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Research Article
Morphological and molecular data reveal Cerrena caulinicystidiata sp. nov. and Polyporus minutissimus sp. nov. in Polyporales from Asia
expand article infoZi-Wei Zheng‡, Qiu-Yue Zhang‡, Li-Rong Zhang§, Hai-Sheng Yuan|, Fang Wu‡
‡ Beijing Forestry University, Beijing, China
§ Chinese Academy of Environmental Planning, Beijing, China
| Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, China

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

Academic editor: María P. Martín
© 2024 Zi-Wei Zheng, Qiu-Yue Zhang, Li-Rong Zhang, Hai-Sheng Yuan, Fang Wu.
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: Zheng Z-W, Zhang Q-Y, Zhang L-R, Yuan H-S, Wu F (2024) Morphological and molecular data reveal Cerrena caulinicystidiata sp. nov. and Polyporus minutissimus sp. nov. in Polyporales from Asia. MycoKeys 106: 1-21. https://doi.org/10.3897/mycokeys.106.121840
Open Access

Abstract

Two new species of Polyporales, Cerrena caulinicystidiata and Polyporus minutissimus, are illustrated and described on the basis of morphological studies and phylogenetic analyses from southern China and Vietnam. C. caulinicystidiata is characterized by annual, resupinate, sometimes effused-reflexed basidiocarps, greyish orange to brownish orange pore surface, irregular pores (3–8 per mm), a trimitic hyphal system, pyriform to ventricose cystidia, and subglobose basidiospores 3.2–4.5 × 2.8–3.5 µm in size. P. minutissimus is characterized by annual, solitary, fan-shaped with a depressed center or infundibuliform basidiocarps, obvious black stipe, cream to buff yellow pileal surface with glabrous, occasionally zonate and radially aligned stripes, angular pores (6–9 per mm), a dimitic hyphal system, and cylindrical basidiospores, 5–9.2 × 2.2–4 μm. Detailed descriptions and illustrations of the two new species are provided. The differences between the two new species and their morphologically similar and phylogenetically related species are discussed.

Key words

Cerrenaceae, phylogeny, Polyporaceae, taxonomy, wood-decaying fungi

Introduction

The order Polyporales presents a great diversity of basidiocarp types and hymenophore configurations (Binder et al. 2013). It is an important group of fungi, as Polyporales species can cause wood-decay and thus play an essential role in the carbon cycle. In addition, some species of Polyporales may have medicinal properties (Dai 1996, 1999, 2012; Dai et al. 2009). The order has long been the subject of research on taxonomic diversity, distribution patterns, and ecological functions (Hibbett et al. 2014). As of early 2024, more than 1,800 species are recognized in the order (Martinez et al. 2004; Martinez et al. 2009; Kirk et al. 2008; Grigoriev et al. 2013; Zhao et al. 2015; Justo et al. 2017). Due to its great diversity, the order is intensively studied worldwide (Justo et al. 2017).

Cerrena Gray is the type genus of Cerrenaceae within the Polyporales (Ryvarden and Gilbertson 1993; Justo et al. 2017), and it is typified by C. unicolor (Bull.) Murrill. It is widely distributed throughout the world. The genus is characterized by resupinate, effused-reflexed or pileate basidiocarps, irregular hymenophore, dimitic or trimitic hyphal systems, cylindric to ellipsoid basidiospores, and white rot (Ryvarden and Gilbertson 1993; Ryvarden et al. 2022). The genus Cerrena published in 1821 has priority above Trametes Fr., published in 1838, and the two genera were considered as a single taxon in several studies. Thus, a huge number of new combinations in Cerrena. (Cunningham and Cunningham1965; Gilbertson and Ryvarden 1987; Corner 1989; Ryvarden and Melo 2022). However, Ryvarden (1991) recommended to keep Cerrena as a separate genus based on their anatomical characters. According to Index Fungorum (http://www.indexfungorum.org) and Yuan (2014), Cerrena currently comprises around 10 species.

Polyporus P. Micheli ex Adans., the type genus of the Polyporaceae, is a well-known polypore genus (Gilbertson and Ryvarden 1987). Given that Micheli (1729) did not originally select a type species for Polyporus, there is no consensus on the selection of the type. Since Donk (1933) selected P. tuberaster (Jacq. ex Pers.) Fr. as the type species, this lectotype was accepted by most subsequent mycologists (Cunningham and Cunningham 1965; Singer 1986; Niemelä and Kotiranta 1991; Ryvarden 1991; Sotome et al. 2008; Ji et al. 2022). Morphologically, Polyporus is a heterogeneous genus including many species belonging to six morphological groups described by Núñez and Ryvarden (1995), viz, the Polyporus group, the Favolus group (= Favolus Fr.), the Melanopus group (= Melanopus Pat.), the Polyporellus group (= Polyporellus Karst.), the Admirabilis group, and the Dendropolyporus group (= Dendropolyporus (Pouz.) Jülich). Phylogenetically, Polyporus s. str. is known as a polyphyletic genus (Krüger et al. 2006; Sotome et al. 2008, 2011; Ji et al. 2022). Phylogenetic analyses of Polyporus did not conform to the six morphological groups, for which further in-depth study of the group is needed (Sotome et al. 2008; Zhou et al. 2016).

