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Campanophyllum microsporum (Agaricales, Agaricomycetes), Calocera multiramosa, and Dacrymyces naematelioides (Dacrymycetales, Dacrymycetes), three new species from Yunnan Province, southwestern China
expand article infoYuan-Hao Ma‡§|, Ping Liu‡§|, Hong-Mei Chai‡§|, Min Zeng‡§|, Yi-Yun Guo‡§|, Wei-Min Chen|‡§, Yong-Chang Zhao‡§|
‡ Biotechnology and Germplasm Resources Institute, Yunnan Academy of Agricultural Sciences, Kunming, China
§ Yunnan Provincial Key Lab of Agricultural Biotechnology, Kunming, China
| Ministry of Agriculture and Rural Affairs, Kunming, China

Corresponding author: Wei-Min Chen ( chwmkm@aliyun.com )

Corresponding author: Yong-Chang Zhao ( yaasmushroom@aliyun.com )

Academic editor: Yupeng Ge
© 2024 Yuan-Hao Ma, Ping Liu, Hong-Mei Chai, Min Zeng, Yi-Yun Guo, Wei-Min Chen, Yong-Chang Zhao.
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: Ma Y-H, Liu P, Chai H-M, Zeng M, Guo Y-Y, Chen W-M, Zhao Y-C (2024) Campanophyllum microsporum (Agaricales, Agaricomycetes), Calocera multiramosa, and Dacrymyces naematelioides (Dacrymycetales, Dacrymycetes), three new species from Yunnan Province, southwestern China. MycoKeys 107: 327-350. https://doi.org/10.3897/mycokeys.107.125571
Open Access

Abstract

Three new species belonging to Basidiomycota from southwestern China are described based on morphological and molecular data. Campanophyllum microsporum is morphologically characterized by dorsally pseudostipitate, pale orange to brownish orange pileus, excentric to lateral pseudostipe, crowded lamellae, cylindrical-ellipsoid basidiospores 3.0–4.2 × 1.7–2.2 µm, narrowly clavate to clavate basidia 14.5–23.0 × 3.0–4.2 µm, and cylindrical to clavate cheilocystidia 22.0–55.0 × 5.0–10.8 µm. Calocera multiramosa is morphologically characterized by stipitate, yellowish to orange, dendroid, and dichotomously branched basidiomata, cylindrical to clavate basidia 36.5–52.5 × 3.8–6.1 µm, navicular or reniform, 1–5-septate mature basidiospores 10.4–16.7 × 5.2–7.4 µm. Dacrymyces naematelioides is morphologically characterized by stipitate and cerebriform, orange to light brown basidiomata, cylindrical to clavate, smooth or roughened basidia 38.5–79.5 × 6.5–10.6 µm, broadly and elliptic-fusiform, 7-septate mature basidiospores 18.5–28.6 × 8.9–13.8 µm. These three new species are supported by the phylogenetic analyses using maximum likelihood (ML) and Bayesian inference (BI) analyses with combined nuclear ribosomal DNA (rDNA) internal transcribed spacer (ITS) and large ribosomal subunit (LSU) sequences. Full descriptions and photographs of these new species are provided.

Key words

Basidiomycota, new taxon, phylogenetic analyses, taxonomy

Introduction

The monotypic genus Campanophyllum Cifuentes & R.H. Petersen was proposed to accommodate Lentinus proboscideus Fr., traditionally contains dorsally pseudostipitate pileus with tricholomataceus, excentric to lateral pseudostipe, crowded lamellae, cylindrical-ellipsoid spores, cylindrical, clavate to utriform cheilocystidia, and grows on rotten wood (Cifuentes et al. 2003). L. proboscideus was combined into the genus Campanophyllum in the family Cyphellaceae, and designated a neotype by Cifuentes et al. (2003). In addition, the authors demonstrated that this species represents a novel species of a novel genus distinct from its closest relatives through a comprehensive analysis of morphological characteristics, molecular data, and sexual compatibility (Cifuentes et al. 2003). The species was found in montane forests in Colombia, Costa Rica, Ecuador, Mexico, and Panama, and is currently being assessed for inclusion in the IUCN red list as endangered (https://redlist.info/iucn/species_view/488791/) (Cifuentes et al. 2003; Reschke et al. 2021). Recently, researchers also collected the species of C. proboscideum in India and studied its fungal extracts, which are rich in natural antioxidants and highly effective antimicrobial activity (Borthakur et al. 2020). However, the specimens collected from India may not be of C. proboscideum, but of another species, due to the huge differences in the ITS sequences. The absence of detailed morphological descriptions precludes the ability to ascertain the specific species to which the specimen belongs.

Calocera (Fr.) Fr. and Dacrymyces Nees are the two major polyphyletic groups in the family Dacrymycetaceae, characterized by pulvinate to dendroid or cerebriform basidiomata (Shirouzu et al. 2007, 2009, 2013, 2017; Zamora and Ekman 2020; Fan et al. 2021; Lian et al 2022; Zamora et al. 2022). Their classifications in Dacrymycetes are based on morphology and have remained unaltered (McNabb 1965, 1973; Shirouzu et al. 2017; Zamora and Ekman 2020). However, this morphology-based classification has often conflicted with the results of molecular phylogenetic analyses, and Calocera, Dacrymyces, and Dacryopinax G.W. Martin have been shown to be non-monophyletic genera (Shirouzu et al. 2013, 2017; Zamora and Ekman 2020). Phylogenetic analysis shows that species of Calocera and Dacrymyces are distributed in many clades of the family Dacrymycetaceae. The three species of C. cornea (Batsch) Fr., C. lutea (Massee) McNabb, and C. fusca Lloyd were clustered into three distinct clades, rather than forming a single clade, despite belonging to the same genus, and many of the species of Dacrymyces were grouped with other genera in one clade (Shirouzu et al. 2017; Zamora and Ekman 2020).

