<?xml version="1.0" encoding="UTF-8" standalone="no"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v2.3 20070202//EN" "journalpublishing.dtd">
<article xml:lang="EN" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" article-type="research-article">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2022.1080290</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Two new species of <italic>Exidia sensu lato</italic> (Auriculariales, Basidiomycota) based on morphology and DNA sequences</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Tohtirjap</surname> <given-names>Ablat</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/2061384/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Hou</surname> <given-names>Shi-Xing</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Rivoire</surname> <given-names>Bernard</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Gates</surname> <given-names>Genevieve</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1185995/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Wu</surname> <given-names>Fang</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x0002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1800487/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Dai</surname> <given-names>Yu-Cheng</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c002"><sup>&#x0002A;</sup></xref>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Institute of Microbiology, School of Ecology and Nature Conservation, Beijing Forestry University</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>Administration of Yuyuantan Park</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>Retired</institution>, <addr-line>Orli&#x000E9;nas</addr-line>, <country>France</country></aff>
<aff id="aff4"><sup>4</sup><institution>Tasmanian Institute of Agriculture</institution>, <addr-line>Hobart, TAS</addr-line>, <country>Australia</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Jes&#x000FA;s Navas-Castillo, Spanish National Research Council (CSIC), Spain</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: R. Greg Thorn, Western University, Canada; &#x00141;ukasz &#x00141;opusiewicz, West Pomeranian University of Technology, Poland</p></fn>
<corresp id="c001">&#x0002A;Correspondence: Fang Wu &#x02709; <email>fangwubjfu2014&#x00040;bjfu.edu.cn</email></corresp>
<corresp id="c002">Yu-Cheng Dai &#x02709; <email>yuchengdai&#x00040;bjfu.edu.cn</email></corresp>
<fn fn-type="other" id="fn001"><p>This article was submitted to Microbe and Virus Interactions with Plants, a section of the journal Frontiers in Microbiology</p></fn></author-notes>
<pub-date pub-type="epub">
<day>14</day>
<month>02</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>13</volume>
<elocation-id>1080290</elocation-id>
<history>
<date date-type="received">
<day>26</day>
<month>10</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>12</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x000A9; 2023 Tohtirjap, Hou, Rivoire, Gates, Wu and Dai.</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Tohtirjap, Hou, Rivoire, Gates, Wu and Dai</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license> </permissions>
<abstract>
<p>In the present study, fourteen <italic>Exidia</italic>-like specimens were collected from China, France, and Australia. Based on morphological characteristics and phylogenetic analyses using the internal transcribed spacer regions (ITS) and the large subunit of nuclear ribosomal RNA gene (nLSU), four species in <italic>Exidia sensu lato</italic>, including <italic>Exidia saccharina</italic> and <italic>Tremellochaete atlantica</italic>, and two new species, <italic>Exidia subsaccharina</italic> and <italic>Tremellochaete australiensis</italic>, were identified. The four species are described and illustrated in detail. <italic>E. saccharina</italic> and <italic>T. atlantica</italic>, two known species from China are reported for the first time. <italic>E. subsaccharina</italic> and <italic>T. australiensis</italic>, two new species from France and Australia, respectively are also described. <italic>E. subsaccharina</italic> is characterized by its reddish brown to vinaceous brown basidiomata, slightly papillate hymenial surface, and narrowly allantoid basidiospores without oil drop measuring 12.5&#x02013;17.5 &#x000D7; 4.2&#x02013;5.5 &#x003BC;m. It differs from the similar species, <italic>E. saccharina</italic>, by distinctly larger basidiospores (12.5&#x02013;17.5 &#x000D7; 4.2&#x02013;5.5 vs. 10&#x02013;14.2 &#x000D7; 3.2&#x02013;4.5 &#x003BC;m). <italic>Tremellochaete australiensis</italic> is characterized by its white to grayish blue basidiomata, obviously and densely papillate hymenial surface, and allantoid basidiospores with oil drop measuring 13.8&#x02013;16.2 &#x000D7; 4.8&#x02013;6.5 &#x003BC;m. It also can be distinguished from the similar species, <italic>T. atlantica</italic> and <italic>T. japonica</italic>, by its distinctly larger basidiospores (13.5&#x02013;17.8 &#x000D7; 4&#x02013;5.2 vs. 10&#x02013;11.8 &#x000D7; 4&#x02013;4.8 &#x003BC;m in <italic>T. atlantica</italic>; 9.4&#x02013;11.8 &#x000D7; 3.5&#x02013;4.2 &#x003BC;m in <italic>T. japonica</italic>).</p></abstract>
<kwd-group>
<kwd>Auriculariaceae</kwd>
<kwd>phylogenetic analysis</kwd>
<kwd>taxonomy</kwd>
<kwd>wood-rotting fungi</kwd>
<kwd>diversity</kwd>
</kwd-group>
<counts>
<fig-count count="6"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="41"/>
<page-count count="12"/>
<word-count count="5624"/>
</counts>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="s1">
<title>Introduction</title>
<p><italic>Exidia</italic> Fr. was proposed by Fries and typified by <italic>Exidia glandulosa</italic> (Bull.) Fr. The genus is characterized by gelatinous bassidiomata, ellipsoid to subglobose, longitudinally cruciate septate, 4-celled basidia, cylindrical to allantoid basidiospores, and the ability to cause white rot in woody plants (Lowy, <xref ref-type="bibr" rid="B9">1971</xref>; Liu, <xref ref-type="bibr" rid="B6">1992</xref>; Roberts, <xref ref-type="bibr" rid="B20">2001</xref>; Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>; Wu et al., <xref ref-type="bibr" rid="B35">2022a</xref>). Because of their morphological similarities, <italic>Exidia sensu lato</italic> traditionally includes three genera <italic>Exidia, Myxarium</italic> Wallr., and <italic>Tremellochaete</italic> Raitv (Roberts, <xref ref-type="bibr" rid="B19">1998</xref>; Wei&#x000DF; and Oberwinkler, <xref ref-type="bibr" rid="B31">2001</xref>; Malysheva, <xref ref-type="bibr" rid="B11">2012</xref>), as confirmed by phylogenetic analyses in recent studies (Malysheva and Spirin, <xref ref-type="bibr" rid="B12">2017</xref>; Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>, <xref ref-type="bibr" rid="B25">2019a</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>). Some species in <italic>Hyaloria</italic> M&#x000F6;ller, <italic>Stypella</italic> M&#x000F6;ller, and Sebacina C. Tul and C. Tul. et al., have waxy, very small, and effused basidiomata different from the usually gelatinous, thick, orbicular basidiomata of <italic>Exidia</italic>, were recently transferred into <italic>Myxarium</italic> based on morphological and phylogenetic analyses (Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>). <italic>Myxarium</italic> phylogenetically forms a monophyletic clade distantly related to <italic>Exidia</italic> and <italic>Tremellochaete</italic> (Spirin et al., <xref ref-type="bibr" rid="B25">2019a</xref>; Stalpers et al., <xref ref-type="bibr" rid="B26">2021</xref>). In addition, <italic>Myxarium</italic> belongs to Hyaloriaceae, whereas <italic>Exidia</italic> and <italic>Tremellochaete</italic> belong to Auriculariaceae, and <italic>Myxarium</italic> can be distinguished from the latter two genera by its distinctly stalked basidia (Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>, <xref ref-type="bibr" rid="B25">2019a</xref>). Therefore, <italic>Exidia sensu lato</italic> is defined here as a group of fungi that includes <italic>Exidia</italic> and <italic>Tremellochaete</italic>.</p>
<p><italic>Exidia</italic> was less studied in the latter part of the 20th century but has received attention more recently (Wei&#x000DF; and Oberwinkler, <xref ref-type="bibr" rid="B31">2001</xref>; Wells et al., <xref ref-type="bibr" rid="B32">2004</xref>; Roberts, <xref ref-type="bibr" rid="B21">2009</xref>) due to its edible species and medicinal values (&#x00141;opusiewicz, <xref ref-type="bibr" rid="B7">2018</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>). One edible species from China, <italic>Exidia yadongensis</italic> described by F. Wu et al., contains rich amino acids and plays a key role in the balance of physiological functions (Chen et al., <xref ref-type="bibr" rid="B2">2019</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>). Recently, <italic>E. reflexa</italic> F. Wu et al., <italic>E. subglandulosa</italic> F. Wu et al., and <italic>E. qinghaiensi</italic> S.R. Wang and Thorn, were described based on multigene phylogenies (Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>; Wang and Thorn, <xref ref-type="bibr" rid="B30">2021</xref>). However, the genus is still polyphyletic in the phylogeny, and species of the genus are scattered in several genera of Auriculariaceae (Yuan et al., <xref ref-type="bibr" rid="B41">2018</xref>; Spirin et al., <xref ref-type="bibr" rid="B25">2019a</xref>,<xref ref-type="bibr" rid="B24">b</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>). <italic>Tremellochaete</italic> was reinstated to accommodate <italic>T. japonica</italic> (Yasuda), Raitv. and <italic>T. nigerrima</italic> (Vi&#x000E9;gas), Spirin and Malysheva (Malysheva and Spirin, <xref ref-type="bibr" rid="B12">2017</xref>), and <italic>T. atlantica</italic> Alvarenga and <italic>T. cerradensis</italic> Alvarenga, and one new combination species, <italic>T. ciliata</italic> (M&#x000F6;ller) Spirin and Alvarenga, were described and proposed in this genus (Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; Phookamsak et al., <xref ref-type="bibr" rid="B16">2019</xref>). In total, there are more than 70 species in <italic>Exidia</italic> and six species in <italic>Tremellochaete</italic> worldwide according to Index Fungorum (<ext-link ext-link-type="uri" xlink:href="http://www.indexfungorum.org">http://www.indexfungorum.org</ext-link>) and MycoBank (<ext-link ext-link-type="uri" xlink:href="https://www.mycobank.org">https://www.mycobank.org</ext-link>), but &#x0003C; 20 species have molecular data (Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>; Wang and Thorn, <xref ref-type="bibr" rid="B30">2021</xref>). <italic>Tremellochaete</italic> was reinstated at the genus level, it is not accepted by some researchers (Wang and Thorn, <xref ref-type="bibr" rid="B30">2021</xref>), and the generic demarcation of <italic>Exidia</italic> and <italic>Tremellochaete</italic> is unclear both in morphology and phylogeny (Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>). Further studies are urgently needed based on more samples and taxa.</p>
