinarundinaria, Dai 26129 (BJFC043679, paratype), Dai 26130 (BJFC, paratype), Dai 26134 (BJFC, paratype), Dai 26136 (BJFC, paratype), Dai 26142 (BJFC043692, paratype); Leshan, Ebian County, Heizhugou, 11 Aug 2023, dead Sinarundinaria, Dai 26115 (BJFC043665, paratype), Dai 26116 (BJFC043666, paratype).

Distribution and ecology

Favolaschia sinarundinariae is distributed in the subtropical area of Sichuan Province, China; it grows on rotten Sinarundinaria and causes white rot.

Discussion

The earliest records of Favolaschia species in China date back to the 20th century (Liu 1994; Liu and Yang 1994). Recently, with the application of molecular systematics, the species of Favolaschia in China have been continually updated (Liu 2020; Tolgor et al. 2021; Zhang and Dai 2021; Zhang et al. 2023). Morphological examination and phylogenetic analyses identified 16 species of Favolaschia in China (Zhang and Dai 2021; Zhang et al. 2023). In this study, three new species of Favolaschia are identified in south China: Favolaschia imbricata, F. miscanthi and F. sinarundinariae.

Phylogenetic studies of Favolaschia largely support the section classification, based on morphology proposed by Singer (1974). Morphologically, the F. sect. Anechinus includes two subsections, F. subsect. Rubrinae and F. subsect. Depauperatae, the former lacks gloeocystidia and gloeoporous hyphae (with granular contents); the latter has these structures, while the two subsections are not well distinguished in the phylogenetic tree. The three new species, Favolaschia imbricata, F. miscanthi and F. sinarundinariae, clustered within section Anechinus (group B), which is characterised by the lack of acanthocysts, gloeocystidia and gloeoporous hyphae replaced by irregular cystidia at the dissepiment edge or pileipellis.

In a combined ITS1-5.8S-ITS2-nLSU dataset-based phylogeny (Fig. 2), Favolaschia imbricata grouped with another new species F. miscanthi, with strong support (100/1.00). However, Favolaschia miscanthi can be easily distinguished from F. imbricata by its larger pores (0.5–1 mm vs. 0.2–0.6 mm), white pileal surface upon drying and growing on rotten Miscanthus, while F. imbricata shares dark grey or mouse-grey pileal surface upon drying and grows on dead bamboo. Based on the sub-generic classification of Favolaschia by Singer (1974), F. imbricata belongs to Favolaschia sect. Anechinus subsect. Rubrinae, because it lacks acanthocystidia, gloeocystidia and gloeoplerous hyphae.

Morphologically, Favolaschia imbricata with F. flabelliformis and F. tonkinensis share grey to greyish-white basidiomata when fresh and occur in south China. However, F. flabelliformis differs from F. imbricata by its flabelliform pilei and smaller pores (0.2–0.25 mm vs. 0.2–0.6 mm, Zhang et al. (2023)). F. tonkinensis differs from F. imbricata by its larger basidiomata (6–24 mm vs. 2–5 mm) and larger basidiospores (8–12.5 × 7–10.5 µm vs. 7–9 × 5–6.8 µm, Singer (1974)). In addition, F. imbricata has numerous cheilocystidia (cylindrical or clavate, with obtuse diverticulate projections at the apex) at the dissepiment edge, while they are absent in F. flabelliformis and F. tonkinensis.

Morphologically, Favolaschia miscanthi is characterised by its white basidiomata, inhabiting rotten Miscanthus, large pores (0.5–1 mm in diam.), broom-shaped cheilocystidia and broadly ellipsoid to subglobose basidiospores (7.5–10 × 5.5–7 µm) and belongs to F. sect. Anechinus subsect. Rubrinae. Four species in F. subsect. Rubrinae, viz. F. bannaensis, F. rigida, F. semicircularis and F. tenuissima, resemble F. miscanthi by sharing smaller (< 2 cm) and pure white to white basidiomata when fresh and occur in south China. However, Favolaschia bannaensis differs from F. miscanthi by its larger pores (up to 1.5 mm vs. up to 1 mm) and inhabit bamboo or rotten angiosperms (Zhang et al. 2023). Favolaschia rigida differs from F. miscanthi by its obvious stipe, smaller pores (0.25–0.33 mm vs. up to 0.5–1 mm) and the absence of cheilocystidia (Zhang et al. 2023). Favolaschia semicircularis is readily distinguished from F. miscanthi by its obvious stipe and shorter basidia (24–32 μm vs. 32–45 μm in length, Zhang et al. (2023)). Favolaschia tenuissima differs from F. miscanthi by its fewer pores (up to 20 vs. up to 40) and the presence of obvious stipe (Zhang et al. 2023).

