Mycoscience VOL.62 (2021) 212-216

Short communication qinghaiensis, a new species from China Shurong Wanga, R. Greg Thornb, * a College of Food Science and Engineering, Shanxi Agricultural University, Taigu, 030801, China b Department of Biology, University of Western Ontario, London, N6A 5B7, Canada.

ABSTRACT A novel, wood-inhabiting jelly from China is described as a new species, Exidia qinghaiensis (: ). Phylogenetic analyses were based on sequences of the nuclear ribosomal DNA internal transcribed spacer (nrITS) and large subunit (nrL- SU), RNA polymerase II second largest subunit (RPB2), and translation elongation factor 1-a (Tef1) regions. Sequences of the new taxon formed a sister group to , a species known from Europe and Asia, and distant to sequences of Exidia repanda from Europe. Fruiting bodies are cushion-shaped to irregularly lobed and yellowish brown, basidiospores are hyaline, allantoid (averaging 12.7 × 3.4 μm; average length/width is 3.7), and the host is Betula. The new species also can be distinguished by nrITS, nrLSU, RPB2, and Tef1 sequences. Our multigene phylogeny supports an Exidia including Exidia japonica, type species of Tremellochaete, but defining generic limits in Auriculariaceae will require more extensive taxon sampling.

Keywords: Auriculariaceae, Basidiomycota, one new species, phylogeny,

Article history: Received: 30 December 2020, Revised: 11 March 2021, Accepted: 11 March 2021, Available online 20 May 2021

Exidia Fr. is a genus of wood-inhabiting fungi (Basidiomycota: field collection and morphological and molecular studies of Chi- Auriculariaceae) (Hibbett et al., 2014) growing on dead branches nese Exidia specimens in herbaria. During our work we detected and logs, and best known from the temperate regions. Exidia forms one additional species-level clade based on phylogenetic analyses a sister group to the much better known “wood ears” of the genus of the nuclear ribosomal DNA internal transcribed spacer region (Weiss & Oberwinkler, 2001). The basidia of Exidia are (nrITS) and large subunit (nrLSU), RNA polymerase II second pear-shaped and have longitudinal septa, unlike the tubular and largest subunit (RPB2), and translation elongation factor 1-a (Tef1). transversely septate basidia of Auricularia Bull. (Weiss & Oberwin- Herein we describe this new species based on specimens from Qin- kler, 2001). As in Auricularia, basidiocarps (fruiting bodies) are ghai Province, China. gelatinous and these are diverse in form, ranging from pustular to For light microscopic observations, freehand sections of a rehy- cup-shaped (Wojewoda, 1977; Moore, 1997); indeed, a few species drated portion of specimens were mounted in 2% (w/v) KOH. All of Exidia with rather ear-shaped fruiting bodies have regularly measurements of and hypobasidia were carried out using oil been misidentified as Auricularia (Barber, Thorn, & Voitk, 2011). immersion at 400 × and 1000 × magnification with differential -in The study of fungal diversity plays an important role in its pres- terference microscopy on a Zeiss AxioImager Z1. All dimen- ervation, not only in China but also on a worldwide scale. The my- sions exclude the hilar appendage and are reported as length, width cota of China, including Exidia, still has not been well investigated and Q (length/width), given as the 80th percentile range with out- because of its great geographical extent, and most Exidia species liers in parentheses; the average value of Q is reported as Qavg. Color reported lack supporting molecular data. On the basis of morpho- codes (e.g., 8B5) follow Kornerup and Wanscher (1978). The speci- logical data, a total of 8 species of Exidia, including synonyms and mens we borrowed and examined are from HMAS (Mycological three recently described species, have been reported in recent years Herbarium, Institute of Microbiology, Academia Sinica, Beijing, from China (Wu et al., 2020; Ye, Zhang, Wu, & Liu, 2020). Region- China). ally, five species have been reported from Japan (Aoki & Tubaki, Genomic DNA was extracted from dried materials using the 1986; Imazeki, Otani, & Hongo, 1988; Aoki, 1991), three from Ko- E.Z.N.A. Forensic DNA extraction kit (Omega Bio-Tek, Norcross, rea (Jung, 1993) and ten species from the Russian Far East (Govor- Georgia, USA). PCR amplification was performed with primers ova, 1998; Malysheva, 2012; Malysheva & Spirin, 2017). To add to ITS8F and ITS6R (Dentinger, Margaritescu, & Moncalvo, 2010) or the knowledge of Exidia in China, the first author has undertaken ITS8F/5.8S and 5.8SR/LS1R (Vilgalys & Hester, 1990; Hausner, Reid, & Klassen, 1993) for the ITS region, primers LS1 and LR3 (Vilgalys & Hester, 1990) for the 5’-LSU region, primers b-6F and * Corresponding author. Department of Biology, University of Western Ontario, London, N6A 5B7, Canada. b-7.1R (Matheny, 2005) or b-6F/f-7cR and b-6.9F/b-7.1R (Raja, E-mail address: [email protected] (G. Thorn). Miller, Pearce, & Oberlies, 2017) for the RPB2 region, and ef1-983-F