<p>In the present study, fourteen <italic>Exidia</italic>-like specimens were collected from China, France, and Australia. After morphological examinations and phylogenetic analyses using the internal transcribed spacer regions (ITS) and the large subunit of the nuclear ribosomal RNA gene (nLSU), four species were identified in <italic>Exidia</italic> and <italic>Tremellochaete</italic>, among which two are new to science, and a detailed description of these species is given in the present study.</p></sec>
<sec sec-type="materials and methods" id="s2">
<title>Materials and methods</title>
<sec>
<title>Morphology</title>
<p>The studied specimens were deposited at the herbarium of the Institute of Microbiology, Beijing Forestry University (BJFC), with color terms following those outlined by Petersen (<xref ref-type="bibr" rid="B15">1996</xref>). Sections mounted in 5% KOH and 2% phloxine B (C<sub>20</sub>H<sub>2</sub>Br<sub>4</sub>C<sub>l4</sub>Na<sub>2</sub>O<sub>5</sub>) were studied at a magnification of up to 1,000 &#x000D7; using a Nikon Eclipse 80i microscope and phase contrast illumination. A Nikon Digital Sight DS-L3 camera was used to photograph microscopic structures. We also used other reagents, including Cotton Blue and Melzer&#x00027;s reagent to observe micromorphology following Wu et al. (<xref ref-type="bibr" rid="B39">2022b</xref>). To show the variation in spore sizes, 5% of measurements were excluded from each end of the range and shown in parentheses. At least thirty basidiospores from each specimen were measured. Stalks were excluded from basidia measurements, and the hilar appendage was excluded from basidiospore measurements. The following abbreviations were used: KOH, potassium hydroxide (5%); L, mean length (arithmetic average of all basidiospores length); W, mean width (arithmetic average of all basidiospores width); Q, L/W ratio for each specimen studied; n (a/b), number of basidiospores (a) measured from a given number of specimens (b).</p>
</sec>
<sec>
<title>DNA extraction, PCR reaction, and sequencing</title>
<p>DNA was extracted from dried specimens using a rapid plant genome extraction kit (Aidlab Biotechnologies Co., Ltd., Beijing, China) and modified following Wu et al. (<xref ref-type="bibr" rid="B36">2021</xref>). The internal transcribed spacer regions (ITS) and the large subunit of the nuclear ribosomal RNA gene (nLSU) were amplified with primer pairs ITS 4 and ITS 5 (White et al., <xref ref-type="bibr" rid="B33">1990</xref>) and LR0R and LR7 (Vilgalys and Hester, <xref ref-type="bibr" rid="B29">1990</xref>), respectively. The PCR (polymerase chain reaction) procedure for ITS was initial denaturation at 95&#x000B0;C for 3 min, followed by 35 cycles at 94&#x000B0;C for 40 s, 58&#x000B0;C for 45 s, and 72&#x000B0;C for 1 min, and a final extension at 72&#x000B0;C for 10 min. The PCR procedure for nLSU was initial denaturation at 94&#x000B0;C for 1 min, followed by 35 cycles at 94&#x000B0;C for 1 min, 48&#x000B0;C for 1 min, and 72&#x000B0;C for 1.5 min, and a final extension at 72&#x000B0;C for 10 min (Wu et al., <xref ref-type="bibr" rid="B34">2018</xref>). The PCR products were purified and sequenced at the BGI (Beijing Genomics Institute, China), with the same primers that are used in the PCR reactions. The nLSU sequences were obtained by splicing bidirectional sequences because LR0R-LR7 is &#x0003E;1,000 bp.</p>
</sec>
<sec>
<title>Phylogenetic analyses</title>
<p>The new sequences generated in this study and reference sequences retrieved from GenBank (<xref ref-type="table" rid="T1">Table 1</xref>) were aligned with MAFFT (version 7; Katoh and Standley, <xref ref-type="bibr" rid="B5">2013</xref>) and then manually adjusted in BioEdit and Mesquite version 3.04 software (Hall, <xref ref-type="bibr" rid="B4">1999</xref>; Maddison and Maddison, <xref ref-type="bibr" rid="B10">2017</xref>). A dataset composed of concatenated ITS&#x0002B;nLSU sequences was used in the phylogenetic analyses using the maximum likelihood (ML), maximum parsimony (MP), and Bayesian inference (BI) methods. <italic>Bourdotia galzinii</italic> (Bres.) Trotter was selected as the outgroup in the phylogenetic analyses because the species was closer to species of Auriculariaceae than others but not closely related to species in <italic>Exidia sensu lato</italic> (Spirin et al., <xref ref-type="bibr" rid="B25">2019a</xref>). Except for the outgroup, sequences from the other eleven Auriculariales genera were added to the phylogenetic analyses because <italic>Exidia</italic> was previously shown to be polyphyletic (Yuan et al., <xref ref-type="bibr" rid="B41">2018</xref>; Spirin et al., <xref ref-type="bibr" rid="B25">2019a</xref>).</p>
<table-wrap position="float" id="T1">
<label>Table 1</label>
<caption><p>Taxa information and GenBank accession numbers used in this study.</p></caption>
<table frame="box" rules="all">
<thead>
<tr>
<th valign="top" align="left" style="background-color:#8f9496"><bold>Species</bold></th>
<th valign="top" align="left" style="background-color:#8f9496"><bold>Sample</bold></th>
<th valign="top" align="left" style="background-color:#8f9496" colspan="2"><bold>GenBank Accession nos</bold>.</th>
<th valign="top" align="left" style="background-color:#8f9496"><bold>Country</bold></th>
</tr>
</thead>
<tbody>
<tr>
<td/>
<td/>
<td valign="top" align="left"><bold>ITS</bold></td>
<td valign="top" align="left"><bold>nLSU</bold></td>
<td/>
</tr> <tr>
<td valign="top" align="left"><italic>Adustochaete nivea</italic></td>
<td valign="top" align="left">RLMA 531</td>
<td valign="top" align="left">MN165954</td>
<td valign="top" align="left">MN165989</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Adustochaete interrupta</italic></td>
<td valign="top" align="left">LR 23435</td>
<td valign="top" align="left">MK391518</td>
<td valign="top" align="left">MK391527</td>
<td valign="top" align="left">Mexico</td>
</tr> <tr>
<td valign="top" align="left"><italic>Adustochaete rava</italic></td>
<td valign="top" align="left">KHL 15526</td>
<td valign="top" align="left">MK391517</td>
<td valign="top" align="left">MK391526</td>
<td valign="top" align="left">Brazil</td>
</tr> <tr>
<td valign="top" align="left"><italic>Amphistereum leveilleanum</italic></td>
<td valign="top" align="left">FP1 06715</td>
<td valign="top" align="left">KX262119</td>
<td valign="top" align="left">KX262168</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Amphistereum schrenkii</italic></td>
<td valign="top" align="left">HHB 8476</td>
<td valign="top" align="left">KX262130</td>
<td valign="top" align="left">KX262178</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Auricularia auricula-judae</italic></td>
<td valign="top" align="left">JT 04</td>
<td valign="top" align="left">KT152099</td>
<td valign="top" align="left">KT152115</td>
<td valign="top" align="left">UK</td>
</tr> <tr>
<td valign="top" align="left"><italic>Auricularia auricula-judae</italic></td>
<td valign="top" align="left">Dai 16353</td>
<td valign="top" align="left">MZ618932</td>
<td valign="top" align="left">MZ669900</td>
<td valign="top" align="left">France</td>
</tr> <tr>
<td valign="top" align="left"><italic>Auricularia cornea</italic></td>
<td valign="top" align="left">Dai 13621</td>
<td valign="top" align="left">MZ618936</td>
<td valign="top" align="left">MZ669905</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Auricularia tibetica</italic></td>
<td valign="top" align="left">Dai 13336</td>
<td valign="top" align="left">MZ618943</td>
<td valign="top" align="left">MZ669915</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Elmerina cladophora</italic></td>
<td valign="top" align="left">Otto Miettinen X1902</td>
<td valign="top" align="left">MG757509</td>
<td valign="top" align="left">MG757509</td>
<td valign="top" align="left">Indonesia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Elmerina efibulata</italic></td>
<td valign="top" align="left">Dai 9322</td>
<td valign="top" align="left">JQ764669</td>
<td valign="top" align="left">JQ764647</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Elmerina sclerodontia</italic></td>
<td valign="top" align="left">Otto Miettinen X3269</td>
<td valign="top" align="left">MG757512</td>
<td valign="top" align="left">MG757512</td>
<td valign="top" align="left">Malaysia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Eichleriella alliciens</italic></td>
<td valign="top" align="left">HHB 7194</td>
<td valign="top" align="left">KX262120</td>
<td valign="top" align="left">KX262169</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Eichleriella flavida</italic></td>
<td valign="top" align="left">LR 49412</td>
<td valign="top" align="left">KX262137</td>
<td valign="top" align="left">KX262185</td>
<td valign="top" align="left">UK</td>
</tr> <tr>
<td valign="top" align="left"><italic>Eichleriella sicca</italic></td>
<td valign="top" align="left">OM 17349</td>
<td valign="top" align="left">KX262143</td>
<td valign="top" align="left">KX262191</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia candida</italic></td>
<td valign="top" align="left">VS 3921</td>
<td valign="top" align="left">KY801867</td>
<td valign="top" align="left">KY801892</td>
<td valign="top" align="left">Russia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia candida</italic></td>
<td valign="top" align="left">VS 8588</td>