Eight specimens from Heizhugou and Micangshan National Forest Park in Sichuan Province formed a support lineage (71/1.00) and there are two base pairs differences between them in the ITS regions and no differences in morphology. We described these as a new species, namely Favolaschia sinarundinariae. Phylogenetically, Favolaschia sinarundinariae and F. tephroleuca cluster together in a supported clade (57/1.00) within Favolaschia sect. Anechinus, with each forming highly-supported branches. However, F. tephroleuca is readily distinguished from F. sinarundinariae by its grey to dark grey pileal surface, when fresh and fewer pores (up to 40 vs. up to 80). Morphologically, Favolaschia flabelliformis is similar to F. sinarundinariae by sharing a similar pileal surface, the absence of stipe and almost the same size basidiospores (Zhang et al. 2023), but the former species has smaller basidiomata (2–3 mm vs. 4–7 mm), smaller pores (0.2–0.25 mm vs. 0.5–1 mm) and fewer pores (10–40 vs. up to 80, Zhang et al. (2023)). In addition, F. flabelliformis and F. sinarundinariae are phylogenetically distantly related.

During the investigations of Favolaschia, the information on distribution areas is obtained (Fig. 1). Regarding geographical distribution, Favolaschia is distributed in south China (the south Qinling-Huai River Line). In addition, an updated key to 19 species of Favolaschia is provided in the following.

A key to 19 species of Favolaschia found in China

1 Acanthocystidia present 2
– Acanthocystidia absent 5
2 Basidiomata apricot orange when fresh 3
– Basidiomata lemon chrome when fresh 4
3 Pilei < 5 mm F. minutissima
– Pilei > 5 mm F. brevibasidiata
4 Stipe < 5 mm in length F. brevistipitata
– Stipe > 5 mm in length F. longistipitata
5 Basidiomata campanulate F. manipularis
– Basidiomata conchoid, flabellate, reniform, semicircular or subcircular 6
6 Mature pilei > 2 cm in length 7
– Mature pilei < 2 cm in length 9
7 Stipe present F. tonkinensis
– Stipe absent 8
8 Mature pores polygonal, 3–6 mm in width F. pustulosa
– Mature pores round, 1–2.3 mm in width F. subpustulosa
9 Gloeocystidia present 10
– Gloeocystidia absent 11
10 Basidiospores 4.5–5.5 μm in width F. peziziformis
– Basidiospores 7–10 μm in width F. crassipora
11 Pilei transparent 12
– Pilei opaque 14
12 Stipe present, 1–3 × 0.5–1 mm F. tenuissima
– Stipe absent 13
13 Basidiomata inhabiting rotten Miscanthus F. miscanthi
– Basidiomata inhabiting dead bamboo (Bambusoideae) F. bannaensis
14 Pilei ash grey to fuscous or dark when fresh F. tephroleuca
– Pilei usually white to pale white when fresh 15
15 Cheilocystidia present at dissepiment edge 16
– Cheilocystidia absent 17
16 Stipe usually present F. semicircularis
– Stipe absent 17
17 Mature pores 0.2–0.6 mm in width F. imbricata
– Mature pores up to 1 mm in width F. sinarundinariae
18 Stipe present, with diverticulate hyphae in pileipellis F. rigida
– Stipe absent, without diverticulate hyphae in pileipellis F. flabelliformis

Acknowledgements

We thank Dr. Chao-Ge Wang (State Key Laboratory of Efficient Production of Forest Resources, School of Ecology and Nature Conservation, Beijing Forestry University) and Gang He (Jiujiang University) for helping in sample collection.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the National Natural Science Foundation of China (Project Nos. U23A20142, 32161143013), Fundamental Research Funds for the Central Non-profit Research Institution of the Chinese Academy of Forestry (Project No. CAFYBB2021MA007) and the Yunnan Province expert workstation programme (No. 202205AF150014).

Author contributions

Qiu-Yue Zhang and Wan-Ying Li designed the research and contributed to data analysis and interpretation. Qiu-Yue Zhang, Xin Zhang, Yu-Cheng Dai and Hong-Gao Liu prepared the samples and drafted the manuscript. Yu-Cheng Dai, Lu-Sen Bian and Fang Wu discussed the results and edited the manuscript. All authors contributed to the article and approved the submitted version.

Author ORCIDs

Qiu-Yue Zhang https://orcid.org/0000-0001-9458-3566

Hong-Gao Liu https://orcid.org/0000-0002-9508-3245

Wan-Ying Li https://orcid.org/0009-0006-7782-6670

Xin Zhang http://orcid.org/0009-0005-8363-7852

Yu-Cheng Dai http://orcid.org/0000-0002-6523-0320

Fang Wu https://orcid.org/0000-0002-6239-3634

Lu-Sen Bian https://orcid.org/0000-0003-3660-9056

Data availability

The sequences are deposited in the GenBank database (Table 1).

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Supplementary material

Supplementary material 1 

Three new species of Favolaschia (Mycenaceae, Agaricales) from South China

Qiu-Yue Zhang, Hong-Gao Liu, Wan-Ying Li, Xin Zhang, Yu-Cheng Dai, Fang Wu, Lu-Sen Bian

Data type: nex

Explanation note: In this study, the final alignments and the retrieved topologies were deposited in TreeBASE (http://treebase.org/treebase-web/home.html), under accession ID: 30973. At the same time, we upload it as an attachment to the editor.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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