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Yu et al. IMA Fungus (2021) 12:19 https://doi.org/10.1186/s43008-021-00071-1 IMA Fungus RESEARCH Open Access Taxonomy and phylogeny of Resinicium sensu lato from Asia-Pacific revealing a new genus and five new species (Hymenochaetales, Basidiomycota) Jia Yu1,2†, Xue-Wei Wang2†, Shi-Liang Liu2†, Shan Shen2,3 and Li-Wei Zhou1,2* ABSTRACT Resinicium, belonging to Hymenochaetales, Agaricomycetes, is a worldwide genus of corticioid wood-inhabiting fungi. To improve the knowledge of species diversity within the Hymenochaetales, two dozen specimens from Asia- Pacific preliminarily identified to be members of Resinicium sensu lato were carefully studied from morphological and phylogenetic perspectives. From these specimens, a new monotypic genus Skvortzoviella, and five new species, viz. Resinicium austroasianum, R. lateastrocystidium, Skvortzovia dabieshanensis, S. qilianensis and Skvortzoviella lenis are described; moreover, a new basal lineage of Resinicium represented by a Vietnam specimen and three Chinese specimens of S. pinicola are identified. The six newly proposed taxa are morphologically compared with related genera and species, while the family positions of Resinicium, Skvortzovia, and Skvortzoviella within the Hymenochaetales are still ambiguous. In addition, the ancestral geographic origin of Resinicium, even though inconclusive, is now thought to be Asia-Pacific instead of tropical America as previously assumed. KEYWORDS: Corticioid fungi, Skvortzovia, Skvortzoviella, Wood-inhabiting fungi, Six new taxa INTRODUCTION at the family level was not fully clarified from the phylo- Resinicium, a worldwide genus of corticioid wood- genetic perspective. inhabiting fungi, was erected for Hydnum bicolor and A total of 22 species have been assigned to Resinicium Corticium furfuraceum with the former as the generic at some stage (Index Fungorum: http://www. type (Parmasto 1968). Although this genus is treated as indexfungorum.org/Names/Names.asp). However, phylo- a member of Rickenellaceae within Hymenochaetales genetic analyses have indicated that Resinicium is not a (He et al. 2019; Olariaga et al. 2020), the corresponding monophyletic genus (Larsson et al. 2006;Nakasone2007). phylogenetic analysis did not have a comprehensive sam- Within Hymenochaetales, R. aculeatum, R. bicolor, R. con- pling throughout this order (Olariaga et al. 2020). Due fertum, R. friabile, R. grandisporum, R. monticola, R. to the ambiguous circumscription of families within mutabile, R. rimulosum, R. saccharicola,andR. tenue were Hymenochaetales, the taxonomic position of Resinicium accepted to be members of Resinicium sensu stricto, whereas R. furfuraceum, R. furfurellum, R. meridionale and R. pinicola were included in the clade of Resinicium * Correspondence: [email protected] sensu lato and now are put in Skvortzovia (Larsson et al. †Jia Yu, Xue-Wei Wang and Shi-Liang Liu contributed equally to this work. 2006;Nakasone2007; Telleria et al. 2008; Gruhn et al. 1 School of life Science, Liaoning University, Shenyang 110036, Liaoning, 2017; Gruhn and Hallenberg 2018). Skvortzovia was ori- China 2State Key Laboratory of Mycology, Institute of Microbiology, Chinese ginally erected as a monotypic genus for Odontia furfur- Academy of Sciences, Beijing 100101, China ella (Hjortstam and Bononi 1987). Besides the above- Full list of author information is available at the end of the article © The Author(s). 2021 Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. Yu et al. IMA Fungus (2021) 12:19 Page 2 of 16 mentioned four species, Phlebia georgica was also com- 94 °C for 40 s, 57.2 °C for 45 s and 72 °C for 1 min, and a bined into Skvortzovia, bringing the number of species in final extension at 72 °C for 10 min; for nLSU region, ini- that genus to five (Gruhn and Hallenberg 2018). tial denaturation at 94 °C for 1 min, followed by 34 cycles Morphologically, Resinicium is characterized by resu- at 94 °C for 30 s, 47.2 °C for 1 min and 72 °C for 1.5 min, pinate, thin, soft basidiomes with smooth to odontioid and a final extension at 72 °C for 10 min. The PCR prod- hymenia, a monomitic hyphal system mainly with clamp ucts were sequenced with the same primers used in PCR connections or with simple septa in few species, the amplification at the Beijing Genomics Institute, Beijing, presence of astrocystidia, and thin-walled, smooth, el- China. All