During investigations on wood-decaying polypores from South China and Vietnam, specimens that morphologically fit the definitions of Cerrena and Polyporus were collected. Phylogenetically, these samples formed two distinct lineages within Cerrena and Polyporus, respectively, and they are different from their morphologically similar and phylogenetically related species. Therefore, we describe and illustrate two new species, Cerrena caulinicystidiata sp. nov. and Polyporus minutissimus sp. nov. within the Polyporales on the basis of morphological studies and phylogenetic analyses.

Materials and methods

Morphological studies

The studied specimens are deposited in the Fungarium of Beijing Forestry University (BJFC) and the Institute of Applied Ecology of the Chinese Academy of Sciences (IFP). Macro-morphological descriptions were based on field notes and voucher herbarium specimens. Microscopic measurements and drawings were made from slides prepared from voucher tissues and stained with Cotton Blue and Melzer’s reagent. The following abbreviations were used: KOH = 5% potassium hydroxide; CB = Cotton Blue; CB+ = cyanophilous in Cotton Blue; CB– = acyanophilous in Cotton Blue; IKI = Melzer’s; IKI– = neither amyloid nor dextrinoid in Melzer’s reagent; L = mean basidiospore length (arithmetic average of basidiospores); W = mean basidiospore width (arithmetic average of basidiospores); Q = variation in the L/W ratios between specimens studied; n (a/b) = number of basidiospores (a) measured from the given number of specimens (b). When we present basidiospore size variation, 5% of measurements were excluded from each end of the range. These excluded values are given in parentheses. Special color terms follow Anonymous (1969) and Petersen (1996).

DNA extraction and sequencing

A CTAB rapid plant genome extraction kit (Aidlab Biotechnologies, Co., Ltd., Beijing, China) was used to obtain DNA products from voucher specimens following the manufacturer’s instructions with some modifications (Wu et al. 2020, 2022). The following primer pairs were used to amplify the DNA: ITS5 and ITS4 for the internal transcribed spacer (ITS) region (White et al. 1990) and LR0R and LR7 for the nuclear large subunit (nLSU) rDNA gene (Vilgalys and Hester 1990).

The procedures for DNA extraction and polymerase chain reaction (PCR) used in this study were the same as described by Wu et al. (2022). The PCR products were purified and sequenced by Beijing Genomics Institute (BGI), China. All newly generated sequences in this study were deposited in GenBank (Sayers et al.2024; http://www.ncbi.nlm.nih.gov/genbank/) and listed in Table 1.

Table 1.
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Taxa information and GenBank accession numbers of the sequences used in this study.