Calocera is ecologically saprobic, causing brown rot except C. viscosa (Pers.) Bory, and C. lutea which are white rot species (Shirouzu et al. 2009, 2013). According to the Index Fungorum (https://www.indexfungorum.org) as of June 2024, 95 species names of Calocera are recorded. In China, only six species have been reported: C. sinensis McNabb, C. hunanensis B. Liu & K. Tao, C. mangshanensis B. Liu & L. Fan, C. morchelloides B. Liu & L. Fan, C. bambusicola Sheng H. Wu, and C. tibetica F. Wu, L.F. Fan & Y.C. Dai (McNabb 1965; Liu et al. 1988; Liu and Fan 1989, 1990; Fan et al. 2021). Dacrymyces described by Nees (1816) based on D. stillatus, is treated as a genus of saprotrophic fungi (Shirouzu et al. 2009; Zamora and Ekman 2020; Zamora et al. 2022). A total of 234 species names of Dacrymyces is recorded in the Index Fungorum in June 2024, and the genus appears to be the most polyphyletic in the phylogeny of the Dacrymycetales (Shirouzu et al. 2017; Zamora and Ekman 2020; Savchenko et al. 2021; Zamora et al. 2022).

The Laojun Mountain is one of the main parts of the Three Parallel Rivers of Yunnan Protected Areas (TPRYPA), the World Natural Heritage Site, in northwest Yunnan Province, southwestern China. The TPRYPA is part of the Mountains of Southwest China Biodiversity Hotspot, which includes 12,000 plant species, 29 percent of which are found nowhere else (Zhang et al. 2010; Mittermeier et al. 2011). The Laojun Mountain is located between 26°2.80'–27°36.60'N, 99°1.20'–99°54.60'E and includes four counties, including Yulong, Jianchuan, Lanping, and Weixi, with an area of about 108,500 hm2 and elevations ranging from 2,100 to 4,513 m (Zhang et al. 2010). The dominant tree species in Laojun Mountain are Abies sp., Acer sp., Betula sp., Cyclobalanopsis sp., Fargesia sp., Lithocarpus sp., Picea sp., Pinus sp., Quercus sp., Rhododendron sp., and Sorbus sp. (Wu and Zhu 1987).

During the investigation of the diversity of macrofungi in the Laojun Mountain, a multitude of specimens, including a dozen belonging to the Campanophyllum genus and several belonging to the genera Calocera and Dacrymyces, were collected from July to September 2019–2023. In this study, the specimens of these three new species were collected from the same position in a deciduous forest of the Laojun Mountain. With the combination of morphological observations and phylogenetic analyses, we described three new species, namely Campanophyllum microsporum, Calocera multiramosa, and Dacrymyces naematelioides.

Materials and methods

Specimen collection, morphological observation, and isolation

The fungal specimens used in this study were collected from the Laojun Mountain in northwestern Yunnan Province, China. After collection, the specimens were dried in an electric drier at ca. 45 °C, and deposited in the Herbarium of Cryptogams, Kunming Institute of Botany of the Chinese Academy of Sciences (HKAS). Macromorphological characteristics and habitats were obtained from field notes and photographs. Color codes were based on Kornerup and Wanscher (1978). Micromorphological features were observed from the dried specimens and measured and photographed in 5% KOH solution (w/v) and 1% Congo Red solution (w/v) using a Leica DM6 B upright light microscope and Leica Application Suite X (LAS X, version 3.7.5). In the description of basidiospores, the abbreviations m/n/p denote m basidiospores measured from n basidiomata of p collections. The dimensions of the microscopic structures are given as (a–) b–c (–d), in which b–c contains at least 90% of the measured values, and (a–) and (–d) are the extreme values provided in parentheses. The Q value stands for the ratio of length/width of an individual basidiospore and basidium, and Lm/Wm/Qm refers to the average length/width/Q value of all basidiospores (Na et al. 2022; Wei et al. 2024). The strains of Campanophyllum microsporum were isolated from the inner tissue of fresh basidiomata using a Yeast Extract Peptone Dextrose Agar (YPD) Medium consisting of 2 g yeast extract (Beijing Aoboxing Biotech Co., Ltd.), 2 g peptone (Beijing Aoboxing Biotech Co., Ltd.), 20 g dextrose (Tianjin Fengchuan Chemical Reagent Co., Ltd.), 13 g agar (Biosharp Life Sciences), and 1000 mL distilled water. The living cultures were preserved at the National Germplasm Bank of Edible Mushroom (Yunnan). Their isolate IDs are YAASM 7490 and 7491.

DNA extraction, PCR amplification and sequencing

The genomic DNA was extracted from the dry specimens and cultured mycelia using the Fungal gDNA kit GD2416 (Biomiga CA, USA) following the manufacturer’s instructions. The entire ITS and partial LSU of the nrDNA region were amplified from the total DNA using the primer pair ITS5/ITS4 (White et al. 1990) and LR0R/LR7 (Vilgalys and Hester 1990; Moncalvo et al. 2000), respectively, and no DNA template was used as the negative control. The PCR cycling for the amplification of both ITS and LSU was as set follows: an initial denaturation at 95 °C for 3 min, followed by 34 cycles of 95 °C for 30 s, 56 °C for 1 min, 72 °C for 1 min, and a final extension at 72 °C for 5 min. The PCR products were sequenced bi-directionally by Tsingke Biotechnology Co., Ltd. Kunming, China. Newly generated sequences of both directions were assembled using the software SeqMan version 11.1.0 (DNASTAR, Inc.), and submitted to GenBank (accession nos. ITS: PP550870–PP550882, LSU: PP550017–PP550027).

Sequence alignment and phylogenetic analyses

The sequences used in this study were those retrieved from GenBank combined with newly generated sequences. Taxon information and GenBank accession numbers of all the sequences are listed in Table 1. All sequences were aligned using the software MAFFT 7.503 (Katoh and Standley 2013) with the default settings and edited manually using BioEdit 7.2.5 (Hall 1999). After alignment, the ITS and LSU datasets were concatenated using the program SequenceMatrix 1.8.1 (Vaidya et al. 2011). Phylogenetic analyses of Cyphellaceae and Dacrymycetaceae were performed using maximum likelihood (ML) and Bayesian inference (BI) analyses based on the sequences matrix on the personal computer. The best-fit models for the concatenated ITS+LSU dataset were selected according to the Akaike Information Criterion (AIC) in jModelTest 2.1.10 (Guindon and Gascuel 2003; Darriba et al. 2012). ML analyses of the concatenated ITS+LSU dataset of Cyphellaceae and Dacrymycetaceae were performed using RAxML-NG 1.1.0 (Kozlov et al. 2019) under the GTR+I+G model with 1,000 bootstrap replicates. BI analyses of Cyphellaceae and Dacrymycetaceae were implemented using MrBayes 3.2.7 (Ronquist et al. 2012) under the GTR+I+G model. There were four independent runs, each of which had four chains for 15,000,000 generations sampling from the posterior distribution every 1000th generation. The first 25% of the sampled trees were discarded as burn-in, while the remaining trees were used to obtain the Bayesian posterior probabilities of the clades. The constructed phylogenetic trees were visualized and edited in FigTree 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/) and Adobe Illustrator 25.3.1. Flammulina velutipes (Curtis) Singer, was used as an outgroup in the phylogeny of Cyphellaceae (Vizzini et al. 2022), while Suillus pictus (Peck) Kuntze, and Coprinus comatus (O.F. Müll.) Pers., were used as the outgroup in the phylogeny of Dacrymycetaceae (Shirouzu et al. 2013). The final alignments and the retrieved topologies were deposited in TreeBASE (http://purl.org/phylo/treebase/phylows/study/TB2:S31416) with submission ID 31416.