<td valign="top" align="left">KY801870</td>
<td valign="top" align="left">KY801895</td>
<td valign="top" align="left">USA</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia candida</italic></td>
<td valign="top" align="left">LE 313211</td>
<td valign="top" align="left">KY801868</td>
<td valign="top" align="left">KY801893</td>
<td valign="top" align="left">Russia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia candida</italic></td>
<td valign="top" align="left">LE 38198</td>
<td valign="top" align="left">KY801871</td>
<td valign="top" align="left">KY801896</td>
<td valign="top" align="left">Russia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia crenata</italic></td>
<td valign="top" align="left">Dai 19464</td>
<td valign="top" align="left">MT663359</td>
<td valign="top" align="left">MT664778</td>
<td valign="top" align="left">Canada</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia crenata</italic></td>
<td valign="top" align="left">Wu 26</td>
<td valign="top" align="left">MT663361</td>
<td valign="top" align="left">MT664780</td>
<td valign="top" align="left">Canada</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia glandulosa</italic></td>
<td valign="top" align="left">MW 355</td>
<td valign="top" align="left">AF291273</td>
<td valign="top" align="left">AF291319</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia glandulosa</italic></td>
<td valign="top" align="left">TUFC 34008</td>
<td valign="top" align="left">AB871761</td>
<td valign="top" align="left">AB871742</td>
<td valign="top" align="left">Japan</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia glandulosa</italic></td>
<td valign="top" align="left">Dai 18024</td>
<td valign="top" align="left">MH213394</td>
<td valign="top" align="left">MH213426</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia glandulosa</italic></td>
<td valign="top" align="left">Wu 265</td>
<td valign="top" align="left">MN850376</td>
<td valign="top" align="left">MN850356</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia pithya</italic></td>
<td valign="top" align="left">MW 313</td>
<td valign="top" align="left">AF291275</td>
<td valign="top" align="left">AF291321</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia qinghaiensis</italic></td>
<td valign="top" align="left">HMAS 156328</td>
<td valign="top" align="left">MW353409</td>
<td valign="top" align="left">MW353409</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia qinghaiensis</italic></td>
<td valign="top" align="left">HMAS 156376</td>
<td valign="top" align="left">MW353408</td>
<td valign="top" align="left">MW353408</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia recisa</italic></td>
<td valign="top" align="left">MW 315</td>
<td valign="top" align="left">AF291276</td>
<td valign="top" align="left">AF291322</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia recisa</italic></td>
<td valign="top" align="left">SL 180317</td>
<td valign="top" align="left">MT663365</td>
<td valign="top" align="left">MT664783</td>
<td valign="top" align="left">Finland</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia reflexa</italic></td>
<td valign="top" align="left">Dai 20833</td>
<td valign="top" align="left">MN850386</td>
<td valign="top" align="left">MN850362</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia reflexa</italic></td>
<td valign="top" align="left">Dai 20861</td>
<td valign="top" align="left">MN850388</td>
<td valign="top" align="left">MN850364</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia reflexa</italic></td>
<td valign="top" align="left">Dai 20874</td>
<td valign="top" align="left">MN850389</td>
<td valign="top" align="left">MN850365</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia repanda</italic></td>
<td valign="top" align="left">LY BR 7046</td>
<td valign="top" align="left">MT663367</td>
<td valign="top" align="left">MT664784</td>
<td valign="top" align="left">France</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia saccharina</italic></td>
<td valign="top" align="left">Roki 88</td>
<td valign="top" align="left">AF291277</td>
<td valign="top" align="left">AF291323</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia saccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 15848</bold></td>
<td valign="top" align="left"><bold>OP605366</bold></td>
<td valign="top" align="left"><bold>OP605350</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia saccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 15890</bold></td>
<td valign="top" align="left"><bold>OP605367</bold></td>
<td valign="top" align="left"><bold>OP605351</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia saccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 21719</bold></td>
<td valign="top" align="left"><bold>OP605368</bold></td>
<td valign="top" align="left"><bold>OP605352</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia saccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 21720</bold></td>
<td valign="top" align="left"><bold>OP605369</bold></td>
<td valign="top" align="left"><bold>OP605353</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia subglandulosa</italic></td>
<td valign="top" align="left">Wu 270</td>
<td valign="top" align="left">MN850381</td>
<td valign="top" align="left">MN850357</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia subglandulosa</italic></td>
<td valign="top" align="left">Wu 272</td>
<td valign="top" align="left">MN850383</td>
<td valign="top" align="left">MN850359</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia subglandulosa</italic></td>
<td valign="top" align="left">Wu 278</td>
<td valign="top" align="left">MN850385</td>
<td valign="top" align="left">MN850361</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia subsaccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 22195</bold></td>
<td valign="top" align="left"><bold>OP605370</bold></td>
<td valign="top" align="left"><bold>OP605354</bold></td>
<td valign="top" align="left"><bold>France</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Exidia subsaccharina</bold></italic></td>
<td valign="top" align="left"><bold>Dai 22187</bold></td>
<td valign="top" align="left"><bold>OP605371</bold></td>
<td valign="top" align="left"><bold>OP605355</bold></td>
<td valign="top" align="left"><bold>France</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia thuretiana</italic></td>
<td valign="top" align="left">Spirin 9999</td>
<td valign="top" align="left">KY801878</td>
<td valign="top" align="left">KY801905</td>
<td valign="top" align="left">Finland</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia thuretiana</italic></td>
<td valign="top" align="left">MW 373</td>
<td valign="top" align="left">AF291278</td>
<td valign="top" align="left">AF291324</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia thuretiana</italic></td>
<td valign="top" align="left">VS 11185</td>
<td valign="top" align="left">KY801889</td>
<td valign="top" align="left">KY801914</td>
<td valign="top" align="left">Norway</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia truncata</italic></td>
<td valign="top" align="left">MW 365</td>
<td valign="top" align="left">AF291279</td>
<td valign="top" align="left">AF291325</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia truncata</italic></td>
<td valign="top" align="left">Dai 21231</td>
<td valign="top" align="left">MT663369</td>
<td valign="top" align="left">MT664785</td>
<td valign="top" align="left">Finland</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia uvapassa</italic></td>
<td valign="top" align="left">TUFC 34007</td>
<td valign="top" align="left">AB871863</td>
<td valign="top" align="left">AB871744</td>
<td valign="top" align="left">Japan</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia uvapassa</italic></td>
<td valign="top" align="left">AFTOL-ID 461</td>
<td valign="top" align="left">DQ241776</td>
<td valign="top" align="left">AY645056</td>
<td valign="top" align="left">Japan</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia yadongensis</italic></td>
<td valign="top" align="left">Dai 17209</td>
<td valign="top" align="left">MT663370</td>
<td valign="top" align="left">MT664786</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia yadongensis</italic></td>
<td valign="top" align="left">Dai 17212</td>
<td valign="top" align="left">MT663373</td>
<td valign="top" align="left">MT664789</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidia yadongensis</italic></td>
<td valign="top" align="left">Dai 17268</td>
<td valign="top" align="left">MT663375</td>
<td valign="top" align="left">MT664791</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidiopsis calcea</italic></td>
<td valign="top" align="left">MW 331</td>
<td valign="top" align="left">AF291280</td>
<td valign="top" align="left">AF291326</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidiopsis effusa</italic></td>
<td valign="top" align="left">OM 19136</td>
<td valign="top" align="left">KX262145</td>
<td valign="top" align="left">KX262193</td>
<td valign="top" align="left">Finland</td>
</tr> <tr>
<td valign="top" align="left"><italic>Exidiopsis grisea</italic></td>
<td valign="top" align="left">RoKi 162</td>
<td valign="top" align="left">AF291281</td>
<td valign="top" align="left">AF291328</td>
<td valign="top" align="left">Germany</td>
</tr> <tr>
<td valign="top" align="left"><italic>Grammatus labyrinthinus</italic></td>
<td valign="top" align="left">Yuan 1600</td>
<td valign="top" align="left">KM379139</td>
<td valign="top" align="left">KM379140</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Grammatus semis</italic></td>
<td valign="top" align="left">OM10618</td>
<td valign="top" align="left">KX262146</td>
<td valign="top" align="left">KX262194</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Heteroradulum adnatum</italic></td>
<td valign="top" align="left">LR 23453</td>
<td valign="top" align="left">KX262116</td>
<td valign="top" align="left">KX262165</td>
<td valign="top" align="left">Mexico</td>
</tr> <tr>
<td valign="top" align="left"><italic>Heteroradulum deglubens</italic></td>
<td valign="top" align="left">LE 38182</td>
<td valign="top" align="left">KX262112</td>
<td valign="top" align="left">KX262162</td>
<td valign="top" align="left">Sweden</td>