This is an open-access paper distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivative 4.0 international license (CC BY-NC-ND 4.0: https://creativecommons.org/licenses/by-nc-nd/4.0/). doi:org/10.47371/mycosci.2021.03.002 ― 212 ― S. Wang, R.G. Thorn / Mycoscience VOL.62 (2021) 212-216 and ef1-1567-R (Rehner & Buckley, 2005) for Tef1. Successfully da [syn. Tremellochaete japonica (Yasuda) Raitv.], was placed with amplified products were cleaned using an EZ-10 Spin Column PCR strong support in the genus Exidia, as the sister to Exidia candida Products Purification Kit (BioBasic Canada, Markham, Ontario) Lloyd. However, a more inclusive taxon sample will be required to and sequenced using dye-terminator sequencing at Robarts Re- define generic limits in the Auriculariaceae. search Institute (London, Canada) or Sangon Biotech Co., Ltd. (Shanghai, China). Following DNA sequencing, chromatograms of Taxonomy partial sequences were cleaned and assembled using SeqEd v1.0.3 (Applied Biosystems, Foster City, California, USA). Newly acquired Exidia qinghaiensis S.R. Wang & Thorn, sp. nov. Figs. 2, 3. sequences have been deposited in GenBank as MW353408– MycoBank no.: MB 838343. MW353409 and MW358923–MW358926. Few Exidia sequences available from GenBank contained all of Diagnosis: Fruiting bodies are cushion-shaped to irregularly the chosen gene regions but, after preliminary analyses based sole- lobed and adpressed, becoming confluent, yellowish brown (drying ly on the ITS region that found no matches to our putative new fuscous), with paler flesh, basidiospores are hyaline, allantoid, av- species (data not shown), we chose to focus on a small dataset that eraging 12.7 × 3.4 μm, with Qavg = 3.7, and the host is Betula. Basid- had coverage of as many of these regions as possible (Table 1). For iospores of Exidia saccharina Fr., on conifers, are slightly wider (Bull.) Fr. (HHB 12029) and the outgroup Auric- (10–14 × 3.5–4.0 μm), and those of E. thuretiana, also on angio- ularia heimuer F. Wu, B.K. Cui & Y.C. Dai (Dai 13782), draft ge- sperms, are both longer and broader (13–19 × 4.5–6.0 μm), but nome sequences are available and were queried using BLASTn to similar in shape to those of E. qinghaiensis. These species can be obtain sequences of RPB2 or Tef1. Sequences of the different gene distinguished by their nrITS, nrLSU, RPB2, and Tef1sequences. regions were separately aligned using MAFFT v7 online (Katoh & Holotype: CHINA, Qinghai Province, Menyuan County, Xianmi Standley, 2013) with the G-INS-i strategy and “leave gappy regions” wood farm, approx. 37°17’ N, 101°57’ E, 2,850 m above sea level option invoked, then the rough ends of alignments trimmed using (a.s.l.), on a fallen branch of Betula, 19 Oct 2004, leg. Zhang Xiao- MEGA X (Kumar, Stecher, & Tamura, 2016) before concatenating qing, HMAS 156328 (Mycological Herbarium, Institute of Microbi- to yield a combined matrix of 2,572 aligned bases. Prior to concate- ology, Academia Sinica). nation, evolutionary models were compared using MEGA X, and Gene sequences ex-holotype: MW353409 (nrITS, nrLSU), since the GTR+G+I model received the best ln(L) score in all cases, MW358924 (RPB2) and MW358926 (Tef1). this model was used in maximum likelihood (ML) analyses imple- Etymology: qinghaiensis (Latin), referring to Qinghai Province, mented in MEGA X for the combined dataset. Bayesian analyses where the holotype was collected. were conducted in MrBayes 3.2.6 (MB; Ronquist et al., 2012) with 5 Basidiomata gelatinous, cushion-shaped to irregularly lobed, 000 000 generations, 4 chains, and a burn-in of 25% (when the av- adpressed, orbicular, typically growing separately and adhering to erage standard deviation of split frequencies between chains had the substrate, sometimes fusing together and becoming confluent, stabilized below 0.001). Node support was determined as posterior up to 3 cm in widest dimension and 0.5 cm thick; when dried uni- probabilities in MrBayes, and as bootstrap support in ML analyses formly greyish or fuscous brown (7F3) on both hymenial and abhy- using 100 replicates. The alignments and trees have been deposited menial surfaces, rehydrating to yellowish brown or caramel (5DE7 to TreeBase (http://www.treebase.org) as S27878. to 6E7), with paler flesh; the hymenial, downward-facing surface Phylogenetic analyses of the combined nrITS, nrLSU, RPB2 and smooth, glabrous and a little shining, becoming slightly eroded in Tef1 data (Fig. 1) supports the segregation of Exidia qinghaiensis as age; abymenial surface slightly granular, usually without wrinkles a sister species to Exidia thuretiana (Lév.) Fr. A sample represent- and alveolate-venose. Flesh thin and pliant-gelatinous; odor and ing the type species of Tremellochaete Raitv., Exidia japonica Yasu- taste not noted.