Table 1.
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Taxa used in the phylogenetic analyses and their corresponding GenBank accession numbers. Newly generated sequences are in bold. Type materials are marked with ‘T’.

Species Isolate ID /Voucher Country GenBank Accession Numbers Reference
ITS LSU
Agaricomycetes
Campanophyllum microsporum – /HKAS 133167 China PP550870 PP550018 this study
C. microsporum – /HKAS 133168 China PP550871 PP550019 this study
C. microsporum – /HKAS 133169 China PP550872 PP550020 this study
C. microsporum – /HKAS 133170T China PP550873 PP550017 this study
C. proboscideum – /TENN56402 Mexico AY230866 AY230866 Cifuentes et al. (2003)
C. proboscideum – /TENN56427 Mexico AY230867 AY230867 Cifuentes et al. (2003)
C. proboscideum – /PA46 Panama MW386067 – Reschke et al. (2021)
C. proboscideum – /PAN327 Panama MW386071 – Reschke et al. (2021)
C. proboscideum – /PAN373 Panama MW386072 – Reschke et al. (2021)
C. proboscideum – /NEHU.MBSRJ. 38 India KP843881 – unpublished
Chondrostereum coprosmae – /PDD: 119544 New Zealand OL709440 – unpublished
C. coprosmae – /PDD: 89940 New Zealand OL709437 OL709436 unpublished
C. purpureum HHB-13334-sp. /– USA AF518607 – Hibbett and Binder (2002)
C. purpureum SFI-B18 /– Ireland MT535785 MT559785 unpublished
C. purpureum 14-2300 /– USA MG774405 – Merlet et al. (2018)
C. purpureum CBS 350.53 /– France MH857241 MH868775 Vu et al. (2019)
C. vesiculosum – /PDD: 119640 New Zealand OR607672 – unpublished
Cunninghammyces umbonatus – /He 5316 China MW557955 – unpublished
C. umbonatus – /He 5311 China MW557940 MW557954 unpublished
C. umbonatus – /He 5313 China MW557941 – unpublished
Cyphella digitalis – /PVKU3421 Czech Republic OM837174 – Holec et al. (2022)
C. digitalis Thorn-617 /– USA AY293175 – Binder et al. (2005)
C. digitalis CBS 679.82 /– USA DQ486698 AY635771 Matheny et al. (2006)
Gloeostereum incarnatum G1905 /HCC-3 Russia MK278092 – Varga et al. (2019)
G. incarnatum – /KUC20131022-28 South Korea KJ668540 KJ668393 Jang et al. (2015)
G. incarnatum 3332 /– Sweden AF141637 – Parmasto and Hallenberg (2000)
G. incarnatum – /NIFoS 1948 South Korea MH992519 – unpublished
G. incarnatum BCC 41461 /– Thailand KY614001 KY614002 unpublished
G. cimri CBS 145006T /– Netherlands MT023735 MN266884 Ahmed et al. (2020)
Granulobasidium vellereum G0482 /DK 2781 Poland MK278094 – Varga et al. (2019)
G. vellereum CBS 52.84 /– USA AY745729 – unpublished
G. vellereum – /B. Gilsenius (GB) Sweden DQ677490 – Larsson (2007)
G. vellereum BAFCcult 4367 /– Argentina KC881193 – Robles et al. (2015)
G. vellereum TJU_NOV19 /– China OM237077 – unpublished
Incrustocalyptella columbiana – /K:237992 United Kingdom MW830122 – unpublished
Flammulina velutipes AFTOL-ID 558 /– USA AY854073 AY639883 unpublished
Dacrymycetes
Calocera cornea CBS 124.84 /– Canada AB712437 AB472738 Shirouzu et al. (2013)
C. cornea ICMP 20465 /PDD 104991 New Zealand LC131403 LC131362 Shirouzu et al. (2017)
C. cornea AFTOL-ID 438 /– unknown AY789083 AY701526 unpublished
C. cornea ICMP 21223 /PDD 107847 New Zealand LC131404 LC131363 Shirouzu et al. (2017)
C. cornea – /UPS F-940774 Sweden MN595626 MN595626 Zamora and Ekman (2020)
C. cornea – /CWU(MYC)6922 Ukraine MW191969 MW159089 Savchenko et al. (2021)
C. furcata – /H:Spirin 10949 Russia MW191975 MW159088 Savchenko et al. (2021)
C. furcata – /TU135016 Estonia MW191958 MW159087 Savchenko et al. (2021)
C. tibetica – /Dai20171T China MW549777 MW750403 Fan et al. (2021)
C. tibetica – /Dai20178 China MW549778 MW750404 Fan et al. (2021)
C. multiramosa – /HKAS 133171T China PP550874 PP550021 this study
C. multiramosa – /HKAS 133172 China PP550875 PP550022 this study
C. multiramosa – /HKAS 133173 China PP550876 PP550023 this study
C. viscosa AFTOL-ID 1679 /MW 591 Germany DQ520102 DQ520102 unpublished
C. viscosa TUFC12873 /TNS-F-15704 Japan AB712439 AB299048 Shirouzu et al. (2013)
C. viscosa – /UPS F-940773 Sweden MN595628 MN595628 Zamora and Ekman (2020)
C. viscosa – /CWU(MYC)6937 Ukraine MW191970 MW159090 Savchenko et al. (2021)
Cerinomyces aculeatus – /TUMH61942 (TUFC50098)T Japan MW191955 MW159053 Savchenko et al. (2021)
C. atrans TUFC 30545 /– Canada AB712443 AB712423 Shirouzu et al. (2013)
C. borealis – /O160848T Norway MW191890 MW159042 Savchenko et al. (2021)
C. brevisetus – /URM:Chikowski 1544T Brazil MW191886 MW159046 Savchenko et al. (2021)
C. creber – /UPS:F-946512T Spain MW191985 MW191985 Savchenko et al. (2021)
C. enatus TUFC12876 /TNS-F-21034 Japan AB712441 AB472696 Shirouzu et al. (2013)
C. ramosissimus CFMR:FP-150848T /– Belize AB712446 AB712426 Shirouzu et al. (2013)
Dacrymyces burdsallii CFMR:HHB-6908T /– USA AB712444 AB712424 Shirouzu et al. (2013)
D. capitatus – /Dai 20023 China OL587808 OL546776 unpublished
D. capitatus CBS 293.82 /– Canada AB712450 AB472741 Shirouzu et al. (2013)
D. ceraceus CFMR:HHB-8969T /– USA AB712442 AB712422 Shirouzu et al. (2013)
D. chrysocomus – /UPS:F-940136 Spain MN595629 MN595629 Zamora and Ekman (2020)
D. chrysocomus – /UPS:F-940134 Sweden MN595630 MN595630 Zamora and Ekman (2020)