</tr> <tr>
<td valign="top" align="left"><italic>Heteroradulum deglubens</italic></td>
<td valign="top" align="left">Solheim 1864</td>
<td valign="top" align="left">KX262133</td>
<td valign="top" align="left">KX262181</td>
<td valign="top" align="left">Norway</td>
</tr> <tr>
<td valign="top" align="left"><italic>Heteroradulum kmetii</italic></td>
<td valign="top" align="left">VS 6466</td>
<td valign="top" align="left">KX262104</td>
<td valign="top" align="left">KX262152</td>
<td valign="top" align="left">Russia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Heteroradulum kmetii</italic></td>
<td valign="top" align="left">He 4915</td>
<td valign="top" align="left">MH178262</td>
<td valign="top" align="left">MH178286</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Proterochaete adusta</italic></td>
<td valign="top" align="left">VS 9021</td>
<td valign="top" align="left">MK391520</td>
<td valign="top" align="left">MK391528</td>
<td valign="top" align="left">Canada</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete atlantica</italic></td>
<td valign="top" align="left">URM 90198</td>
<td valign="top" align="left">MG594382</td>
<td valign="top" align="left">MG594384</td>
<td valign="top" align="left">Brazil</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete atlantica</italic></td>
<td valign="top" align="left">URM 90199</td>
<td valign="top" align="left">MG594381</td>
<td valign="top" align="left">MG594383</td>
<td valign="top" align="left">Brazil</td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete atlantica</bold></italic></td>
<td valign="top" align="left"><bold>Dai 22363</bold></td>
<td valign="top" align="left"><bold>OP605374</bold></td>
<td valign="top" align="left"><bold>OP605358</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete atlantica</bold></italic></td>
<td valign="top" align="left"><bold>Dai 22375</bold></td>
<td valign="top" align="left"><bold>OP605375</bold></td>
<td valign="top" align="left"><bold>OP605359</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete atlantica</bold></italic></td>
<td valign="top" align="left"><bold>Wu 539</bold></td>
<td valign="top" align="left"><bold>OP605373</bold></td>
<td valign="top" align="left"><bold>OP605357</bold></td>
<td valign="top" align="left"><bold>China</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete australiensis</bold></italic></td>
<td valign="top" align="left"><bold>Dai 18601A</bold></td>
<td valign="top" align="left"><bold>OP605376</bold></td>
<td valign="top" align="left"><bold>OP605360</bold></td>
<td valign="top" align="left"><bold>Australia</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete australiensis</bold></italic></td>
<td valign="top" align="left"><bold>Dai 18664</bold></td>
<td valign="top" align="left"><bold>OP605377</bold></td>
<td valign="top" align="left"><bold>OP605361</bold></td>
<td valign="top" align="left"><bold>Australia</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete australiensis</bold></italic></td>
<td valign="top" align="left"><bold>Dai 18704</bold></td>
<td valign="top" align="left"><bold>OP605378</bold></td>
<td valign="top" align="left"><bold>OP605362</bold></td>
<td valign="top" align="left"><bold>Australia</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete australiensis</bold></italic></td>
<td valign="top" align="left"><bold>Dai 18714</bold></td>
<td valign="top" align="left"><bold>OP605379</bold></td>
<td valign="top" align="left"><bold>OP605363</bold></td>
<td valign="top" align="left"><bold>Australia</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic><bold>Tremellochaete australiensis</bold></italic></td>
<td valign="top" align="left"><bold>Dai 18758</bold></td>
<td valign="top" align="left"><bold>OP605380</bold></td>
<td valign="top" align="left"><bold>OP605364</bold></td>
<td valign="top" align="left"><bold>Australia</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete cerradensis</italic></td>
<td valign="top" align="left">URM 90200</td>
<td valign="top" align="left">MK391524</td>
<td valign="top" align="left">MK391530</td>
<td valign="top" align="left">Brazil</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete ciliata</italic></td>
<td valign="top" align="left">SP 467241</td>
<td valign="top" align="left">MK391523</td>
<td valign="top" align="left">MK391529</td>
<td valign="top" align="left">Brazil</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete japonica</italic></td>
<td valign="top" align="left">TAA 42689</td>
<td valign="top" align="left">AF291274</td>
<td valign="top" align="left">AF291320</td>
<td valign="top" align="left">Russia</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete japonica</italic></td>
<td valign="top" align="left">Wu 251</td>
<td valign="top" align="left">MN850378</td>
<td valign="top" align="left">MN850367</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Tremellochaete japonica</italic></td>
<td valign="top" align="left">Wu 254</td>
<td valign="top" align="left">MN850379</td>
<td valign="top" align="left">MN850368</td>
<td valign="top" align="left">China</td>
</tr> <tr>
<td valign="top" align="left"><italic>Bourdotia galzinii</italic> (out group)</td>
<td valign="top" align="left">Otto MiettinenX3067</td>
<td valign="top" align="left">MG757511</td>
<td valign="top" align="left">MG757511</td>
<td valign="top" align="left">Spain</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>New sequences are in bold.</p>
</table-wrap-foot>
</table-wrap>
<p>Maximum likelihood (ML), Bayesian inference (BI), and maximum parsimony (MP) phylogenetic analyses were performed using RAxML (version 8; Stamatakis, <xref ref-type="bibr" rid="B27">2014</xref>), MrBayes (version 3.2.7a; Ronquist et al., <xref ref-type="bibr" rid="B22">2012</xref>), and PAUP (version 4.0b10; Swofford, <xref ref-type="bibr" rid="B28">2002</xref>), respectively, following the study of Wu et al. (<xref ref-type="bibr" rid="B37">2020b</xref>). The optimal substitution models for the combined dataset are determined using the Akaike information criterion (AIC) implemented in MrModeltest 2.3 (Posada and Crandall, <xref ref-type="bibr" rid="B17">1998</xref>; Nylander, <xref ref-type="bibr" rid="B14">2004</xref>) after scoring 24 models of evolution by PAUP (version 4.0b10; Swofford, <xref ref-type="bibr" rid="B28">2002</xref>). The GTR &#x0002B; I &#x0002B; G model was applied in the BI and ML analyses.</p>
<p>Branches that received bootstrap support for maximum likelihood (BS), Bayesian posterior probabilities (BPP), and maximum parsimony (BP) &#x0003E;50% (BS), 0.90 (BPP), and 50% (BP), respectively, are considered to be significantly supported. The phylograms were viewed using FigTree version 1.4.2 (Rambaut, <xref ref-type="bibr" rid="B18">2012</xref>).</p></sec></sec>
<sec sec-type="results" id="s3">
<title>Results</title>
<sec>
<title>Phylogenetic analyses</title>
<p>The combined ITS&#x0002B;nLSU dataset included 81 fungal specimens representing 44 species in the Auriculariales. The dataset had an aligned length of 1,898 characters, including 1,406 constants, 156 parsimony-uninformative characters, and 336 parsimony-informative characters. MP analysis yielded four equally parsimonious trees (tree length = 1,607, consistency index = 0.432, retention index = 0.725, rescaled consistency index = 0.313, and homoplasy index = 0.568). The average standard deviation of split frequencies in BI analysis was 0.005425. The topology of the ML tree with bootstrap values for BP, BS, and BPP was chosen to represent the phylogenetic relationship of species in the Auriculariales since ML, MP, and BI resulted in similar topologies (<xref ref-type="fig" rid="F1">Figure 1</xref>). The phylogeny demonstrated our fourteen <italic>Exidia</italic>-like specimens were clustered into four different lineages with high support, including two new lineages that represented two new species, <italic>E. subsaccharina</italic> (100% BS, 1.00 BPP, and 100% BP; <xref ref-type="fig" rid="F1">Figure 1</xref>) and <italic>Tremellochaete australiensis</italic> (100% BS, 1.00 BPP, and 100% BP; <xref ref-type="fig" rid="F1">Figure 1</xref>). The four Chinese specimens and one German sample (Roki 88) identified as <italic>E. saccharina</italic> by Wei&#x000DF; and Oberwinkler (<xref ref-type="bibr" rid="B31">2001</xref>) were nested in the same lineage with high support in the phylogeny (<xref ref-type="fig" rid="F1">Figure 1</xref>), so these specimens were treated as <italic>E. saccharina</italic>, and this represents the first record of the species in China.</p>
<fig id="F1" position="float">
<label>Figure 1</label>
<caption><p>Maximum likelihood tree illustrating the phylogeny of <italic>Exidia sensu lato</italic> based on the combined ITS&#x0002B;nLSU dataset. Branches are labeled with maximum likelihood bootstrap &#x0003E;50%, Bayesian posterior probabilities &#x0003E;0.90, and maximum parsimony bootstrap &#x0003E;50%, respectively. New species are in blue.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0001.tif"/>
</fig>
</sec>
<sec>
<title>Taxonomy</title>
<p><italic><bold>Exidia saccharina</bold></italic> Fr., Syst. mycol. (Lundae) 2(1): 225 (1822), <xref ref-type="fig" rid="F2">Figures 2A</xref>, <xref ref-type="fig" rid="F3">3</xref>.</p>
<fig id="F2" position="float">
<label>Figure 2</label>
<caption><p>Basidiomata of species in <italic>Exidia</italic> and <italic>Tremellochaete</italic>. <bold>(A)</bold> <italic>Exidia saccharina</italic> (Dai 21719); <bold>(B)</bold> <italic>E. subsaccharina</italic> (Dai 22187 and LY BR 337, holotype); <bold>(C)</bold> <italic>Tremellochaete atlantica</italic> (Dai 22375); <bold>(D)</bold> <italic>T. australiensis</italic> (Dai 18664, holotype). Scale bars: 1 cm.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0002.tif"/>
</fig>
<fig id="F3" position="float">
<label>Figure 3</label>
<caption><p>Microscopic structures of <italic>E. saccharina</italic> (Dai 21720). <bold>(A)</bold> A section of hymenium; <bold>(B)</bold> basidia; <bold>(C)</bold> basidiospores; <bold>(D)</bold> hyphidia; <bold>(E)</bold> hyphae. Scale bars: 10 &#x003BC;m.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0003.tif"/>
</fig>