Species Specimen No. Country ITS LSU
Cerrena albocinnamomea Miettinen 10511 China OR262168 OR262168
Cerrena albocinnamomea NIBRFG0000102423 South Korea FJ821532 –
Cerrena albocinnamomea Dai 12892 China KC485522 KC485539
Cerrena albocinnamomea KUC20121102-06 South Korea KJ668561 –
Cerrena caulinicystidiata Yuan 12664 Vietnam MT269762 MT259328
Cerrena caulinicystidiata Yuan 12666 Vietnam MT269763 MT259329
Cerrena caulinicystidiata (Cerrena sp. 1) BJ2-11 China KX527879 –
Cerrena caulinicystidiata (Cerrena sp. 1) G1669 China MK247953 –
Cerrena caulinicystidiata (Cerrena sp. 1) Otu0185 China MT908560 –
Cerrena caulinicystidiata * Wu 661 China PP035831 PP035828
Cerrena cystidiata 548/17 Brazil MZ649034 MZ649034
Cerrena gilbertsonii JV 1609/29 Guadeloupe OR262202 –
Cerrena gilbertsonii Vandevender 94-144 Mexico OR262171 OR262171
Cerrena multipileata JV 1407/63 Costa Rica OR262201 OR262201
Cerrena multipileata Ryvarden 43881 Costa Rica OR262155 OR262155
Cerrena multipileata Kout A36 Guatemala OR262203 –
Cerrena sp. 2 F12 China OP022000 –
Cerrena sp. 2 7-SU-3-B-77(M)-B Indonesia KJ654531 –
Cerrena sp. 2 NTOU5117 Taiwan MN592928 –
Cerrena unicolor B2 Antarctica HM589361 –
Cerrena unicolor D.T6.5_2 Argentina MH019790 –
Cerrena unicolor CBS 154.29 Canada MH855029 –
Cerrena unicolor He6082 China OM100740 OM083972
Cerrena unicolor GSM-10 China JQ798288 –
Cerrena unicolor Han 849 China MW467890 –
Cerrena unicolor CU2 Czech FJ821536 –
Cerrena unicolor H:Otto Miettinen 9443 Finland FN907915 FN907915
Cerrena unicolor MUT<ITA_:5063 Italy MK581063 –
Cerrena unicolor FCG-1937 Japan LC415531 –
Cerrena unicolor Pertti Uotila 47558 (H) Kyrgyzstan OR262167 –
Cerrena unicolor Feketic Serbia MW485440 –
Cerrena unicolor KA17-0024 South Korea MN294859 –
Cerrena unicolor 3115 Sweden JN710525 JN710525
Cerrena unicolor CUZFVG179 Turkey MK120293 –
Cerrena unicolor K(M):249944 UK MZ159683 –
Cerrena unicolor FD-299 USA KP135304 KP135209
Cerrena unicolor TASM: YG/PS79 Uzbekistan MT526291 –
Cerrena zonata Gates 2008-4-17 (H) Australia OR262160 OR262160
Cerrena zonata Otto Miettinen 9773 (H) China OR262157 OR262157
Cerrena zonata Otto Miettinen 9889 (H) China OR262158 OR262158
Cerrena zonata Otto Miettinen 13798 (H) Indonesia OR262166 OR262166
Cerrena zonata WS36_1_2_B_As Japan LC631683 –
Cerrena zonata PDD:95790 New Zealand HQ533016 –
Cerrena zonata KA17-0224 South Korea MN294861 –
Cerrena zonata LE-BIN 4492 Vietnam OP985107 –
Datroniella scutellata RLG9584T USA JN165004 JN164792
Datroniella tropica Dai 13147 China KC415181 KC415189
Echinochaete brachypora TFM:F 24996 Japan AB462321 AB462309
Echinochaete russiceps TFM:F 15716 Japan AB462310 AB368065
Echinochaete russiceps TFM:F 24250 Japan AB462313 AB462301
Favolus acervatus Cui 11053 China KU189774 KU189805
Favolus acervatus Dai 10749b China KX548953 KX548979
Favolus gracilisporus Cui 4292 China KX548970 KX548992
Favolus gracilisporus Li 1938 China KX548971 KX548993
Hexagonia glabra Dai 10691 China JX569733 JX569750
Hexagonia tenuis Cui 8468 China JX559277 JX559302
Irpex latemarginatus Dai 8289 China KY131835 –
Lentinus longiporus DAOM:229479 Canada AB478880 LC052217
Lentinus longiporus WD2579 Japan AB478879 LC052218
Lentinus substrictus Wei 1582 China KU189767 KU189798
Lentinus substrictus Wei 1600 China KC572022 KC572059
Microporus affinis Cui 7714 China JX569739 JX569746
Microporus flabelliformis Dai 11574 China JX569740 JX569747
Mycobonia flava CulTENN10256 Costa Rica AY513570 AJ487934
Mycobonia flava TENN59088 Argentina AY513571 AJ487933
Neodatronia gaoligongensis* Cui 8055 China JX559269 JX559286
Neodatronia sinensis* Dai 11921 China JX559272 JX559283
Neofavolus cremeoalbidus Cui 12412 China KX899982 KX900109
Neofavolus cremeoalbidus* TUMH:50009 Japan AB735980 AB735957
Neofavolus mikawai Cui 11152 China KU189773 KU189804
Neofavolus mikawai Dai 12361 China KX548975 KX548997
Physisporinus lineatus JV_1008_18 Costa Rica OM669902 –
Physisporinus lineatus JV_1407_37 Costa Rica OM669903 –
Physisporinus vinctus JV0610_A31B-1 Mexico JQ409460 –
Physisporinus vinctus JV0610_A31B-2 Mexico JQ409461 –
Picipes ailaoshanensis Cui 12585 China KX900068 KX900183
Picipes ailaoshanensis* Cui 12578 China KX900067 KX900182
Picipes americanus JV 0809-104 USA KC572003 KC572042
Picipes americanus* JV 0509-149 USA KC572002 KC572041
Picipes annularius* Cui 10123 China KX900060 KX900176
Picipes atratus Dai 13375 China KX900042 KX900158
Picipes atratus* Cui 11289 China KX900043 KX900159
Picipes auriculatus Yuan 4221 China KX900064 KX900180
Picipes auriculatus* Cui 13616 China KX900063 KX900179
Picipes badius Cui 10853 China KU189780 KU189811
Picipes badius Cui 11136 China KU189781 KU189812
Picipes baishanzuensis Cui 11395 China KU189763 KU189794
Picipes baishanzuensis* Dai 13418 China KU189762 KU189793
Picipes brevistipitatus Cui 11345 China KX900074 KX900188
Picipes brevistipitatus* Cui 13652 China KX900075 KX900189
Picipes cf. dictyopus Cui 11109 China KX900025 KX900145
Picipes cf. dictyopus Cui 11092 China KX900026 KX900146
Picipes conifericola Cui 9950 China KU189783 KU189814
Picipes conifericola* Dai 11114 China JX473244 KC572061
Picipes dictyopus TENN 59385 Belize AF516561 AJ487945
Picipes fraxinicola Dai 2494 China KC572023 KC572062
Picipes fraxinicola Wei 6025 China KC572024 KC572063
Picipes melanopus H 6003449 Finland JQ964422 KC572064
Picipes melanopus MJ 372-93 Czech KC572026 KC572065
Picipes nigromarginatus* Cui 8113 China KX900062 KX900178
Picipes pumilus Cui 5464 China KX851628 KX851682
Picipes pumilus Dai 6705 China KX851630 KX851684
Picipes rhizophilus Dai 11599 China KC572028 KC572067
Picipes rhizophilus Dai 16082 China KX851634 KX851687
Picipes subdictyopus Cui 11220 China KX900057 KX900173
Picipes subdictyopus Cui 12539 China KX900058 KX900174
Picipes submelanopus Dai 13294 China KU189770 KU189801
Picipes submelanopus Dai 13296 China KU189771 KU189802
Picipes subtropicus Li 1928 China KU189758 KU189790
Picipes subtropicus* Cui 2662 China KU189759 KU189791
Picipes subtubaeformis Cui 10793 China KU189753 KU189785
Picipes subtubaeformis* Dai 11870 China KU189752 KU189784
Picipes taibaiensis Dai 5741 China JX489169 KC572071
Picipes taibaiensis* Dai 5746 China KX196783 KX196784
Picipes tibeticus Cui 12225 China KU189756 KU189788
Picipes tibeticus* Cui 12215 China KU189755 KU189787
Picipes tubaeformis Niemela 6855 Finland KC572036 KC572073
Picipes tubaeformis JV 0309-1 USA KC572034 KC572072
Picipes ulleungus Cui 12410 China KX900022 KX900142
Picipes virgatus CulTENN11219 Argentina AF516581 AJ488122
Picipes virgatus CulTENN11406 Argentina AF516582 AJ488122
Picipes wuyishanensis* Dai 7409 China KX900061 KX900177
Podofomes mollis RLG6304sp USA JN165002 JN164791
Podofomes stereoides Holonen Finland KC415179 KC415196
Polyporus auratus* Dai 13665 China KX900056 KX900172
Polyporus austrosinensis Cui 11140 China KX900046 KX900162
Polyporus austrosinensis* Cui 11126 China KX900045 KX900161
Polyporus cuticulatus Cui 8637 China KX851614 KX851668
Polyporus cuticulatus Dai 13141 China KX851613 KX851667
Polyporus guianensis TENN 58404 Venezuela AF516566 AJ487948
Polyporus guianensis TENN 59093 Argentina AF516564 AJ487947
Polyporus hapalopus* Yuan 5809 China KC297219 KC297220
Polyporus hemicapnodes Cui 11259 China KX851625 KX851679
Polyporus hemicapnodes Dai 13403 China KX851627 KX851681
Polyporus lamelliporus Dai 12327 China KX851622 KX851676
Polyporus lamelliporus* Dai 15106 China KX851623 KX851677
Polyporus leprieurii TENN 58579 Costa Rica AF516567 AJ487949
Polyporus mangshanensis* Dai 15151 China KX851796 KX851797
Polyporus minutissimus Wu 970 China PP035829 PP035826
Polyporus minutissimus * Wu 971 China PP035830 PP035827
Polyporus parvovarius Yuan 6639 China KX900049 KX900165
Polyporus parvovarius Dai 13948 China KX900050 KX900166
Polyporus radicatus DAOM198916 Canada AF516584 AJ487955
Polyporus radicatus TENN 58831 USA AF516585 AJ487956
Polyporus sp.1 Cui 11071 China KX851642 KX851695
Polyporus sp.1 Cui 11045 China KX851643 KX851696
Polyporus sp.2 Dai 13585A China KX900055 KX900171
Polyporus squamosus Cui 10394 China KX851635 KX851688
Polyporus squamosus Cui 10595 China KU189778 KU189809
Polyporus subvarius WD2368 Japan AB587643 AB587638
Polyporus subvarius* Yu 2 China AB587632 AB587621
Polyporus tuberaster Dai 11271 China KU189769 KU189800
Polyporus tuberaster Dai 12462 China KU507580 KU507582
Polyporus umbellatus Pen 13513 China KU189772 KU189803
Polyporus varius Cui 12249 China KU507581 KU507583
Polyporus varius Dai 13874 China KU189777 KU189808
Pseudofavolus cucullatus Dai 13584A China KX900071 KX900185
Pseudofavolus cucullatus WD2157 Japan AB587637 AB368114
Trametes conchifer FP106793sp USA JN164924 JN164797
Trametes elegans FP105679sp USA JN164944 JN164799
Trametes polyzona Cui 11040 China KR605824 KR605767