D. chrysospermus TUFC13115 /TNS-F-15712 Japan AB712452 AB299073 Shirouzu et al. (2013)
D. chrysospermus – /H:Spirin 10795 Russia MW191974 MW159078 Savchenko et al. (2021)
D. chrysospermus – /H:Miettinen 14818 USA MW191961 MW159077 Savchenko et al. (2021)
D. aff. Chrysospermus – /UPS:F-593536 Japan MN595631 MN595631 Zamora and Ekman (2020)
D. dictyosporus CFMR:HHB-8618 /– USA AB712454 AB712429 Shirouzu et al. (2013)
D. estonicus – /UPS:F-940137 Sweden MN595632 MN595632 Zamora and Ekman (2020)
D. estonicus – /UPS:F-940138 Sweden MN595633 MN595633 Zamora and Ekman (2020)
D. fennicus – /H:Miettinen 21174 Finland MW191957 MW159071 Savchenko et al. (2021)
D. fennicus – /UPS:F-946596 Sweden MZ147627 MZ147627 Savchenko et al. (2021)
D. grandinioides – /H7008841 Kenya MW191950 MW159076 Savchenko et al. (2021)
D. lacrymalis TUFC13327 /TNS-F-15719 Japan AB712456 AB299069 Shirouzu et al. (2013)
D. cf. minor – /H:Miettinen 19137 Finland MW191967 MW159080 Savchenko et al. (2021)
D. cf. minor – /H:Miettinen 20591 Finland MW191965 MW159079 Savchenko et al. (2021)
D. naematelioides – /HKAS 133174aT China PP550877 PP550024 this study
D. naematelioides – /HKAS 133174bT China PP550878 PP550025 this study
D. naematelioides – /HKAS 133174cT China PP550879 PP550026 this study
D. naematelioides – /HKAS 133174dT China PP550880 PP550027 this study
D. naematelioides YAASM 7490 /– China PP550881 – this study
D. naematelioides YAASM 7491 /– China PP550882 – this study
D. ovisporus – /H:Miettinen 20787 Finland MW191964 MW159074 Savchenko et al. (2021)
D. ovisporus – /H:Spirin 11145 Norway MW191960 MW159073 Savchenko et al. (2021)
D. pinacearum – /UPS:F-593533 Japan MN595637 MN595637 Zamora and Ekman (2020)
D. pinacearum – /UPS:F-593535 Japan MN595638 MN595638 Zamora and Ekman (2020)
D. puniceus TUFC12833 /TNS-F-15711 Japan AB712449 AB299057 Shirouzu et al. (2013)
D. puniceus – /Wu180 China OL587812 OL546780 unpublished
D. sinostenosporus – /Dai 20003T China MW540888 MW540890 Lian et al. (2022)
D. sinostenosporus – /Dai 20008 China MW540889 MW540891 Lian et al. (2022)
D. sobrius CFMR:RLG-13487T /– USA AB712445 AB712425 Shirouzu et al. (2013)
D. stenosporus ICMP 20488 /PDD 105018T New Zealand LC131433 LC131396 Shirouzu et al. (2017)
D. stenosporus ICMP 21237 /PDD 107970 New Zealand LC131434 LC131397 Shirouzu et al. (2017)
D. stillatus (anamorph) – /UPS:F-939814 Sweden MN595676 MN595676 Zamora and Ekman (2020)
D. stillatus (anamorph) – /UPS:F-939816 Sweden – MN593494 Savchenko et al. (2021)
D. stillatus (teleomorph) – /UPS:F-939814 Sweden MN595677 MN595677 Zamora and Ekman (2020)
D. stillatus (teleomorph) – /UPS:F-939816 Sweden – MN593495 Savchenko et al. (2021)
D. subalpinus TUFC12834 /TNS-F-15730 Japan AB712465 AB299060 Shirouzu et al. (2013)
D. venustus – /O:Adane 150T Ethiopia MW191949 MW159075 Savchenko et al. (2021)
Dacryonaema macnabbii – /UPS:F-940949 Sweden MN595650 MN595650 Zamora and Ekman (2020)
D. macnabbii – /UPS:F-940992 Sweden MN595653 MN595653 Zamora and Ekman (2020)
D. macrosporum – /UPS:F-940998 Finland MN595660 MN595660 Zamora and Ekman (2020)
D. macrosporum – /UPS:F-941001 Finland MN595661 MN595661 Zamora and Ekman (2020)
D. rufum – /UPS:F-941005 Sweden MN595646 MN595646 Zamora and Ekman (2020)
D. rufum – /UPS:F-941012 Finland MN595649 MN595649 Zamora and Ekman (2020)
Dacryopinax elegans – /TENN 066927 USA MN595640 MN595640 Zamora and Ekman (2020)
Dacryopinax sp. – /H7008759 Kenya MW191959 MW159091 Savchenko et al. (2021)
D. spathularia TUFC12846 /TNS-F-21048 Japan AB712473 AB472710 Shirouzu et al. (2013)
D. spathularia FCME 27539 /– Mexico MN733711 MN733722 Castro-Santiuste et al. (2020)
D. spathularia – /H:Miettinen 20559 Indonesia MW191976 MW159092 Savchenko et al. (2021)
Dendrodacrys ciprense – /UPS:F-946590T Cyprus OM519385 OM519385 Zamora et al. (2022)
D. ciprense – /UPS:F-946591 Cyprus OM519386 OM519386 Zamora et al. (2022)
D. concrescens – /UPS:F-946602T Sweden OM519390 OM519390 Zamora et al. (2022)
D. ellipsosporum – /UPS:F-946604T Spain OM519392 OM519392 Zamora et al. (2022)
D. oblongisporum – /UPS:F-979568T Spain OM519400 OM519400 Zamora et al. (2022)
Ditiola peziziformis – /H:Haikonen 24269 Finland MW191972 MW159070 Savchenko et al. (2021)
D. peziziformis – /H:Haikonen 30097 Finland MN595642 MN595642 Zamora and Ekman (2020)
D. radicata – /H:Miettinen 20590.2 Finland MW191966 MW159083 Savchenko et al. (2021)
D. radicata – /UPS:F-939957 Sweden MN595641 MN595641 Zamora and Ekman (2020)
Guepiniopsis buccina – /CWU(MYC)7014 Ukraine MW191971 MW159086 Savchenko et al. (2021)
G. buccina – /UPS:F-940947 Spain MN595643 MN595643 Zamora and Ekman (2020)
Unilacryma unispora – /UPS:F-941279 Sweden MN595667 MN595667 Zamora and Ekman (2020)
U. bispora – /UPS:F-941254 Sweden MN595670 MN595670 Zamora and Ekman (2020)
U. bispora – /UPS:F-941266 Sweden MN595674 MN595674 Zamora and Ekman (2020)
Dacrymycetes sp. NBRC 110592 /– Japan LC004003 LC003884 Shirouzu et al. (2016)
Coprinus comatus AFTOL-ID 626 /– USA AY854066 AY635772 Shirouzu et al. (2013)
Suillus pictus AFTOL-ID 717 /– USA AY854069 AY684154 Shirouzu et al. (2013)