<p><italic>Basidiomata:</italic> When fresh, the basidiomata are gelatinous, fawn to orange-brown, suborbicular to cerebriform, sessile, usually remaining separate, occasionally coalescing; are up to 10 cm wide and 1.5 cm thick; have free margins; have a hymenial surface that is clearly ridged, with sparse papillae, becoming vinaceous brown when dry; and are absent of mineral inclusions.</p>
<p><italic>Internal features:</italic> Hyphal structure is monomitic; hyphae are clamped (clamps are usually open), usually branched, hyaline, thin-walled, 0.5&#x02013;2.5 &#x003BC;m in diameter, and embedded in a gelatinous matrix. Basidia are longitudinally cruciate septate, 4-celled, subglobose to ovoid, and thin-walled, measuring 13&#x02013;15.5 &#x000D7; 8.5&#x02013;11.8 &#x003BC;m. Hyphidia are simple, thin-walled, and hyaline. Basidiospores are narrowly allantoid, slightly to distinctly curved, hyaline, thin-walled, smooth, usually without oil drop, neither amyloid, dextrinoid, nor cyanophilous, measuring (9.8&#x02013;)10&#x02013;14.2(&#x02212;14.5) &#x000D7; (3&#x02013;)3.2&#x02013;4.5 &#x003BC;m, L = 11.71 &#x003BC;m, W = 3.82 &#x003BC;m, and Q = 2.98&#x02013;3.12 (<italic>n</italic> = 60/2).</p>
<p><italic>Specimens examined</italic>: CHINA. Hebei Province, Weichang County, Saihanba National Forest Park, on fallen trunk of <italic>Larix</italic>, 27.VIII.2020, Dai 21719 (BJFC 035621), and Dai 21720 (BJFC 035622); Xinjiang Autonomous Region, Burjin County, Karnas Nature Reserve, on fallen trunk of <italic>Larix</italic>, 11.IX.2015, Dai 15890 (BJFC 019991); Habahe County, Baihaba River Forest Park, on the fallen trunk of <italic>Larix</italic>, 10.IX.2015, Dai 15848 (BJFC 019949).</p>
<p><italic><bold>Exidia subsaccharina</bold></italic> F. Wu, B. Rivoire, A. Tohtirjap, and Y.C. Dai, sp. nov. <xref ref-type="fig" rid="F2">Figures 2B</xref>, <xref ref-type="fig" rid="F4">4</xref>.</p>
<fig id="F4" position="float">
<label>Figure 4</label>
<caption><p>Microscopic structures of <italic>E. subsaccharina</italic> (Dai 22187 and LY BR 337, holotype). <bold>(A)</bold> A section of hymenium; <bold>(B)</bold> basidia; <bold>(C)</bold> basidiospores; <bold>(D)</bold> hyphidia; <bold>(E)</bold> hyphae. Scale bars: 10 &#x003BC;m.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0004.tif"/>
</fig>
<p><italic>MycoBank</italic>: MB846784.</p>
<p><italic>Holotype</italic>: FRANCE. Chaussan, on dead tree of <italic>Pinus sylvestris</italic>, 10.VIII.2008, Dai 22187 and LY BR 337 (BJFC 036778).</p>
<p><italic>Etymology</italic>: <italic>Subsaccharina</italic> (Latin) refers to the micromorphology being similar to that of <italic>E. saccharina</italic>.</p>
<p><italic>Diagnosis: E. subsaccharina</italic> may be confused with <italic>E. saccharina</italic> when fresh, but <italic>E. saccharina</italic> differs from the species by its slightly smaller basidia (13&#x02013;15.5 &#x000D7; 8.5&#x02013;11.8 &#x003BC;m), usually simple hyphidia, and distinctly smaller basidiospores (10&#x02013;14.2 &#x000D7; 3.2&#x02013;4.5 &#x003BC;m).</p>
<p><italic>Basidiomata:</italic> When fresh, the basidiomata are gelatinous, reddish brown to vinaceous brown, orbicular to suborbicular, sessile, usually remaining coalescing, occasionally separate; are fused together with up to 10 cm in width and 1 cm in thickness; have free margins; have a hymenial surface that is slightly ridged, with papillae, becoming fuscous when dry; and are absent of mineral inclusions.</p>
<p><italic>Internal features:</italic> Hyphal structure is monomitic; hyphae are clamped (clamps are usually open), usually branched, hyaline, thin-walled, 0.5&#x02013;3 &#x003BC;m in diameter, and embedded in a gelatinous matrix. Basidia are longitudinally cruciate septate, 4-celled, subglobose to ovoid, and thin-walled, measuring 13.5&#x02013;19.2 &#x000D7; 9.2&#x02013;14.2 &#x003BC;m. Hyphidia are usually branched, sometimes simple, thin-walled, and hyaline. Basidiospores are narrowly allantoid, slightly to distinctly curved, hyaline, thin-walled, smooth, usually without oil drop, neither amyloid, dextrinoid, nor cyanophilous, measuring (12&#x02013;)12.5&#x02013;17.5(&#x02212;18.5) &#x000D7; (4&#x02013;)4.2&#x02013;5.5(&#x02212;5.8) &#x003BC;m, L = 15.78 &#x003BC;m, W = 4.73 &#x003BC;m, and Q = 3.23&#x02013;3.44 (<italic>n</italic> = 60/2).</p>
<p><italic>Additional specimen examined (paratype)</italic>: FRANCE. Orli&#x000E9;nas, on dead tree of <italic>Pinus sylvestris</italic>, 22.XII.2011, Dai 22195 and LY BR 4290 (BJFC 036786).</p>
<p><italic><bold>Tremellochaete atlantica</bold></italic> Alvarenga, in Phookamsak et al., Fungal Diversity 95: 242 (2019) <xref ref-type="fig" rid="F2">Figures 2C</xref>, <xref ref-type="fig" rid="F5">5</xref>.</p>
<fig id="F5" position="float">
<label>Figure 5</label>
<caption><p>Microscopic structures of <italic>T. atlantica</italic> (Dai 23375). <bold>(A)</bold> A section of hymenium; <bold>(B)</bold> basidia; <bold>(C)</bold> basidiospores; <bold>(D)</bold> hyphidia; <bold>(E)</bold> hyphae. Scale bars: 10 &#x003BC;m.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0005.tif"/>
</fig>
<p><italic>Basidiomata:</italic> When fresh, the basidiomata are gelatinous, white to ash-gray or brownish, suborbicular to slightly cerebriform, sessile, usually remaining coalescing, occasionally separate; are fused, with up to 10 cm width and 1 cm thickness; have free margins; have occasionally ridged hymenial surface, are obviously and densely studded with irregular papillae, becoming dark gray or grayish brown when dry; and are absent of mineral inclusions.</p>
<p><italic>Internal features:</italic> Hyphal structure is monomitic; hyphae are usually simple septate, rarely clamped, branched, hyaline, thin-walled, 0.5&#x02013;2 &#x003BC;m in diameter, and embedded in a gelatinous matrix. Basidia are longitudinally cruciate septate, 4-celled, subglobose to globose, and thin-walled, measuring 10&#x02013;12.8 &#x000D7; 9&#x02013;10.8 &#x003BC;m. Hyphidia are distinctly branched, thin-walled, and hyaline. Basidiospores are allantoid, slightly to distinctly curved, hyaline, thin-walled, smooth, usually with oil drop, neither amyloid, dextrinoid, nor cyanophilous, measuring (9.8&#x02013;)10&#x02013;11.8(&#x02212;12.8) &#x000D7; (3.8&#x02013;)4&#x02013;4.8(&#x02212;5) &#x003BC;m, L = 10.95 &#x003BC;m, W = 4.24 &#x003BC;m, and Q = 2.58 (<italic>n</italic> = 30/1).</p>
<p><italic>Specimens examined</italic>: CHINA. Fujian Province, Yongtai County, Tianmenshan National Forest Park, on fallen angiosperm branch, 5.VI.2021, Dai 22363 (BJFC 036947) and Dai 22375 (BJFC 036959); Yunnan Province, Xishuangbanna, Mengla County, Rainforest Valley Scenic Area, on fallen angiosperm branch, 3.VII.2021, Wu 539 (BJFC 036394).</p>
<p><italic><bold>Tremellochaete australiensis</bold></italic> F. Wu, G.M. Gates, A. Tohtirjap, and Y.C. Dai, sp. nov. <xref ref-type="fig" rid="F2">Figures 2D</xref>, <xref ref-type="fig" rid="F6">6</xref>.</p>
<fig id="F6" position="float">
<label>Figure 6</label>
<caption><p>Microscopic structures of <italic>Tremellochaete australiensis</italic> (Dai 18664, holotype). <bold>(A)</bold> A section of hymenium; <bold>(B)</bold> basidia; <bold>(C)</bold> basidiospores; <bold>(D)</bold> hyphidia; <bold>(E)</bold> hyphae. Scale bars: 10 &#x003BC;m.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-13-1080290-g0006.tif"/>
</fig>
<p><italic>MycoBank</italic>: MB846785.</p>
<p><italic>Holotype</italic>: AUSTRALIA. Melbourne, Dandenong Ranges Botanical Garden, on the dead tree of <italic>Rhododendron</italic>, 12.V.2018, Dai 18664 (BJFC 027132).</p>
<p><italic>Etymology</italic>: <italic>Australiensis</italic> (Latin) refers to the species being found in Australia.</p>
<p><italic>Diagnosis: Tremellochaete australiensis</italic> is morphologically similar to <italic>T. atlantica</italic> and <italic>T. japonica</italic>, but the latter two species have shorter basidia (&#x0003C; 13 &#x003BC;m in length) and basidiospores (&#x0003C; 12 &#x003BC;m in length) and branched hyphidia.</p>
<p><italic>Basidiomata:</italic> When fresh, the basidiomata are gelatinous, white to grayish blue, suborbicular to slightly cerebriform, sessile, usually remaining coalescing, occasionally separate; are fused together with up to 20 cm in width and 0.5&#x02013;1 cm in thickness; have free margins; have a hymenial surface occasionally ridged, is clearly and densely studded with irregular papillae, becoming dark gray to black when dry; and are absent of mineral inclusions.</p>
<p><italic>Internal features:</italic> Hyphal structure is monomitic; hyphae are clamped (clamps are usually open), usually branched, hyaline, thin-walled, 0.5&#x02013;2.5 &#x003BC;m in diameter, and embedded in a gelatinous matrix. Basidia are longitudinally cruciate septate, 4-celled, subglobose to globose, and thin-walled, measuring 13&#x02013;15.8 &#x000D7; 11.5&#x02013;15 &#x003BC;m. Hyphidia are simple, cylindrical, thin-walled, and hyaline. Basidiospores are allantoid, slightly to distinctly curved, hyaline, thin-walled, smooth, usually with oil drop, neither amyloid, dextrinoid, nor cyanophilous, measuring (12.8&#x02013;)13.8&#x02013;16.2(&#x02212;18) &#x000D7; (4.5&#x02013;)4.8&#x02013;6.5 &#x003BC;m, L = 14.94 &#x003BC;m, W = 5.59 &#x003BC;m, and Q = 2.6 (<italic>n</italic> = 30/1).</p>
<p><italic>Additional specimens examined (paratypes)</italic>: AUSTRALIA. Tasmania, Hobart, Mt. Wellington, on rotten wood of <italic>Olearia</italic>, 13.V.2018, Dai 18704 (BJFC 027173) and Dai 18714 (BJFC 027183); Mount Field Forest, close to Mount National Park, on the fallen trunk of <italic>Nothofagus</italic>, 14.V.2018, Dai 18758 (BJFC 027226); Victoria, Yarra Ranges National Park, on rotten wood of <italic>Eucalyptus</italic>, 9.V.2018, Dai 18601A (BJFC 027070).</p></sec></sec>
<sec sec-type="discussion" id="s4">
<title>Discussion</title>