Notes: New sequences are in bold; “–” represents missing data; * represents type specimens

Phylogenetic analysis

Phylogenetic trees of Cerrena and Polyporus were constructed using the two concatenated ITS1-5.8S-ITS2-nLSU sequences dataset, respectively, and phylogenetic analyses were performed with Maximum Likelihood (ML) and Bayesian Inference (BI) methods. New sequences generated in this study and reference sequences retrieved from GenBank (Table 1) were partitioned to ITS1, 5.8S, ITS2, nLSU and then aligned separately using MAFFT v.74 (Katoh et al. 2019; http://mafft.cbrc.jp/alignment/server/) 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). Unreliably aligned sections were removed before the analyses, and efforts were made to manually inspect and improve the alignment. The data matrix was edited in Mesquite v3.70. Irpex latemarginatus (Durieu & Mont.) C.C. Chen & Sheng H. Wu was used as an outgroup in the phylogenetic analysis of Cerrena (Parmasto and Hallenberg 2000). Trametes conchifer (Schwein.) Pilát, T. elegans (Spreng.) Fr. and T. polyzona (Pers.) Justo were selected as outgroups in the phylogenetic analysis of Polyporus (Ji et al. 2022). The final alignments and the retrieved topologies were deposited in TreeBASE (http://www.treebase.org) under accessions 31102, 31103.