Results

Phylogenetic analyses

In the phylogeny of Cyphellaceae, 36 sequences were used for phylogenetic analyses, of which four sequences were newly generated in this study. The concatenated dataset of ITS and LSU sequences comprised a total of 1695 characters. ML and BI analyses generated similar topologies, so only the ML tree is presented along with the support values from the Maximum likelihood bootstrap (BS, >75%) values and Bayesian inference (BI) posterior probabilities (PP, >0.95) (Fig. 1). The phylogeny revealed that Cyphellaceae was divided into three clades, and these three genera of Incrustocalyptella Agerer, Cyphella Fr., and Campanophyllum constituted one of the three major clades with strong statistical supports (99% BS, 1.00 PP). In the ML tree, the phylogenetic results demonstrated that Campanophyllum microsporum formed a distinct lineage closely related to C. proboscideum and Campanophyllum sp. (Voucher NEHU.MBSRJ. 38) with strong statistical supports (98% BS, 0.99 PP). The ITS sequences from C. microsporum and C. proboscideum were markedly different, with ca. 105 different nucleobases, and the ITS sequences of C. microsporum and Campanophyllum sp. (Voucher NEHU.MBSRJ. 38) were different with about 20 different nucleobases.

Figure 1. 

Maximum likelihood (ML) tree of Cyphellaceae based on the combined ITS+LSU dataset. ML bootstrap values (BS > 75%) and Bayesian posterior probabilities (PP > 0.95) are shown at the nodes in the order of BS/PP. The tree is rooted with Flammulina velutipes. The new taxon is indicated in bold.

In the phylogeny of Dacrymycetaceae, the concatenated dataset of LSU and ITS sequences comprised a total of 1649 characters. 94 sequences were used for phylogenetic analyses, of which nine sequences were newly generated in this study. ML and BI analyses generated similar topologies, so only the ML tree is presented (Fig. 2). Within Dacrymycetes, we distinguish four main groups, Dacrymycetaceae (clade A), Cerinomycetaceae (clade B), Dacryonaemataceae (clade C), and Unilacrymaceae (clade D). The clade A included several genera and the majority of species, and formed a sister group to the clades B, C, and D with strong statistical support (95% BS, 1.00 PP). Samples of the two new species were placed in the clade A, and one of the new species of Dacrymyces naematelioides formed a non-monophyletic, strongly supported group. The new species Calocera multiramosa was found to be closely related to C. tibetica with high supports (85% BS, 1.00 PP), and the two species clustered together with C. viscosa with strong supports (91% BS, 1.00 PP). The new species D. naematelioides formed a sister lineage to D. chrysospermus with 78% bootstrap support and 1.00 posterior probability.

Figure 2. 

Maximum likelihood (ML) tree of Dacrymycetaceae based on the combined ITS+LSU dataset. ML bootstrap values and Bayesian posterior probabilities are shown at the nodes in the order of BS/PP. The tree is rooted with Suillus pictus and Coprinus comatus. The new taxon is indicated in bold.

Taxonomy

Campanophyllum microsporum Y.H. Ma, W.M. Chen & Y.C. Zhao, sp. nov.

MycoBank No: 853503
Figs 3, 4, 5

Diagnosis

Campanophyllum microsporum is characterized by dorsally pseudostipitate pileus, excentric to lateral pseudostipe, crowded lamellae, cylindrical-ellipsoid basidiospores (3.0–4.2 × 1.7–2.2 µm), narrowly clavate to clavate basidia (14.5–23.0 × 3.0–4.0 µm), and cylindrical to clavate cheilocystidia (22.0–55.0 × 5.0–11.0 µm); occurrence in a deciduous forest and solitary, cespitose, scattered, or gregarious habit on rotten wood.

Type

China. Yunnan Province: Jianchuan County, Laojunshan Town (26°35.85'N, 99°40.44'E, elev. 3100 m), on rotten wood, 21 September 2023, Yuan-Hao Ma, Min Zeng & Wei-Min Chen (Holotype: HKAS 133170!, ex-type: YAASM 7187).