Table 1. Sequences of Auricularia and Exidia used in phylogenetic analyses. New sequences derived for this study are in bold font.

Name Voucher Country ITS LSU RPB2 Tef1 A. americana P. B. Matheny 2295 USA: WA DQ200918 AY634277 DQ366278 DQ408143 A. heimuer Y. C. Dai 13782 China MG925288 KM396842a MH020878 NEKD01000005 E. candida V. Spirin 3921 Russia: KHA KY801867 KY801892 — KY801918 E. candida V. Spirin 8588 USA KY801870 KY801895 — KY801920 E. candida var. cartilaginea V. Spirin 10105 Russia: LEN KY801873 KY801898 — KY801923 E. crenata F. Wu 26 Canada: ON MT663361 MT664780 MT679213 MW358922b E. glandulosa H. H. Burdsall 12029 USA: WI MW353407 MW353407 LOAW01000284 LOAW01000070 E. glandulosa M. Weiss 355 Germany AF291273 AF291319 — — E. glandulosa Y. C. Dai 17633 China MH213393 MH213425 MH213456 — E. japonica F. Wu 251 China MN850378 MN850367 MN819823 — E. qinghaiensis HMAS 156376 China MW353408 MW353408 MW358923 MW358925 E. qinghaiensis HT HMAS 156328 China MW353409 MW353409 MW358924 MW358926 E. recisa M. Weiss 315 Germany AF291276 AF291322 — — E. repanda LY BR 7046 France MT663367 MT664784 MT679214 — E. saccharina Roki 88 Germany AF291277 AF291323 — — E. thuretiana M. Weiss 373 Germany AF291278 AF291324 — — E. thuretiana V. Spirin 9999 Finland KY801878 KY801905 — KY801927 E. truncata M. Weiss 365 Germany AF291279 AF291325 — — E. uvapassa AFTOL-ID 461 Japan DQ241776 AY645056 — — a Sequence derived from Y. C. Dai 13648, China b Sequence derived from R. G. Thorn 200424/02 (University of Western Ontario herbarium), Canada: ON Abbreviations: AFTOL-ID, Assembling the Fungal Tree of Life identifier; HMAS, Mycological Herbarium, Institute of Microbiology, Academia Sinica, Beijing, China; HT, holo- type; KHA, Khabarovsk Krai; LEN, Leningrad Oblast; LY BR, unknown (Wu et al., 2020); ON, Ontario; WA, Washington; WI, Wisconsin. doi:org/10.47371/mycosci.2021.03.002 ― 213 ― S. Wang, R.G. Thorn / Mycoscience VOL.62 (2021) 212-216