RAxML 7.2.8 was used to infer ML trees for both datasets with the GTR+I+G model of site substitution, including estimation of Gamma-distributed rate heterogeneity and a proportion of invariant sites (Stamatakis 2006). The branch support was evaluated with a bootstrapping method of 1,000 replicates (Hillis and Bull 1993).

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. The BI was conducted with MrBayes 3.2.6 in two independent runs, each of which had four chains for 20 million generations and started from random trees (Ronquist et al. 2012). Trees were sampled every 1,000 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.

Phylogenetic trees were visualized using FigTree version 1.4.4 (Rambaut 2018). Branches that received bootstrap support (BS) for ML and BPPs greater than or equal to 75% (ML) and 0.95 (BPP) were considered significantly supported, respectively.

Results

Phylogenetic analyses

In the phylogenetic analysis of Cerrena (Fig. 1), the combined ITS1-5.8S-ITS2-nLSU dataset included sequences from 50 fungal collections representing 11 taxa, and one sample of Irpex latemarginatus was used as an outgroup. ModelFinder proposed models were HKY+F+G4 for ITS1, GTR+F+I+G4 for 5.8s, HKY+F+G4 for ITS2 and GTR+F+I for nLSU, for Bayesian analysis. The BI analysis resulted in an average standard deviation of split frequencies = 0.008865. As both ML and BI trees resulted in similar topologies, only the topology of the ML analysis is presented together with the statistical values of the ML (≥75%) and BPP (≥0.90) algorithms (Fig. 1). The phylogeny inferred from ITS1-5.8S-ITS2-nLSU sequences (Fig. 1) showed that our three newly sequenced samples together with three samples defined as Cerrena sp. 1 by Miettinen et al. (2023) formed an independent lineage with strong support (97/0.98, Fig. 1). The lineage is defined as the new species Cerrena caulinicystidiata.

Figure 1. 

Maximum Likelihood (ML) phylogenetic tree illustrating the phylogeny of Cerrena and related genera in five families based on the combined ITS1-5.8S-ITS2-nLSU dataset. Branches are labeled with maximum likelihood bootstrap values (ML) higher than 75% and Bayesian posterior probabilities above 0.90. The new species is given in bold.

In the phylogenetic analysis of Polyporus (Fig. 2), the combined ITS1-5.8S-ITS2-nLSU dataset included sequences from 113 fungal collections representing 71 species, and three samples of Trametes were used as outgroups. ModelFinder suggested models were GTR+F+I+G4 for ITS1, K2P+I+G4 for 5.8s, K2P+I+G4 for ITS2 and GTR+F+I+G4 for nLSU, for Bayesian analysis. The BI analysis resulted in an average standard deviation of split frequencies = 0.009675. The ML and BI trees were similar in topology, and only the topology of the ML analysis is presented along with the statistical values of the ML (≥75%) and BPP (≥0.90) algorithms (Fig. 2). The phylogeny inferred from the combined ITS1-5.8S-ITS2-nLSU sequences (Fig. 2) revealed that a new lineage with high support (100/1.00,) nests in the squamosus clade in Polyporus, namely Polyporus minutissimus. The new species is closely related to P. hemicapnodes Berk. & Broome and P. parvovarius H. Lee, N.K. Kim & Y.W. Lim.

Figure 2. 

Maximum Likelihood (ML) phylogenetic tree illustrating the phylogeny of Polyporus and related genera based on the combined ITS1-5.8S-ITS2-nLSU dataset. Branches are labeled with maximum likelihood bootstrap values (ML) higher than 75% and Bayesian posterior probabilities (BPPs) more than 0.90. The new species is given in bold.

Taxonomy

Cerrena caulinicystidiata T. Cao, F. Wu & H.S. Yuan, sp. nov.

MycoBank No: 853719
Figs 3, 4

Holotype

China • Zhejiang Province, Hangzhou, Xiaoshan District, Yangjingwu Forest Park; 30°4'1"N, 120°19'35"E; 134 m a.s.l.; 27 Mar. 2023; on fallen angiosperm branch; F. Wu leg., Wu 661 (BJFC040654).

Etymology

Caulinicystidiata (Lat.): Refers to the cystidia with a tapering base.

Description

Basidiocarps. Annual, resupinate, sometimes effused-reflexed, continuous, easily separable, without special odor or taste when fresh, corky when dry, up to 10 cm long, 3 cm wide and 0.5 mm thick. Pore surface greyish orange to brownish orange; pores irregular, 3–8 per mm, partly split up to 2 mm long; dissepiments thin. Sterile margin finely fimbriated. Subiculum very thin, yellowish white, ca. 0.5 mm thick, a very thin brownish red crust present in the bottom next to wood. Tubes concolorous with pore surface, corky, 0.5–1 mm long.

Hyphal structure. Hyphal system trimitic, generative hyphae with clamp connections; skeletal and binding hyphae CB+, IKI–; tissues unchanged in KOH.

Subiculum. Generative hyphae thin- to slightly thick-walled, hyaline, clamped, frequently branched, 2–5 µm in diam; skeletal hyphae dominant, thick-walled to subsolid, unbranched, interwoven, 2.5–6 µm in diam; binding hyphae hyaline, thick-walled to subsolid, tortuose, moderately branched, 1.5–2.5 μm diam. The thin crust made up of subsolid, brownish and strongly agglutinated hyphae.