newly generated sequences are deposited in lipsoid to cylindrical basidiospores. Skvortzovia is quite GenBank (https://www.ncbi.nlm.nih.gov/genbank/; similar to Resinicium but differs in the absence of Table 1). astrocystidia. In this study, we focus on Resinicium sensu lato. repre- Phylogenetic analyses sented by specimens from the Asia-Pacific region includ- Besides the newly generated sequences, additional re- ing China, Vietnam, Malaysia, and Australia. Two new lated sequences were also downloaded from GenBank species are described in each of Resinicium and Skvortzo- (Table 1) for inclusion in the phylogenetic analyses. via, while a new monotypic genus typified by a new spe- Firstly, the combined dataset of ITS and nLSU regions cies without a confirmed position at the family level is (1) was used to explore the phylogenetic positions of the introduced. newly studied specimens within Hymenochaetales. All vouchers of Hymenochaetales and Polyporales listed in MATERIALS AND METHODS Table 1, each with both ITS and nLSU sequences avail- Morphological examination able, were included as ingroup taxa, while Auricularia The studied specimens are deposited at the Fungarium, cornea from Auriculariales was selected as an outgroup Institute of Microbiology, Chinese Academy of Sciences taxon (Hibbett et al. 2007). Due to previous phylogenetic (HMAS), Beijing, China. The specimens were observed studies focusing on Resinicium being mainly based solely with Leica M125 (Wetzlar, Germany) and Nikon SMZ on the ITS region, a voucher- and species-abundant ITS 1500 (Tokyo, Japan) stereomicroscopes and an Olympus dataset of Resinicium (2), comprising all vouchers of BX 43 light microscope (Tokyo, Japan) at magnifications Resinicium in Table 1, was used to further differentiate up to 1000×. Special color terms follow Petersen (1996). species identities within this genus. Finally, another Microscopic procedures followed Wang et al. (2020). combined dataset of ITS and nLSU regions (3) was used Specimen sections were mounted in Cotton Blue (CB), to perform a biogeographic analysis of Resinicium. All Melzer’s reagent (IKI), and 5% potassium hydroxide vouchers of Resinicium listed in Table 1, each with both (KOH). All measurements were made from materials in ITS and nLSU sequences available, were included in this CB. When presenting the variation of basidiospore sizes, dataset. No outgroup taxa were selected for datasets 2 5% of the measurements were excluded from each end and 3. of the range and are given in parentheses. Drawings All datasets were aligned using MAFFT 7.110 (Katoh were made with the aid of a drawing tube. The following and Standley 2013) under the G-INS-i option (Katoh abbreviations are used in the descriptions: L = mean ba- et al. 2005). Regarding the combined datasets of the ITS sidiospore length (arithmetic average of all measured ba- and nLSU regions, each region was aligned separately sidiospores), W = mean basidiospore width (arithmetic and then the alignments of the two regions were average of all measured basidiospores), Q = variation in concatenated as a single alignment. The best-fit evolu- the L/W ratios between the studied specimens, and (a/ tionary models of alignments for phylogenetic analyses b) = number of basidiospores (a) measured from given were estimated using jModelTest (Guindon and Gascuel number (b) of specimens. 2003; Posada 2008) under Akaike information criterion. Regarding datasets 1 and 2, Maximum Likelihood Molecular sequencing (ML) and Bayesian Inference (BI) methods were utilized Crude DNA was extracted from dry specimens as tem- for phylogenetic analyses. The ML method was con- plates for subsequent PCR amplification using CTAB ducted using raxmlGUI 1.2 (Silvestro and Michalak rapid plant genome extraction kit-DN14 (Aidlab Bio- 2012; Stamatakis 2006) with calculation of bootstrap technologies, Beijing, China). The primer pairs ITS5/ (BS) replicates under the auto FC option (Pattengale ITS4 (White et al. 1990; Gardes and Bruns 1993) and et al. 2010). The BI method was conducted using LR0R/LR7 (Vilgalys and Hester 1990) were selected for MrBayes 3.2 (Ronquist et al. 2012). Two independent amplifying ITS and nLSU regions, respectively. The PCR runs were employed, and each run had four chains and procedures were as follows: for ITS region, initial de- started from random trees. Trees were sampled every naturation