68 Wu 26 angiosperm C O

100 100 Wu 305 China 100 78 Weiss 315 66 100 LY BR 7046 F 99

66 AFTOL 461 Japan

Wu 251 China 98 var Spirin 10105 R 100 Spirin 3921 R 100 100 Spirin 8588 USA W

Weiss 355 100 71 Burdsall 12029 USA WI 93 100 Dai 17633 China

Roki 88

72 100 R10 HMAS 156376 China 67 100 86 HT R12 HMAS 156328 China

100 98 Weiss 373

100 Spirin 9999 Fi

Weiss 365

100 Matheny 2295 USA W 100 Dai 13782 China

0.050

Fig. 1 – Phylogeny of selected members of Exidia based on a maximum likelihood (ML) analysis of nrITS, nrLSU, RPB2 and Tef1 gene regions, with ML boot- strap support (bss) values above 50% shown above the nodes and posterior probabilities from Bayesian analysis below the nodes. The new species E. qinghaien- sis clustered with 100% bss as sister to E. thuretiana from Russia and Finland. All terminals are labelled by their collection number and name by which they were identified. Accession numbers are listed in Table 1. HT = holotype.

Fig. 2 – Rehydrated Exidia qinghaiensis (HMAS 156328, holotype), on dead stem of Betula sp. A: Glossy . B: Slightly granular, areolate abhymenium. Bars: A 5 mm; B 1 mm.

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Fig. 3 – Microscopy of Exidia qinghaiensis (HMAS 156328, holotype). A–E: Ba- sidiospores; F, G: Immature hypobasidia with arrows indicating basal . Bars: 5 μm.

Basidiospores (30 measured from 2 collections) narrowly allan- can be distinguished by their ITS sequences, so sequence data for toid, smooth, hyaline in KOH, nonamyloid, (10.4–)11.2–13.5(– Japanese and topotypical Swedish collections of E. repanda are 14.9) × (3.0–)3.1–3.8(–4.1) μm, on average 12.7 × 3.4 μm, Q = highly desirable.

(2.7–)3.1–4.3(–4.5), Qavg = 3.7; probasidia amygdaloid-limoniform; Within the genus Exidia, there seems to be a range in host spec- hypobasidia pyriform to subglobose, cruciately septate at maturity, ificity reported, from quite narrow to very broad, but only a few astipitate or substipitate, with basal clamp connection, (10.5–)11.3– host records are supported by molecular data. Exidia thuretiana 15.5(–17.2) × (7.5–)7.6–9.4(–10.0) μm, on average 12.8 × 8.5 μm; occurs on a broad range of hardwood species, particularly Fagus,