Etymology

The epithet “microsporum” refers to the smaller basidiospores compared to Campanophyllum proboscideum.

Description

Basidiomata pseudostipitate, dorsally and eccentrically or laterally attached to substrate, occasionally central, pendent, broadly cyphelloid to crepidotoid, lamellate. Pileus 5.0–12.0 × 4.0–9.0 cm, spathulate, flabelliform to rounded-flabelliform, sometimes subcircular; plano-convex when young and applanate when older, margin inrolled, lobate when fully expanded; surface moist, initially pale orange (5A2–4), greyish orange (5B2–3), or light orange (6A2–5), then brownish orange (6C5–6), light brown (6D5–8), often with small stains of darker colors. Context thick, fleshy, whitish, and unchanging in color when injured. Lamellae extending radially from attachment point within pseudostipe, very crowded, sometimes forked, white to off-white, sometimes with small blackish stains. Pseudostipe 0.5–2.5 × 0.4–1.0 cm, concolorous with pileus, discolouring to blackish-ochre (6E5-7, 6F7). Spore print white. Taste mild, odor indistinct.

Figure 3. 

Basidiomata of Campanophyllum microsporum in the field A HKAS 133170 (Holotype) B HKAS 133169. Photos by Y.H. Ma. Scale bars: 3 cm.

Basidiospores [149/7/4] (2.7–)3.0–4.2(–4.5) × (1.5–)1.7–2.2(–2.6) µm, Lm = 3.5 µm, Wm = 1.9 µm, Q = 1.4–2.5, Qm = 1.8, cylindrical-ellipsoid, smooth, hyaline, thin-walled, inamyloid. Basidia (13.5–)14.5–23.0(–26.0) × (2.3–)3.0–4.2(–4.6) µm, Lm = 17.6 µm, Wm = 3.6 µm, Q = 3.6–7.4, Qm = 4.9, narrowly clavate to clavate, 4-spored, sterigma 0.9-2.2 µm. Cheilocystidia abundant, (17.5–)22.0–55.0(–59.0) × (4.2–)5.0–10.8(–13.9) µm, Lm = 37.0 µm, Wm = 7.2 µm, hyaline, thin-walled, mostly cylindrical to clavate, sometimes lageniform, rod-like, or beaked-utriform, pedunculate (1.9–12.5 × 1.8–4.1 µm). Pleurocystidia not observed. Lamellar trama hyaline, parallel, hyphae 3.6–17.6 µm in diameter, thin- to thick-walled. Pileipellis composed of repent, parallel hyphae, 4.2–11.5 (–17.0) µm in diameter, sometimes with yellow-brown, intracellular pigments. Clamp connections present in all tissues of basidiomata.

Figure 4. 

Morphological features of Campanophyllum microsporum on YPD medium after 20 days in the dark in a 9 cm Petri plate (ex-type YAASM 7187) A surface of colony B reverse of colony. Photos by Y.H. Ma. Scale bars: 2 cm.

Culture characteristics. Colonies grown on YPD reaching 40 mm radius within 20 days at 22 °C in the dark, forming abundant aerial mycelium, usually zonate. Mycelium irregularly cottony, with common clamp connections, pallid mouse gray to pale brown in aerial mycelium with age, easily forming basidiomata in the Petri plate.

Figure 5. 

Microscopic structures of Campanophyllum microsporum (Holotype HKAS 133170) A basidiospores in Congo red B basidia in Congo red C–H cheilocystidia (C–E in KOH solution F–H in Congo red). Photos by Y.H. Ma. Scale bars: 10 µm.

Habitat and distribution

Solitary, cespitose, scattered, or gregarious on rotten wood in a deciduous forest; known from Yunnan, China.

Additional specimens examined

China, Yunnan Province: Jianchuan County, Laojunshan Town, 7 July 2022, Yuan-Hao Ma, Ping Liu & Yong-Chang Zhao (HKAS 133167, HKAS 133168); 26 July 2023, Yuan-Hao Ma & Ping Liu (HKAS 133169).

Notes

Campanophyllum microsporum is similar to C. proboscideum in both macro- and micro-morphology, including broadly cyphelloid to crepidotoid basidiomata, spathulate, flabelliform to rounded-flabelliform pileus, and very crowded lamellae; cylindrical-ellipsoid basidiospores, narrowly clavate to clavate basidia. However, several other features can distinguish the two species. Morphologically, the new species have smaller basidiospores (3.0–4.2 × 1.7–2.2 µm vs. 4–4.5 × 2–3 µm), slenderer and longer basidia (14.5–23.0 × 3.0–4.2 µm vs. 14–17 × 4.5–5.0 µm), and larger cheilocystidia (22.0–55.0 × 5.0–10.8 µm vs. 18–25 × 9–11 µm) (Cifuentes et al. 2003).

Calocera multiramosa Y.H. Ma, W.M. Chen & Y.C. Zhao, sp. nov.

MycoBank No: 853504
Figs 6, 7, 8

Diagnosis

Calocera multiramosa differs from other species of the genus by yellowish to orange basidiomata, dendroid and dichotomously branches, branched, smooth, thin-walled marginal hyphae (2.0–4.8 µm), branched, thin-walled internal hyphae (2.9–10.0 µm), cylindrical to clavate basidia (36.5–52.5 × 4.0–6.0 µm), 1–5-septate, navicular or reniform basidiospores (10.4–16.7 × 5.2–7.4 µm), occurrence in a deciduous or coniferous forest, occasionally scattered habit on standing timber.

Type

China. Yunnan Province: Shangri-La County, Pudacuo National Park (27°50.61'N, 99°57.03'E, elev. 3800 m), on standing timber, 17 August 2020, Yuan-Hao Ma, Hong-Mei Chai & Wei-Min Chen (Holotype: HKAS 133171!).

Etymology

The epithet “multiramosa” refers to abundant branches of basidiomata.