<p><italic>Exidia sensu lato</italic> is a genus of wood-inhabiting fungi that grows on dead branches and logs and is best known in the temperate regions of Europe, America, and Asia (Malysheva, <xref ref-type="bibr" rid="B11">2012</xref>; Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>; Wang and Thorn, <xref ref-type="bibr" rid="B30">2021</xref>). Although nearly eighty taxa were recorded in <italic>Exidia sensu lato</italic>, most species were described in the 20th century (Fries, <xref ref-type="bibr" rid="B3">1822</xref>; Lowy, <xref ref-type="bibr" rid="B8">1964</xref>, <xref ref-type="bibr" rid="B9">1971</xref>). In recent years, four species in <italic>Exidia</italic> and two species in <italic>Tremellochaete</italic> were described based on morphology and phylogenetic analyses (Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; Wu et al., <xref ref-type="bibr" rid="B38">2020a</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>; Wang and Thorn, <xref ref-type="bibr" rid="B30">2021</xref>), which improved knowledge of <italic>Exidia sensu lato</italic> across the world. However, since the demarcation of <italic>Exidia</italic> and <italic>Tremellochaete</italic> is still ambiguous, multilocus analyses based on taxonomically and geographically broad sampling are needed.</p>
<p><italic>Tremellochaete</italic> was accepted by most researchers (Malysheva and Spirin, <xref ref-type="bibr" rid="B12">2017</xref>; Malysheva et al., <xref ref-type="bibr" rid="B13">2018</xref>; Yuan et al., <xref ref-type="bibr" rid="B41">2018</xref>; Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>). However, Wang and Thorn (<xref ref-type="bibr" rid="B30">2021</xref>) rejected <italic>Tremellochaete</italic> because its type species <italic>T. japonica</italic> was closely related to <italic>E. candida</italic> Lloyd in their phylogeny. In our phylogeny (<xref ref-type="fig" rid="F1">Figure 1</xref>), three <italic>Tremellochaete</italic> species, <italic>T. atlantica, T. australiensis</italic>, and <italic>T. japonica</italic>, are also closely related to <italic>E. candida</italic>, but they formed a separate clade with robust support; two other species placed in <italic>Tremellochaete, T. cerradensis</italic> and <italic>T. ciliata</italic>, are distantly related (<xref ref-type="fig" rid="F1">Figure 1</xref>). <italic>Tremellochaete</italic> may be a polyphyletic genus like other genera, e.g., <italic>Exidia</italic> and <italic>Exidiopsis</italic> (Yuan et al., <xref ref-type="bibr" rid="B41">2018</xref>; Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; Spirin et al., <xref ref-type="bibr" rid="B24">2019b</xref>). In addition, <italic>Tremellochaete</italic> can be distinguished from <italic>Exidia</italic> by its clear and dense papillae on the hymenial surface (Malysheva and Spirin, <xref ref-type="bibr" rid="B12">2017</xref>; Alvarenga et al., <xref ref-type="bibr" rid="B1">2019</xref>; <xref ref-type="fig" rid="F2">Figure 2</xref>).</p>
<p><italic>Exidia subsaccharina</italic> is morphologically similar to <italic>E. saccharina</italic> by sharing gelatinous and brownish basidiomata, a slightly papillate hymenial surface, and narrowly allantoid basidiospores usually without oil drop and grows on rotten conifer wood (Spirin et al., <xref ref-type="bibr" rid="B23">2018</xref>), and both species are closely related in the phylogeny (<xref ref-type="fig" rid="F1">Figure 1</xref>). However, <italic>E. subsaccharina</italic> can be distinguished from <italic>E. saccharina</italic> by its slightly larger basidia (13.5&#x02013;19.2 &#x000D7; 9.2&#x02013;14.2 vs. 13&#x02013;15.5 &#x000D7; 8.5&#x02013;11.8 &#x003BC;m), usually branched hyphidia (usually simple in <italic>E. saccharina</italic>), and distinctly bigger basidiospores (12.5&#x02013;17.5 &#x000D7; 4.2&#x02013;5.5 vs. 10&#x02013;14.2 &#x000D7; 3.2&#x02013;4.5 &#x003BC;m), and they form two distinct lineages with robust support (<xref ref-type="fig" rid="F1">Figure 1</xref>). <italic>Exidia pithya</italic> (Alb. and Schwein.) Fr. usually grows on conifer wood too, but it differs from <italic>E. subsaccharina</italic> by its resupinate and black basidiomata and distinctly smaller basidiospores (10&#x02013;13 &#x000D7; 3&#x02013;5 &#x003BC;m; Malysheva, <xref ref-type="bibr" rid="B11">2012</xref>), and it is distantly related to <italic>E. subsaccharina</italic> in the phylogeny (<xref ref-type="fig" rid="F1">Figure 1</xref>).</p>
<p><italic>Tremellochaete australiensis</italic> may be confused with <italic>T. atlantica</italic> and <italic>T. japonica</italic> due to their gelatinous and white to gray basidiomata, densely papillated hymenial surface, and allantoid basidiospores usually with oil drop (Phookamsak et al., <xref ref-type="bibr" rid="B16">2019</xref>; Ye et al., <xref ref-type="bibr" rid="B40">2020</xref>), but it has longer basidia (13&#x02013;15.8 &#x000D7; 11.5&#x02013;15 &#x003BC;m in <italic>T. australiensis</italic>; 10&#x02013;12.8 &#x000D7; 9&#x02013;10.8 &#x003BC;m in <italic>T. atlantica</italic>; 9.4&#x02013;12.4 &#x000D7; 9.1&#x02013;14.2 &#x003BC;m in <italic>T. japonica</italic>) and basidiospores (13.8&#x02013;16.2 &#x000D7; 4.8&#x02013;6.5 &#x003BC;m in <italic>T. australiensis</italic>; 10&#x02013;11.8 &#x000D7; 4&#x02013;4.8 &#x003BC;m in <italic>T. atlantica</italic>; 9.4&#x02013;11.8 &#x000D7; 3.5&#x02013;4.2 &#x003BC;m in <italic>T. japonica</italic>). Furthermore, <italic>T. australiensis</italic> usually has cylindrical hyphidia, but they are distinctly branched in <italic>T. atlantica</italic> and <italic>T. japonica</italic>.</p>
<p><italic>Tremellochaete atlantica</italic> was originally described from Brazil by Phookamsak et al. (<xref ref-type="bibr" rid="B16">2019</xref>), and our Chinese samples and the type of <italic>T. atlantica</italic> share almost the same ITS sequences, with &#x0003C; 2 base pair differences in the ITS region between the Chinese samples and the type of <italic>T. atlantica</italic>. The morphology of the Chinese samples fits the descriptions of <italic>T. atlantica</italic> except for slightly longer basidiospores (10&#x02013;11.8 &#x000D7; 4&#x02013;4.8 vs. 7.75&#x02013;10 &#x000D7; 2&#x02013;5 &#x003BC;m) and usually simple septate hyphae, with Brazillian specimens usually having clamped hyphae) (Phookamsak et al., <xref ref-type="bibr" rid="B16">2019</xref>). These minor differences are considered intraspecific, so we consider this the first report of <italic>T. atlantica</italic> from China.</p></sec>
<sec sec-type="data-availability" id="s5">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found in the article/supplementary material.</p></sec>
<sec sec-type="author-contributions" id="s6">
<title>Author contributions</title>
<p>FW and Y-CD coordinated the project, designed the experimental plan, and acquired funding. AT and FW analyzed the data and prepared the original draft. BR and Y-CD collected the samples from the field. S-XH, GG, BR, and Y-CD reviewed and edited the manuscript. All authors contributed to the study and approved the submitted version.</p></sec>
</body>
<back>
<sec sec-type="funding-information" id="s7">
<title>Funding</title>
<p>This study was supported by the National Natural Science Foundation of China (Project Nos. 32070006 and 32270011), the Tibet Autonomous Region Science and Technology Project (XZ202201ZY0006N), and the Fundamental Research Funds for the Central Universities (No. 2021ZY91).</p>
</sec>
<ack><p>We thank Long-Fei Fan, Zhan-Bo Liu, and Ya-Ping Lian for their guidance on DNA extraction, PCR reaction, sequencing, and illustration.</p>
</ack>
<sec sec-type="COI-statement" id="conf1">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="s8">
<title>Publisher&#x00027;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<ref-list>
<title>References</title>
<ref id="B1">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Alvarenga</surname> <given-names>R. L. M.</given-names></name> <name><surname>Spirin</surname> <given-names>V.</given-names></name> <name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Gibertoni</surname> <given-names>T. B.</given-names></name> <name><surname>Larsson</surname> <given-names>K. H.</given-names></name></person-group> (<year>2019</year>). <article-title>Two new genera and six other novelties in <italic>Heterochaete</italic> sensu lato (Auriculariales, Basidiomycota)</article-title>. <source>Botany</source> <volume>97</volume>, <fpage>439</fpage>&#x02013;<lpage>451</lpage>. <pub-id pub-id-type="doi">10.1139/cjb-2019-0046</pub-id></citation>
</ref>
<ref id="B2">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chen</surname> <given-names>H. Y.</given-names></name> <name><surname>Bao</surname> <given-names>D. P.</given-names></name> <name><surname>Yang</surname> <given-names>R. H.</given-names></name> <name><surname>Wang</surname> <given-names>Y.</given-names></name> <name><surname>Gao</surname> <given-names>Y. Y.</given-names></name> <name><surname>Li</surname> <given-names>Y.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Amino acid profile and protein quality of <italic>Exidia</italic> sp</article-title>. <source>Acta Agric.Nucl. Sin.</source> <volume>33</volume>, <fpage>81</fpage>&#x02013;<lpage>87</lpage>.</citation>
</ref>
<ref id="B3">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Fries</surname> <given-names>E. M.</given-names></name></person-group> (<year>1822</year>). <article-title>Systema</article-title>. <source>Mycologicum</source>. <volume>2</volume>, <fpage>1</fpage>&#x02013;<lpage>274</lpage>.</citation>
</ref>
<ref id="B4">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hall</surname> <given-names>T. A.</given-names></name></person-group> (<year>1999</year>). <article-title>Bioedit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT</article-title>. <source>Nucleic Acids Symp. Ser</source>. <volume>41</volume>, <fpage>95</fpage>&#x02013;<lpage>98</lpage>.</citation>
</ref>
<ref id="B5">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Katoh</surname> <given-names>K.</given-names></name> <name><surname>Standley</surname> <given-names>D. M.</given-names></name></person-group> (<year>2013</year>). <article-title>MAFFT multiple sequence alignment software version 7: improvements in performance and usability</article-title>. <source>Mol. Biol. Evol.</source> <volume>30</volume>, <fpage>772</fpage>&#x02013;<lpage>780</lpage>. <pub-id pub-id-type="doi">10.1093/molbev/mst010</pub-id><pub-id pub-id-type="pmid">23329690</pub-id></citation></ref>