Figure 3. 

Basidiocarps of Cerrena caulinicystidiata (Holotype, Wu 661).

Tubes. Generative hyphae infrequent, hyaline, thin- to slightly thick-walled, clamped, rarely branched, 2–3 µm diam; skeletal hyphae dominant, hyaline, thick-walled to subsolid, rarely branched, sometimes with septate, interwoven, 2–4 µm in diam; binding hyphae rare. Cystidia clavate to pyriform to ventricose, mostly thin-walled, occasionally thick-walled, smooth, 13–20 × 6–12 µm; encrusted cystidia numerous, clavate, originated from and tightly embedded in trama, 10–25 × 7–15 µm (with encrustation). Basidia short clavate, with four sterigmata and a basal clamp, 8–11 × 4–5 µm, basidioles in shape similar to basidia, but slightly smaller.

Figure 4. 

Microscopic structures of Cerrena caulinicystidiata (Wu 661) a basidiospores b basidia and basidioles c cystidia d encrusted cystidia e hyphae from subiculum f hyphae from trama.

Basidiospores. Basidiospores broadly-ellipsoid to ovoid, hyaline, thin-walled, smooth, CB–, IKI–, (3–)3.2–4.5(–4.8) × (2.5–)2.8–3.5(–3.9) µm, L = 3.94 µm, W = 2.84 µm, Q = 1.38–1.44 (n = 60/2).

Additional specimens examined

(paratypes). Vietnam • Lam dong Province (Lat.), Lac Duong District, Bidoup Nui Ba National Park; 12°11'8"N, 108°40'41"E; 1495 m a.s.l.; 15 Oct. 2017; on fallen angiosperm branch; H.S. Yuan leg., Yuan 12666 (IFP 019379), Yuan 12664 (IFP 019378).

Polyporus minutissimus Q.Y. Zhang, Z.W. Zheng & F. Wu, sp. nov.

MycoBank No: 853720
Figs 5, 6

Holotype

China • Zhejiang Province, Hangzhou, Yuhang District, Luniao Town; 30°25'50"N, 119°42'38"E; 158.47 m a.s.l.; 9 Jun. 2023; on ground of Bamboo forest; F. Wu leg., Wu 971 (BJFC040963, holotype).

Etymology

Minutissimus (Lat.): Referring to the species having tiny basidiocarps.

Description

Basidiocarps. Annual, centrally stipitate, solitary, fleshy to soft leathery when fresh, becoming fragile when dry. Pilei flat with a depressed center or infundibuliform, up to 1.5 cm in diam and 0.5–1 mm thick. Pileal surface cream to buff yellow when dry, glabrous, occasionally zonate and with radially aligned stripes; margin sharp, incurved upon drying. Pore surface cream when dry; pores angular, 6–9 per mm; dissepiments thin, entire. Context buff cream to pale neutral when dry, fragile upon drying, up to 0.5 mm thick. Tubes white to cream when dry, decurrent, up to 0.5 mm thick. Stipe dark violet, glabrous, 0.3–0.5 cm long and 1–2 mm in diam.

Figure 5. 

Basidiocarps of Polyporus minutissimus (Holotype, Wu 971).

Hyphal structure. Hyphal system dimitic; generative hyphae bearing clamp connections, thin-walled, hyaline; skeleton-binding hyphae thick-walled with a wide lumen, with arboriform branches, IKI–, CB+; tissue unchanged in KOH.

Context. Generative hyphae frequent, colorless, thin-walled, 2.5–4 μm in diam; skeleto-binding hyphae dominant, colorless, thick-walled with a wide lumen, moderately branched, strongly interwoven, 2–4.5 μm diam.

Figure 6. 

Microscopic structures of Polyporus minutissimus (Wu970/Wu 971) a basidiospores b basidia and basidioles c hyphae from trama.

Tubes. Generative hyphae frequent, colorless, thin-walled, 2–3 μm in diam; skeleto-binding hyphae dominant, colorless, thick-walled with a wide lumen, moderately branched, interwoven, 1–3 μm in diam. Cystidia and cystidioles absent. Basidia clavate, with four sterigmata and a basal clamp connection, 22–28 × 7–9 μm; basidioles in shape similar to basidia, but slightly smaller.

Stipe. Generative hyphae frequent, colorless, thin-walled, rarely branched, 3–4 μm in diam; skeleto-binding hyphae dominant, colorless, thick-walled with a wide lumen, moderately branched, interwoven, 1.5–4 μm in diam.

Basidiospores. Basidiospores cylindrical to oblong, colorless, thin-walled, smooth, IKI–, CB–, 5–9.2(–10) × (2–)2.2–4(–4.2) μm, L = 7.30 μm, W = 3.23 μm, Q = 2.25–2.27 (n = 60/2).

Additional specimen examined

(paratype). China • Zhejiang Province, Hangzhou, Yuhang District, Luniao Town; 30°25'50"N, 119°42'38"E; 155.11 m a.s.l; on ground of bamboo forest, 9 Jun. 2023; F. Wu leg., Wu 970 (BJFC040962).