Description

Basidiomata stipitate, fasciculate, usually geminate, occasionally scattered, gelatinous, 1.5–4.0 cm in height, tough, dendroid and dichotomously branched, cylindrical or flattened, surface smooth, yellowish to orange (5B8, 6A8, 6B7–8), 0.3–0.5 cm in diameter at the upper branching part. Marginal hyphae on sterile surfaces cylindrical, branched, smooth, straight or flexuous, septate, thin-walled, hyaline, 2.0–4.8 µm in diameter. Internal hyphae branched, septate, thin-walled, hyaline, 2.9–10.0 µm in diameter. Hymenium limited to the upper surface of basidomata, amphigenous, composed of basidia and simple cylindrical hyphidia; hyphidia hyaline or pale yellow, smooth, thin-walled. Subhymenial hyphae hyaline, smooth or scabrous, thin- or slightly thick-walled, 2.5–7.3 µm in diameter. Basidia cylindrical to clavate, hyaline or pale yellow, thin-walled, becoming bifurcate when mature, (33.5–)36.5–52.5(–55.0) × (3.5–)3.8–6.1(–6.4) µm, Lm = 45.1 µm, Wm = 4.9 µm, sometimes with many septa. Basidiospores [102/3/3], navicular or reniform, straight or curved, with a small apiculum at the top, thin-walled with thin septa, hyaline to pale yellow, sometimes with oil drops when young and in the germination stage, (6.5–)10.4–16.7(–17.0) × (4.5–)5.2–7.4(–8.8) µm, Lm = 14.3 µm, Wm = 6.3 µm, Q = (1.4–)1.6–2.7(–2.8), Qm = 2.3, 1–5-septate at maturity. Germination with conidia by abnormally developing basidia with lots of septa, by hyphae with septa, or by germ tubes. Clamp connections absent in all tissues of the basidiomata.

Figure 6. 

Basidiomata of Calocera multiramosa A HKAS 133171 (Holotype) B HKAS 133172 C HKAS 133173. Photos by Y.H. Ma. Scale bars: 3 cm.

Habitat and distribution

Geminate, occasionally scattered on standing timber in a deciduous or coniferous forest; known from Yunnan, China.

Figure 7. 

Microscopic structures of Calocera multiramosa in Congo red (Holotype HKAS 133171) A, B basidiospores C germinating basidiospores D probasidia, developing basidia and hyphidia E abnormal developing basidia with septa and geminations. Photos by Y.H. Ma. Scale bars: 10 µm.

Additional specimens examined

China. Yunnan Province: Shangri-La County, Pudacuo National Park, 28 August 2021, Yuan-Hao Ma, Ping Liu & Yong-Chang Zhao (HKAS 133172); Jianchuan County, Laojunshan Town, 26 July 2023, Yuan-Hao Ma & Ping Liu (HKAS 133173).

Figure 8. 

Microscopic structures of Calocera multiramosa in Congo red (Holotype HKAS 133171) A marginal hyphae B internal hyphae C subhymenial hyphae, probasidia, developing basidia, and hyphidia. Photos by Y.H. Ma. Scale bars: 10 µm.

Notes

Calocera multiramosa resembles C. tibetica, C. viscosa and C. mangshanensis in dendrite basidiomata. However, C. multiramosa is distinguished from C. tibetica by larger basidiospores (10.4–16.7 × 5.2–7.4 µm vs. 9.0–13.0 × 5.0–6.0 µm) with different septa (1–5 vs. 3–4) (Fan et al. 2021); C. multiramosa differs from C. viscosa in larger basidia (36.5–52.5 × 3.8–6.1 µm vs. 23–42 × 3–4.5 µm) and basidiospores with different septa (1–5 vs. 0–1) (McNabb 1965; Shirouzu et al. 2009). C. multiramosa can be distinguished from C. mangshanensis by larger (10.4–16.7 × 5.2–7.4 µm vs. 10.0–13.0 × 4.5–5.5 µm), more septate (1–5 vs. 0–1) basidiospores (Liu and Fan 1989). The new species grows on angiosperm and gymnosperm wood, while C. tibetica and C. viscosa only grows on gymnosperm wood and C. mangshanensis only grows on decayed angiosperm wood (McNabb 1965; Liu and Fan 1989; Oberwinkler 2014). C. multiramosa can be distinguished from C. cornea by the size of the basidiomata (1.5–4.0 cm vs. 0.1–0.5 cm high) (Shirouzu et al. 2009), and C. furcata by the mature basidiospores with different septa (1–5 vs. 1–3) (McNabb 1965). The specimen of C. multiramosa, collected from the Laojun Mountain could not be designated as the holotype because of many immature basidiospores. Therefore, the specimen of C. multiramosa collected from a coniferous forest in the Pudacuo National Park was designated as the holotype.

Dacrymyces naematelioides Y.H. Ma, W.M. Chen & Y.C. Zhao, sp. nov.

MycoBank No: 853505
Figs 9, 10, 11

Diagnosis

Dacrymyces naematelioides differs from other species of the genus by stipitate and cerebriform basidiomata, smooth or roughened, simple or branched, septate marginal hyphae (3.0–8.5 µm), smooth or roughened, thin-walled, branched, and septate internal hyphae (2.3–11.0 µm), cylindrical to clavate, smooth or roughened basidia (38.5–79.5 × 6.5–10.6 µm), broadly elliptic-fusiform, 7-septate mature basidiospores (18.5–28.6 × 8.9–13.8 µm), the absence of clamp connections, occurrence in a deciduous forest, and fasciculate, gregarious, or scattered habit on rotten wood.

Figure 9. 

Basidiomata of Dacrymyces naematelioides A–D HKAS 133174 (Holotype). Photos by Y.H. Ma. Scale bars: 3 cm.

Type

China. Yunnan Province: Jianchuan County, Laojunshan Town (26°35.86'N, 99°40.46'E, elev. 3100 m), 21 September 2023, Yuan-Hao Ma, Min Zeng & Wei-min Chen (Holotype: HKAS 133174!).

Figure 10. 

Microscopic structures of Dacrymyces naematelioides (Holotype HKAS 133174) A immature and mature basidiospores in Congo red and KOH solution B probasidia, developing basidia and hyphidia in Congo red. Photos by Y.H. Ma. Scale bars: 10 µm.

Etymology

The epithet “naematelioides” refers to the similarity of the new species in terms of macromorphological features to Naematelia aurantialba.