<ref id="B6">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Liu</surname> <given-names>B.</given-names></name></person-group> (<year>1992</year>). <source>Flora of Fungorum Sinicorum, vol. 2 (Tremellales and Dacrymycetales). Beijing: Science Press</source>. p. 93.</citation>
</ref>
<ref id="B7">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>&#x00141;opusiewicz</surname> <given-names>&#x00141;.</given-names></name></person-group> (<year>2018</year>). <article-title>Isolation, characterisation and biological activity of melanin from <italic>Exidia nigricans</italic></article-title>. <source>World Sci. News</source> <volume>91</volume>, <fpage>111</fpage>&#x02013;<lpage>129</lpage>.</citation>
</ref>
<ref id="B8">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lowy</surname> <given-names>B.</given-names></name></person-group> (<year>1964</year>). <article-title>New species of Tremellales from Guatemala</article-title>. <source>J. Elisha Mitchell sci. Soc.</source> <volume>80</volume>, <fpage>65</fpage>&#x02013;<lpage>70</lpage>.</citation>
</ref>
<ref id="B9">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lowy</surname> <given-names>B.</given-names></name></person-group> (<year>1971</year>). <source>Flora Neotropica Monograph 6 (Tremellales). New York: Hafner Publishing Co., Inc</source>. 153.</citation>
</ref>
<ref id="B10">
<citation citation-type="web"><person-group person-group-type="author"><name><surname>Maddison</surname> <given-names>W. P.</given-names></name> <name><surname>Maddison</surname> <given-names>D. R.</given-names></name></person-group> (<year>2017</year>). <source>Mesquite: a Modular System for Evolutionary Analysis, version 3.2</source>. Available online at: <ext-link ext-link-type="uri" xlink:href="http://mesquiteproject.org">http://mesquiteproject.org</ext-link></citation>
</ref>
<ref id="B11">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Malysheva</surname> <given-names>V.</given-names></name></person-group> (<year>2012</year>). <article-title>A revision of the genus <italic>Exidia</italic> (Auriculariales, Basidiomycota) in Russia</article-title>. <source>Mikol. Fitopat.</source> <volume>46</volume>, <fpage>365</fpage>&#x02013;<lpage>376</lpage>.</citation>
</ref>
<ref id="B12">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Spirin</surname> <given-names>V.</given-names></name></person-group> (<year>2017</year>). <article-title>Taxonomy and phylogeny of the Auriculariales (Agaricomycetes, Basidiomycota) with stereoid basidiocarps</article-title>. <source>Fungal Biol.</source> <volume>121</volume>, <fpage>689</fpage>&#x02013;<lpage>715</lpage>. <pub-id pub-id-type="doi">10.1016/j.funbio.2017.05.001</pub-id><pub-id pub-id-type="pmid">28705397</pub-id></citation></ref>
<ref id="B13">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Spirin</surname> <given-names>V.</given-names></name> <name><surname>Miettinen</surname> <given-names>O.</given-names></name> <name><surname>Motato-V&#x000E1;squez</surname> <given-names>V.</given-names></name> <name><surname>Hernawati</surname> <given-names>Seelan, J. S. S.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Revision of <italic>Protohydnum</italic> (Auriculariales, Basidiomycota)</article-title>. <source>Mycol. Prog.</source> <volume>17</volume>, <fpage>805</fpage>&#x02013;<lpage>814</lpage>. <pub-id pub-id-type="doi">10.1007/s11557-018-1393-6</pub-id><pub-id pub-id-type="pmid">35903469</pub-id></citation></ref>
<ref id="B14">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Nylander</surname> <given-names>J. A. A.</given-names></name></person-group> (<year>2004</year>). <italic>MrModeltest v2</italic>. <source>Program Distributed by the Author</source>. <publisher-loc>Uppsala</publisher-loc>: <publisher-name>Uppsala University, Evolutionary Biology Centre</publisher-name>.</citation>
</ref>
<ref id="B15">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Petersen</surname> <given-names>J. H.</given-names></name></person-group> (<year>1996</year>). <article-title>The Danish Mycological Society&#x00027;s colour-chart</article-title>. <source>Foreningen til Svampekundskabens Fremme</source> (Greve), <fpage>1</fpage>&#x02013;<lpage>6</lpage>.</citation>
</ref>
<ref id="B16">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Phookamsak</surname> <given-names>R.</given-names></name> <name><surname>Hyde</surname> <given-names>K. D.</given-names></name> <name><surname>Jeewon</surname> <given-names>R.</given-names></name> <name><surname>Bhat</surname> <given-names>J.</given-names></name> <name><surname>Jones</surname> <given-names>E. B.</given-names></name> <name><surname>Maharachchikumbura</surname> <given-names>S. S.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Fungal diversity notes 929&#x02013;1035: taxonomic and phylogenetic contributions on genera and species of fungi</article-title>. <source>Fungal Divers</source>. <volume>95</volume>, <fpage>1</fpage>&#x02013;<lpage>273</lpage>. <pub-id pub-id-type="doi">10.1007/s13225-019-00421-w</pub-id><pub-id pub-id-type="pmid">34899100</pub-id></citation></ref>
<ref id="B17">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Posada</surname> <given-names>D.</given-names></name> <name><surname>Crandall</surname> <given-names>K. A.</given-names></name></person-group> (<year>1998</year>). <article-title>Modeltest: testing the model of DNA substitution</article-title>. <source>Bioinformatics</source>. <volume>14</volume>, <fpage>817</fpage>&#x02013;<lpage>818</lpage>. <pub-id pub-id-type="doi">10.1093/bioinformatics/14.9.817</pub-id><pub-id pub-id-type="pmid">9918953</pub-id></citation></ref>
<ref id="B18">
<citation citation-type="web"><person-group person-group-type="author"><name><surname>Rambaut</surname> <given-names>A.</given-names></name></person-group> (<year>2012</year>). <source>Molecular Evolution, Phylogenetics and Epidemiology. FigTree ver. 1.4 Software</source>. Available online at: <ext-link ext-link-type="uri" xlink:href="http://tree.bio.ed.ac.uk/software/figtree/">http://tree.bio.ed.ac.uk/software/figtree/</ext-link> (accessed 7 August, 2022).</citation>
</ref>
<ref id="B19">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Roberts</surname> <given-names>P.</given-names></name></person-group> (<year>1998</year>). <article-title>A revision of the genera <italic>Heterochaetella, Myxarium, Protodontia</italic> and <italic>Stypella</italic> (Heterobasidiomycetes)</article-title>. <source>Mycotaxon</source> <volume>69</volume>, <fpage>209</fpage>&#x02013;<lpage>248</lpage>.</citation>
</ref>
<ref id="B20">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Roberts</surname> <given-names>P.</given-names></name></person-group> (<year>2001</year>). <article-title>A key to British <italic>Exidia</italic> species</article-title>. <source>Field Mycol.</source> <volume>2</volume>, <fpage>134</fpage>&#x02013;<lpage>135</lpage>. <pub-id pub-id-type="doi">10.1016/S1468-1641(10)60535-X</pub-id></citation>
</ref>
<ref id="B21">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Roberts</surname> <given-names>P.</given-names></name></person-group> (<year>2009</year>). <article-title><italic>Exidia nigricans</italic>: a new and legitimate name for <italic>Exidia plana</italic></article-title>. <source>Mycotaxon</source> <volume>109</volume>, <fpage>219</fpage>&#x02013;<lpage>220</lpage>. <pub-id pub-id-type="doi">10.5248/109.219</pub-id></citation>
</ref>
<ref id="B22">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ronquist</surname> <given-names>F.</given-names></name> <name><surname>Teslenko</surname> <given-names>M.</given-names></name> <name><surname>Van Der Mark</surname> <given-names>P.</given-names></name> <name><surname>Ayres</surname> <given-names>D. L.</given-names></name> <name><surname>Darling</surname> <given-names>A.</given-names></name> <name><surname>H&#x000F6;hna</surname> <given-names>S.</given-names></name> <etal/></person-group>. (<year>2012</year>). <article-title>MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space</article-title>. <source>Syst. Biol.</source> <volume>61</volume>, <fpage>539</fpage>&#x02013;<lpage>542</lpage>. <pub-id pub-id-type="doi">10.1093/sysbio/sys029</pub-id><pub-id pub-id-type="pmid">22357727</pub-id></citation></ref>
<ref id="B23">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Spirin</surname> <given-names>V.</given-names></name> <name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Larsson</surname> <given-names>K. H.</given-names></name></person-group> (<year>2018</year>). On some forgotten of <italic>Exidia</italic> and <italic>Myxarium</italic> (Auriculariales, Basidiomycota). <italic>Nord. J. Bot</italic>. 36, e01601. <pub-id pub-id-type="doi">10.1111/njb.01601</pub-id></citation>
</ref>
<ref id="B24">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Spirin</surname> <given-names>V.</given-names></name> <name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Miettinen</surname> <given-names>O.</given-names></name> <name><surname>Alvarenga</surname> <given-names>R. L. M.</given-names></name> <name><surname>Gibertoni</surname> <given-names>T. B.</given-names></name> <name><surname>Ryvarden</surname> <given-names>L.</given-names></name> <etal/></person-group>. (<year>2019b</year>). <article-title>On <italic>Protomerulius</italic> and <italic>Heterochaetella</italic> (Auriculariales, Basidiomycota)</article-title>. <source>Mycol. Prog.</source> <volume>18</volume>, <fpage>1079</fpage>&#x02013;<lpage>1099</lpage>. <pub-id pub-id-type="doi">10.1007/s11557-019-01507-0</pub-id><pub-id pub-id-type="pmid">23709572</pub-id></citation></ref>
<ref id="B25">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Spirin</surname> <given-names>V.</given-names></name> <name><surname>Malysheva</surname> <given-names>V.</given-names></name> <name><surname>Roberts</surname> <given-names>P.</given-names></name> <name><surname>Trichies</surname> <given-names>G.</given-names></name> <name><surname>Savchenko</surname> <given-names>A.</given-names></name> <name><surname>Larsson</surname> <given-names>K. H.</given-names></name></person-group> (<year>2019a</year>). <article-title>A convolute diversity of the Auriculariales (Agaricomycetes, Basidiomycota) with sphaeropedunculate basidia</article-title>. <source>Nord. J. Bot</source>. 37, e02394. <pub-id pub-id-type="doi">10.1111/njb.02394</pub-id></citation>
</ref>