Discussion

In this study, two new species of the Polyporales - Cerrena caulinicystidiata and Polyporus minutissimus - are proposed based on morphological and phylogenetic evidence. Our three newly sequenced Cerrena samples together with three samples which were defined as Cerrena sp. 1 by Miettinen et al. (2023) formed an independent well-supported lineage in our phylogeny (Fig. 1). The lineage is proposed as the new species C. caulinicystidiata. Another lineage which is defined as Cerrena sp. 2 in our phylogeny is closely related to C. caulinicystidiata, but we didn’t collect one specimen within the lineage, so the lineage is considered to be Cerrena sp.

Cerrena caulinicystidiata is characterized by its resupinate, sometimes effused-reflexed basidiocarps, greyish orange to brownish orange pore surface, 3–8 per mm pores, and subglobose basidiospores, 3.2–4.5 × 2.8–3.5 µm in size. C. albocinnamomea (Y.C. Dai & Niemelä) H.S. Yuan originally described from Northeast China resembles C. caulinicystidiata by sharing resupinate and easily separable basidiocarps. However, C. albocinnamomea differs from C. caulinicystidiata by its clavate to pyriform cystidia, slightly smaller ellipsoid basidiospores (2.8–3.5 × 2–3 µm vs. 3.2–4.5 × 2.8–3.5 µm), and a dimitic hyphal system (Yuan 2014).

In addition, Rigidoporus vinctus (Berk.) Ryvarden [≡ Physisporinus vinctus (Berk.) Murrill, Wu et al. 2017] resembles C. caulinicystidiata by having resupinate basidiocarps, ochraceous pore surface, ventricose cystidia with a subcylindric appendage, encrusted cystidia, and subglobose basidiospores, but it can be distinguished from the latter species by its smaller pores (6–12 per mm vs. 3–8 per mm) and generative hyphae with simple septa (Ryvarden and Johansen 1980).

The genus Cerrena is widely distributed and has diverse morphological characteristics. Currently, there are 13 records according to Index Fungorum (http://www.indexfungorum.org). However, C. ‘gilbertsonii’ Ryvarden cannot be distinguished from C. cystidiata Rajchenb. & De Meijer by morphological characteristics, and C. ‘multipileata’ (C.L. Leite & J.E. Wright) Miettinen cannot be distinguished from C. zonata (Berk.) H.S. Yuan (Miettinen et al. 2023). Cerrena aurantiopora J.S. Lee & Y.W. Lim is a synonym of C. albocinnamomea (Lee and Lim 2010; Miettinen et al. 2023). Therefore, we provide a Key to 11 undisputed Cerrena species including the new species.

Key to species of the Cerrena

1 Paleotropical or temperate-boreal species 2
– Neotropical (South American) species 9
2 Basidiocarp poroid, occasionally lacerate 3
– Basidiocarp irpicoid 8
3 Pore surface umbrinous to bay or blackish C. subglabrescens
– Pore surface white, light orange to brown 4
4 Pores umber, round, 1–2 per mm C. drummondii
– Pores round to angular, > 3 per mm 5
5 Basidiospores narrowly ellipsoid, 7.5–10 × 2.5–3.5 µm C. caperata
– Basidiospores ellipsoid to broadly-ellipsoid, < 6 µm in length 6
6 Pores angular, dissepiments even or lacerate C. albocinnamomea
– Pores rounded to irregular 7
7 Basidiocarps coriaceous, imbricate C. fulvocinerea
– Basidiocarps resupinate, sometimes effused-reflexed C. caulinicystidiata
8 Pore surface white to cream C. unicolor
– Pore surface first white to pale ochraceous C. zonata
9 Cystidia present C. cystidiata
– Cystidia absent 10
10 Pore surface pale cinnamon to brown C. sclerodepsis
– Pore surface dark brown to almost black C. hydnoides

In the phylogenetic analysis of Polyporus, P. minutissimus was assigned to the squamosus clade with strong support (100/1.00, Fig. 2). The squamosus clade has always been supported by phylogenetic analysis based on the ITS+nLSU or eight-gene datasets, but the species within this clade cannot be combined into a monophyletic genus because they manifest greatly diverse morphology (Ji et al. 2022). Phylogenetically, P. minutissimus is closely related to P. hemicapnodes and P. parvovarius (Fig. 2). P. hemicapnodes was described from Dolosbagey (Sri Lanka). For some time, it was treated as a synonymy of P. leprieurii (Núñez and Ryvarden 1995), which differs from P. minutissimus by its larger basidiocarps (up to 10 cm vs. up to 1.5 cm), cream to tan pore surface, and longer stipe (up to 5 cm vs. up to 0.5 cm, Berkeley and Broome 1873). Polyporus parvovarius has microscopic features similar to P. minutissimus. However, P. parvovarius differs by its smaller basidiocarps (up to 0.35 cm vs. up to 1.5 cm) and light buff to brown pileal surface (Tibpromma et al. 2017).