Description

Basidiomata stipitate, fasciculate and conspicuous, gregarious or scattered, gelatinous when fresh, cerebriform, 2.5–4.5 cm high, surface smooth, orange to light brown (6A8, 6D7–8), occasionally colorless, stipe flat cylindrical, usually with white hairs. Marginal hyphae on sterile surfaces of basidiocarps cylindrical, simple or branched, smooth or roughened, straight or flexuous, septate, thick-walled, hyaline, 3.0–8.5 µm in diameter. Internal hyphae branched, septate, thin-walled, hyaline, smooth or roughened, 2.3–11.0 µm in diameter. Hymenium limited to the upper surface of the basidoma, amphigenous, composed of basidia and simple cylindrical hyphidia; hyphidia hyaline or pale yellow, smooth, thin-walled. Subhymenial hyphae, smooth or roughened, thin- to thick-walled, 2.5–5.3 µm in diameter. Basidia cylindrical to clavate, smooth or roughened, pale yellow, thin-walled, becoming bifurcate, (30.0–)38.5–79.5(–83.5) × (5.5–)6.5–10.6(–11.1) µm, Lm = 60.2 µm, Wm = 8.3 µm. Basidiospores [95/5/1], broadly and elliptic-fusiform, with a small apiculum at the base, thin-walled, pale yellow, with oil drops when young, (16.5–)18.5–28.5(–29.5) × (8.7–)8.9–13.8(–14.6) µm, Lm = 23.9 µm, Wm = 11.0 µm, Q = (1.5–)1.8–2.4(–2.5), Qm = 2.2, usually 7–septate, rarely 3– or 4– septate at maturity. Germination not observed. Clamp connections absent in all tissues of the basidiomata.

Figure 11. 

Microscopic structures of Dacrymyces naematelioides (Holotype HKAS 133174) A marginal hyphae B internal hyphae C subhymenial hyphae. Photos by Y.H. Ma. Scale bars: 10 µm.

Habitat and distribution

Fasciculate, gregarious, or scattered habit on rotten wood, and occurrence in a deciduous forest; known from Yunnan, China.

Notes

Dacrymyces naematelioides resembles D. chrysospermus and D. dictyosporus in shape and size of basidiomata. Microscopically, D. chrysospermus differs from D. naematelioides by narrower basidia (4–6.5 µm vs. 6.5–10.6 µm in width) and smaller basidiospores (16.5–23 × 5–7.5 µm vs. 18.5–28.6 ×8.9–13.8 µm), and D. dictyosporus differs by smooth basidia and thick-walled basidiospores (Martin et al. 1958; McNabb 1973).

Discussion

In this study, we described three new species from Yunnan Province, China, based on morphological evidence and multi-locus phylogenic analyses. The identified morphological features of Campanophyllum include dorsally pseudostipitate pileus, excentric to lateral pseudostipe, and crowded lamellae (Cifuentes et al. 2003; Reschke et al. 2021). The specimen of Campanophyllum sp. (Voucher NEHU.MBSRJ. 38) reported from India formed a sister lineage to C. microsporum with strong supports (100% BS, 1.00 PP). The specimen (Voucher NEHU.MBSRJ. 38) is presumably a new species in the genus Campanophyllum based on the phylogenetic trees (Fig. 1), but it needs to be further confirmed. Meanwhile, our research indicates that more new species of this genus will be discovered in China. However, their habitat is in decline and disappearing.

The species of Calocera in the family Dacrymycetaceae are typically distinguished morphologically based on simple or forked clavarioid basidiocarps (Oberwinkler 2014), but the genus Dacrymyces includes more than 30 species with variable basidiomata including pulvinate, discoid, turbinate, spathulate, flabellate, and cylindrical forms (McNabb 1973; Shirouzu et al. 2009). Several species of Dacrymyces are morphologically close to Calocera by sharing spathulate or cylindrical basidioma, and yet they can be distinguished by some other morphological features, such as the septa of the basidiospores, morphology of the basidia, and morphology of the marginal hyphae. More appropriate genus boundaries and definitions can be obtained by studying detailed morphological and molecular data on more specimens of Dacrymycetaceae. Continuing collection efforts and herbarium searches in unidentified Dacrymycetaceae will certainly uncover more new species (Savchenko et al. 2021).

Phylogenetic analyses, based on two combined loci (ITS, LSU), as well as morphological characteristics, are important for the identification of Calocera and Dacrymyces species. The two newly proposed species formed separate branches on the phylogenetic trees with strong statistical support, and the phylogenies for the genera presented here were found to be similar to those of previous studies (Shirouzu et al. 2013). The results of our study indicated that the specimens of Dacrymyces naematelioides collected from the same locality formed two distinct clades in the phylogenetic analysis (Fig. 2) with strong statistical support (100% BS, 1.00 PP), but there is no marked difference in their morphological characteristics. This suggests that there may be some variation in this species at the molecular level.

The abnormal developing basidia and probasidia with a lot of septa in specimens of Calocera multiramosa were also observed clearly under the microscope, and sometimes they can germinate with microconidia in the basidiomata. This microscopic feature may also be useful in identifying species in the genus Calocera. The surface of the basidia of the Dacrymyces naematelioides is smooth or roughened (Fig. 9). However, the surface features of the basidia did not seem to have been noted in much of the literature, and it is likely that in most species the surface of the basidia is smooth or has only one morphological feature.

Acknowledgments

We would like to express our sincerest gratitude to Dr. Konstanze Bensch for her invaluable advice regarding the naming problem. Furthermore, we would like to express our gratitude to the two reviewers for their meticulous examination of the manuscript and their invaluable recommendations for enhancing the quality of this paper.

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 financially supported by the China Agriculture Research System of MOF and MARA (CARS-20); Innovation Guidance and Scientific and Technological Enterprises Cultiva-tion Plan of Yunnan province (202204BP090018); Scientific Talents and Platform Plan of Yun-nan province (202105AC160086).

Author contributions

Investigation: YHM, WMC, YCZ. Methodology: YHM. Resources: PL, MZ, HMC, YYG. Su-pervision: WMC, YCZ. Writing – original draft: YHM.

Author ORCIDs

Yuan-Hao Ma https://orcid.org/0000-0001-6638-8931

Ping Liu https://orcid.org/0000-0002-1345-9887

Hong-Mei Chai https://orcid.org/0000-0003-4893-4315

Yi-Yun Guo https://orcid.org/0009-0005-3326-9588

Wei-Min Chen https://orcid.org/0000-0001-5595-1514

Yong-Chang Zhao https://orcid.org/0000-0003-1494-4259

Data availability

All of the data that support the findings of this study are available in the main text.

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