<ref id="B26">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Stalpers</surname> <given-names>J.</given-names></name> <name><surname>Redhead</surname> <given-names>S.</given-names></name> <name><surname>May</surname> <given-names>T. W.</given-names></name> <name><surname>Rossman</surname> <given-names>A. Y.</given-names></name> <name><surname>Crouch</surname> <given-names>J. A.</given-names></name> <name><surname>Cubeta</surname> <given-names>M. A.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Competing sexual-asexual generic names in Agaricomycotina (Basidiomycota) with recommendations for use</article-title>. <source>IMA Fungus</source> <volume>12</volume>, <fpage>1</fpage>&#x02013;<lpage>31</lpage>. <pub-id pub-id-type="doi">10.1186/s43008-021-00061-3</pub-id><pub-id pub-id-type="pmid">34380577</pub-id></citation></ref>
<ref id="B27">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Stamatakis</surname> <given-names>A.</given-names></name></person-group> (<year>2014</year>). <article-title>RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies</article-title>. <source>Bioinformatics</source> <volume>30</volume>, <fpage>1312</fpage>&#x02013;<lpage>1313</lpage>. <pub-id pub-id-type="doi">10.1093/bioinformatics/btu033</pub-id><pub-id pub-id-type="pmid">24451623</pub-id></citation></ref>
<ref id="B28">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Swofford</surname> <given-names>D. L.</given-names></name></person-group> (<year>2002</year>). <source>PAUP</source><sup>&#x0002A;</sup><italic>: Phylogenetic Analysis Using Parsimony (</italic><sup>&#x0002A;</sup> <italic>and Other Methods); Version 4.0b10; Sinauer Associates</italic>. Sunderland, MA, USA: Sinauer Associates.</citation>
</ref>
<ref id="B29">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Vilgalys</surname> <given-names>R.</given-names></name> <name><surname>Hester</surname> <given-names>M.</given-names></name></person-group> (<year>1990</year>). <article-title>Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several <italic>Cryptococcus</italic> species</article-title>. <source>J. Bacteriol</source>. <volume>172</volume>, <fpage>4238</fpage>&#x02013;<lpage>4246</lpage>. <pub-id pub-id-type="doi">10.1128/jb.172.8.4238-4246.1990</pub-id><pub-id pub-id-type="pmid">2376561</pub-id></citation></ref>
<ref id="B30">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>S. R.</given-names></name> <name><surname>Thorn</surname> <given-names>R. G.</given-names></name></person-group> (<year>2021</year>). <article-title><italic>Exidia qinghaiensis</italic>, a new species from China</article-title>. <source>Mycoscience</source> <volume>62</volume>, <fpage>212</fpage>&#x02013;<lpage>216</lpage>. <pub-id pub-id-type="doi">10.47371/mycosci.2021.03.002</pub-id></citation>
</ref>
<ref id="B31">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wei&#x000DF;</surname> <given-names>M.</given-names></name> <name><surname>Oberwinkler</surname> <given-names>F.</given-names></name></person-group> (<year>2001</year>). <article-title>Phylogenetic relationships in Auriculariales and related groups&#x02013;hypotheses derived from nuclear ribosomal DNA sequences</article-title>. <source>Mycol. Res.</source> <volume>105</volume>, <fpage>103</fpage>&#x02013;<lpage>415</lpage>. <pub-id pub-id-type="doi">10.1017/S095375620100363X</pub-id></citation>
</ref>
<ref id="B32">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wells</surname> <given-names>K.</given-names></name> <name><surname>Bandoni</surname> <given-names>R. J.</given-names></name> <name><surname>Lim</surname> <given-names>S. R.</given-names></name> <name><surname>Berbee</surname> <given-names>M. L.</given-names></name></person-group> (<year>2004</year>). <article-title>&#x0201C;Observations on some species of <italic>Myxarium</italic> and reconsideration of the Auriculariaceae and Hyaloriaceae (Auriculariales),&#x0201D;</article-title> in <source>Frontiers Basidiomycete Mycol. Eching: IHW-Verlag</source>. eds, Agerer, R., Piepenbring, and M., Blanz, P. p. <fpage>237</fpage>&#x02013;<lpage>248</lpage>.</citation>
</ref>
<ref id="B33">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>White</surname> <given-names>T. J.</given-names></name> <name><surname>Bruns</surname> <given-names>T.</given-names></name> <name><surname>Lee</surname> <given-names>S.</given-names></name> <name><surname>Taylor</surname> <given-names>J.</given-names></name></person-group> (<year>1990</year>). <source>Amplification and Direct Sequencing of Fungal Ribosomal RNA Genes for Phylogenetics</source>. <publisher-loc>PCR protocols: a guide to methods and applications. New York, NY</publisher-loc>: <publisher-name>Academic Press</publisher-name>. p. <fpage>315</fpage>&#x02013;<lpage>322</lpage>.</citation>
</ref>
<ref id="B34">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Chen</surname> <given-names>J. J.</given-names></name> <name><surname>Ji</surname> <given-names>X. H.</given-names></name> <name><surname>Vlas&#x000E1;k</surname> <given-names>J.</given-names></name> <name><surname>Dai</surname> <given-names>Y. C.</given-names></name></person-group> (<year>2018</year>). Phylogeny and diversity of morphologically similar polypore genera <italic>Rigidoporus, Physisporinus, Oxyporus</italic> and <source>Leucophellinus. Mycologia</source> <volume>109</volume>, <fpage>749</fpage>&#x02013;<lpage>765</lpage>. <pub-id pub-id-type="doi">10.1080/00275514.2017.1405215</pub-id><pub-id pub-id-type="pmid">29336678</pub-id></citation></ref>
<ref id="B35">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Man</surname> <given-names>X. W.</given-names></name> <name><surname>Tohtirjap</surname> <given-names>A.</given-names></name> <name><surname>Dai</surname> <given-names>Y. C.</given-names></name></person-group> (<year>2022a</year>). <article-title>A comparison of polypore funga and species composition in forest ecosystems of China, North America, and Europe</article-title>. <source>For. Ecosyst.</source> <volume>9</volume>, <fpage>100051</fpage>. <pub-id pub-id-type="doi">10.1016/j.fecs.2022.100051</pub-id></citation>
</ref>
<ref id="B36">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Tohtirjap</surname> <given-names>A.</given-names></name> <name><surname>Fan</surname> <given-names>L. F.</given-names></name> <name><surname>Zhou</surname> <given-names>L. W.</given-names></name> <name><surname>Alvarenga</surname> <given-names>R. L. M.</given-names></name> <name><surname>Gibertoni</surname> <given-names>T. B.</given-names></name> <etal/></person-group>. (<year>2021</year>). Global diversity and updated phylogeny of <italic>Auricularia</italic> (Auriculariales, Basidiomycota). <italic>J. Fungi</italic> 7, 933. <pub-id pub-id-type="doi">10.3390/jof7110933</pub-id><pub-id pub-id-type="pmid">34829220</pub-id></citation></ref>
<ref id="B37">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Yuan</surname> <given-names>Y.</given-names></name> <name><surname>Chen</surname> <given-names>J. J.</given-names></name> <name><surname>Cui</surname> <given-names>B. K.</given-names></name> <name><surname>Zhou</surname> <given-names>M.</given-names></name> <name><surname>Dai</surname> <given-names>Y. C.</given-names></name></person-group> (<year>2020b</year>). <article-title>Terrestriporiaceae fam. nov., a new family of Russulales (Basidiomycota)</article-title>. <source>Mycosphere</source> <volume>11</volume>, <fpage>2755</fpage>&#x02013;<lpage>2766</lpage>. <pub-id pub-id-type="doi">10.5943/mycosphere/11/1/21</pub-id><pub-id pub-id-type="pmid">33622540</pub-id></citation></ref>
<ref id="B38">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Zhao</surname> <given-names>Q.</given-names></name> <name><surname>Yang</surname> <given-names>Z. L.</given-names></name> <name><surname>Ye</surname> <given-names>S. Y.</given-names></name> <name><surname>Rivoire</surname> <given-names>B.</given-names></name> <name><surname>Dai</surname> <given-names>Y. C.</given-names></name></person-group> (<year>2020a</year>). <article-title><italic>Exidia yadongensis</italic>, a new edible species from East Asia</article-title>. <source>Mycosystema</source> <volume>39</volume>, <fpage>1203</fpage>&#x02013;<lpage>1214</lpage>. <pub-id pub-id-type="doi">10.13346/j.mycosystema.200205</pub-id></citation>
</ref>
<ref id="B39">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Zhou</surname> <given-names>L. W.</given-names></name> <name><surname>Vlas&#x000E1;k</surname> <given-names>J.</given-names></name> <name><surname>Dai</surname> <given-names>Y. C.</given-names></name></person-group> (<year>2022b</year>). <article-title>Global diversity and systematics of Hymenochaetaceae with poroid hymenophore</article-title>. <source>Fungal Diver.</source> <volume>113</volume>, <fpage>1</fpage>&#x02013;<lpage>192</lpage>. <pub-id pub-id-type="doi">10.1007/s13225-021-00496-4</pub-id></citation>
</ref>
<ref id="B40">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ye</surname> <given-names>S. Y.</given-names></name> <name><surname>Zhang</surname> <given-names>Y. B.</given-names></name> <name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Liu</surname> <given-names>H. X.</given-names></name></person-group> (<year>2020</year>). <article-title>Multi-locus phylogeny reveals two new species of <italic>Exidia</italic> (Auriculariales, Basidiomycota) from China</article-title>. <source>Mycol. Prog</source>. <volume>19</volume>, <fpage>859</fpage>&#x02013;<lpage>868</lpage>. <pub-id pub-id-type="doi">10.1007/s11557-020-01601-8</pub-id></citation>
</ref>
<ref id="B41">
<citation citation-type="journal"><person-group person-group-type="author"><name><surname>Yuan</surname> <given-names>H. S.</given-names></name> <name><surname>Lu</surname> <given-names>X.</given-names></name> <name><surname>Decock</surname> <given-names>C.</given-names></name></person-group> (<year>2018</year>). <article-title>Molecular and morphological evidence reveal a new genus and species in auriculariales from tropical China</article-title>. <source>Mycokeys</source> <volume>35</volume>, <fpage>27</fpage>&#x02013;<lpage>39</lpage>. <pub-id pub-id-type="doi">10.3897/mycokeys.35.25271</pub-id><pub-id pub-id-type="pmid">30622400</pub-id></citation></ref>
</ref-list> 
</back>
</article> 