Macro-morphologically, Polyporus minutissimus has a depressed center or infundibuliform basidiocarps and black stipe, cream to buff yellow pileal surface, and 6–9 per mm pores. Microscopically, it has a dimitic hyphal system, strongly branched skeleton-binding hyphae in both trama and context, and cylindrical basidiospores. Morphologically, P. lamelliporus B.K. Cui, Xing Ji & J.L. Zhou is similar to P. minutissimus by sharing depressed center or infundibuliform basidiocarps, cream to buff yellow pileal surface, and similar-sized basidiospores, but the former differs through its larger basidiocarps (up to 5.2 cm vs. up to 1.5 cm), longer stipe (1–3.5 cm vs. 0.3–0.5 cm), and larger pores (0.5–1 per mm vs. 6–9 per mm, Ji et al. 2022). In addition, Picipes baishanzuensis J.L. Zhou & B.K. Cui, which is similar to P. minutissimus and shares infundibuliform basidiocarps and a black stipe, has also been reported from Baishanzu nature reserve, which is the type producing area of our new species. However, P. baishanzuensis differs from P. minutissimus by its larger basidiocarps (up to 5.5 cm vs. up to 1.5 cm) and smaller basidiospores (6.6–7.9 × 2.5–3.1 µm vs. 5–9.2 × 2.2–4 µm; Zhou et al. 2016).

Polyporus is a very complicated genus with more than 3000 records according to the Index Fungorum. However, studies on Polyporus species in China are gradually being carried out, with some Chinese species having been described in Cui et al. (2019) and Ji et al. (2022). Therefore, we provide a Key to Chinese Polyporus species including the new species.

Key to species of Polyporus in China

1 Stipe absent P. megasporoporus
– Stipe present 2
2 Stipe bearing black cuticle 3
– Stipe white to ochraceous 7
3 Pileal surface covered with dark-brown to reddish-brown squamules P. squamosus
– Pileal surface glabrous 4
4 Pores more than 5 per mm 5
– Pores less than 5 per mm 6
5 Pileal surface concentrically zonate; basidiospores 5.4–7.6 × 2.9–3.8 μm P. hemicapnodes
– Pileal surface azonate; basidiospores 7.5–9 × 2.5–3.3 μm P. varius
6 Pores 3–5 per mm P. mangshanensis
– Pores 1–2 per mm P. subvarius
7 Stipes numerous and branched P. umbellatus
– Stipes usually single and not branched 8
8 Basidiospores < 8 μm in length 9
– Basidiospores > 8 μm in length 11
9 Basidiocarps imbricate P. hapalopus
– Basidiocarps solitary 10
10 Pores angular, 2–3 per mm P. brumalis
– Pores round, 4–5 per mm P. ciliatus
11 Pileal surface with radial stripes 12
– Pileal surface without radial stripes 13
12 Pores 2–5 per mm P. cuticulatus
– Pores 6–9 per mm P. minutissimus
13 Basidiospores usually < 10 μm in length 14
– Basidiospores usually > 10 μm in length P. tuberaster
14 Cystidioles absent 15
– Cystidioles infrequent P. austrosinensis
15 Basidia < 27 μm in length 16
– Basidia > 27 μm in length P. lamelliporus
16 Basidiospores smaller, 6–8.3 × 2.2–3 μm P. arcularius
– Basidiospores larger, 7.7–10 × 3–3.9 μm P. auratus

Polyporales is a large group of Basidiomycota with diverse morphology and phylogeny. There have been over 577 taxonomic proposals in the Polyporales and 2,183 publications with the keyword ‘Polyporales’ over the past decade (Binder et al. 2013; Justo et al. 2017). However, the species in the order are still not sufficiently investigated in Asia, especially in the subtropics and tropics (Li et al. 2016; Hyde 2022). New DNA sequencing techniques have revolutionized fungal taxonomy and diversity, with multi-marker datasets. In the present study, two new polypore species, C. caulinicystidiata and P. minutissimus were found in subtropical regions, which enriches our understanding of the fungal diversity of the Polyporales in Asia.

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 financed by the National Natural Science Foundation of China (Project Nos. 32070006, 32270011, 31701978), the Tibet Autonomous Region Science and Technology Project (XZ202201ZY0006N), and the Fundamental Research Funds for the Central Universities (No. QNTD202307).

Author contributions

Data curation: LRZ. Investigation: HSY, QYZ, FW, ZWZ. Methodology: QYZ, ZWZ. Resources: HSY. Supervision: LRZ, FW. Validation: QYZ, ZWZ. Writing - original draft: ZWZ, QYZ. Writing - review and editing: FW.

Author ORCIDs

Zi-Wei Zheng https://orcid.org/0009-0006-8442-408X

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

Hai-Sheng Yuan https://orcid.org/0000-0001-7056-140X

Fang Wu https://orcid.org/0000-0002-1455-6486

Data availability

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

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1Zi-Wei Zheng, Qiu-Yue Zhang contributed equally to this work.

Supplementary materials

Supplementary material 1 

31102 Treebase

Zi-Wei Zheng, Qiu-Yue Zhang, Li-Rong Zhang, Hai-Sheng Yuan, Fang Wu

Data type: nxs

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.
Download file (46.70 kb)
Supplementary material 2 

31103 Treebase

Zi-Wei Zheng and Qiu-Yue Zhang, Li-Rong Zhang, Hai-Sheng Yuan, Fang Wu

Data type: nxs

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.
Download file (273.03 kb)
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