Qavg = 1.5, with longitudinal septa, developing 4 long, finger-like Ribes and Ulmus (Spirin et al., 2018). Exidia saccharina is reported epibasidia (sterigmata), 8–18 × 1.5–2.2 μm. Tramal hyphae hyaline, on dead branches of Abies, Larix, Picea and Pinus (Wojewoda, 1977; thin-walled and 1.5–2.5 μm diam, embedded in gelatinous matrix, Govorova, 1998; Kirschner, 2010) in Europe and Asia. The two col- with simple clamp connections. Dikaryophyses tortuous-cylindri- lections that we studied of E. qinghaiensis were recorded from Bet- cal and 2.0–3.2 μm diam at their base, some with brownish con- ula, the same host genus as reported for E. repanda in Japan (Aoki tents, sparingly and irregularly branched, the branches narrowing and Tubaki, 1986; Aoki, 1991). Further studies based on more ex- to 0.4 μm diam; gloeocystidia not seen. A thin, apparently acellular, tensive collecting, with culturing, mating studies, and DNA se- brownish epithecium that is present in some mounts (but poor- quence data, may elucidate the possible presence of other members ly-preserved and with abundant rod-shaped and spherical bacteria of Exidia in Asia, and the substrates used by E. qinghaiensis. present) appears responsible for the laccate, caramel color of the hymenial surface. Disclosure Habitat and distribution: Saprobic; growing on attached or fall- en hardwood branches, the substrate identified as Betula. Collected The authors declare no conflicts of interest. All the experiments in fall, distributed in Qinghai Province, China. undertaken in this study comply with the current laws of the coun- Additional specimen examined (paratype): CHINA, Qinghai tries where they were performed. Province, Menyuan County, Xianmi wood farm, approx. 37°17’ N, 101°57’ E, 2,850 m a.s.l., on dead branch of Betula, 19 Oct 2004, Acknowledgments Zhang Xiaoqing, HMAS 156376. Comments: Exidia thuretiana, reported on hardwood branches This study was supported in part by the Scientific Start-up Pro- in Europe and Asia (Wojewoda, 1977; Moore, 1997; Spirin, Maly- gram of Shanxi Agricultural University (No. 2014YJ18), the Collab- sheva, & Larsson, 2018) has the greatest tendency to form extensive orative Innovation Center of The Loess Plateau Edible Fungi and cushion-shaped to irregularly lobed, adpressed fruiting bodies, (Shanxi Province) and the Faculty of Science and Department of but its spores are longer and broader (13–19 × 4.5–6.0 μm; Maly- Biology at the University of Western Ontario. We thank T.-Z. Wei, sheva, 2012) than those of this species. Exidia saccharina, reported Y.-J. Yao and L. Cai (Curator, HMAS) for the loan of specimens for in Europe (Wojewoda, 1977; Moore, 1997) and Asia (Russia; Govo- our study. SW also thanks K. Nygard at the Biotron, University of rova, 1998), has fruiting bodies of similar colors and spores of sim- Western Ontario for guidance in the use of the microscope. ilar size and shape (10–14 × 3.5–4.0 μm; Malysheva, 2012) but oc- curs on conifers. Exidia repanda Fr., as reported from Japan, References resembles E. qinghaiensis in both macromorphology and size of basidiospores (8.5–16 × 2–4.0 μm; 12 × 3 μm on average; Aoki & Aoki, T. (1991). Heterobasidiomycetes in and around Sugadaira, Central Japan. I. Exidia. Bulletin of the Sugadaira Montane Research Center, 12, 13–26. Tubaki, 1986). Wojewoda (1977) reported E. repanda from Betula, Aoki, T., & Tubaki, K. (1986). Cultural study of Exidia repanda, previously unreport- Alnus, , Prunus, and Tilia and compared the species with ed from Japan. Mycologia, 78, 877–887. E. recisa (Ditmar) Fr., which occurs primarily on Salix. Sequences Barber, M., Thorn, G., & Voitk, A. (2011). The mushroom basket: when is an ear not of material from France on Prunus identified as E. repanda (Wu et an ear? Osprey, 42(3), 25. al. 2020) are very distinct and were placed in the E. recisa complex Dentinger, B. M., Margaritescu, S., & Moncalvo, J. M. (2010). Rapid and reliable by Wu et al. (2020) and in our analyses (Fig. 1). All of these species high-throughput methods of DNA extraction for use in barcoding and molecu- doi:org/10.47371/mycosci.2021.03.002 ― 215 ― S. Wang, R.G. Thorn / Mycoscience VOL.62 (2021) 212-216

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