683 MycoKeys

MycoKeys 114: 277-298 (2025) DOI: 10.3897/mycokeys.114.146031

Research Article

Lijiangomyces laojunensis gen. et sp. nov. (Mytilinidiaceae), and Sclerococcum stictae (Dactylosporaceae), a new lichenicolous species from Yunnan, China

Qingfeng Meng'?2®, Paul Diederich*®, Vinodhini Thiyagaraja°®, Damien Ertz®’7®, Xinyu Wang>®®, Natsaran Saichana®®, Kevin D. Hyde?2©, Ruvishika S. Jayawardena?2°®, Shaobin Ful?®

oO AN DT oO FP WO YN —

School of Public Health, Zunyi Medical University, Zunyi City, Guizhou Province 563000, China

Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand

School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand

Musée national d'histoire naturelle, 25 rue Munster, L - 2160 Luxembourg, Luxembourg

Key Laboratory of Phytochemistry and Natural Medicines, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China

Meise Botanic Garden, Research Department, Nieuwelaan 38, 1860 Meise, Belgium

Fédération Wallonie-Bruxelles, Service Général de I'Enseignement Supérieur et de la Recherche Scientifique, rue A. Lavallée 1, 1080 Bruxelles, Belgium Yunnan Key Laboratory for Fungal Diversity and Green Development, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China Kyung Hee University, 26 Kyungheedae-ro, Dongdaemun-gu, Seoul 02447, Republic of Korea

10 School of Pharmacy, Zunyi Medical University, Zunyi City, Guizhou Province 563000, China Corresponding authors: Ruvishika S. Jayawardena (ruvishika.jay@mfu.ac.th); Shaobin Fu (fushb@126.com)

OPEN Qaceess

This article is part of:

Exploring the Hidden Fungal Diversity: Biodiversity, Taxonomy, and Phylogeny of Saprobic Fungi

Edited by Samantha C. Karunarathna, Danushka Sandaruwan Tennakoon, Ajay Kumar Gautam

Academic editor: Samantha C. Karunarathna Received: 6 January 2025 Accepted: 2 February 2025 Published: 4 March 2025

Copyright: © Qingfeng Meng et al.

This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0).

Abstract

Lijiangomyces laojunensis gen. et sp. nov. and Sclerococcum stictae sp. nov. are reported from China and identified through DNA sequence analyses (LSU, ITS, and tefl-a) and morphological characteristics. Phylogenetic analysis showed that L. laojunensis forms a distinct lineage within Mytilinidiaceae, closely related to the Mytilinidion subclade, leading to the establishment of a new genus within this fam- ily. This saprotrophic species grows on the bark of Abies fabri, often surrounded by the thallus of Pertusaria species. Lijiangomyces laojunensis is characterized by broadly open, black hysterothecia, clavate asci, and uniseriate, hyaline muriform as- cospores. Sclerococcum stictae, a new lichenicolous species, forms a sister clade relationship to a lichenicolous fungus, S. ricasoliae. It was found on the thallus of Sticta, and is characterized by black apothecia, elongate, cylindrical asci, and brown, elliptical, and 1-septate ascospores. Descriptions, illustrations, and phylogenetic analysis results of the new taxa are provided.

Key words: 2 new taxa, lichen, morphology, phylogeny, saprobe, taxonomy

Introduction

Mytilinidiaceae belongs to Mytilinidiales, Dothideomycetes (Hyde et al. 2024), and was introduced by Kirschstein (1924) to accommodate four genera that were originally classified under Hysteriaceae. Boehm et al. (2009a) introduced Mytilinidiales based on multigene phylogenetic analysis to accommodate Mytilinidiaceae. Pem et al. (2024) included eight genera

poe:

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Citation: Meng Q, Diederich

P Thiyagaraja V, Ertz D, Wang X, Saichana N, Hyde KD, S. Jayawardena R, Fu S (2025) Lijiangomyces laojunensis gen. et sp. nov. (Mytilinidiaceae), and Sclerococcum stictae (Dactylosporaceae), a new lichenicolous species from Yunnan, China. MycoKeys 114: 277-298. https://doi.org/10.3897/ mycokeys.114.146031

within Mytilinidiaceae: Actidium, Lophium, Mytilinidion, Ostreola, Peyronelia, Pseudocamaropycnis, Quasiconcha and Zoggium. Hyde et al. (2024) added three more genera: Bullatosporium, Camaroglobulus and Halokirschsteinio- thelia, increasing the total to eleven genera within this family. Members of Mytilinidiaceae are distinguished by their globose or obovoid pseudothecia which are typically oyster- or hatchet-shaped with a longitudinal keel and crested apex. They feature bitunicate asci containing eight ascospores that are arranged uniseriately, biseriately, or in aggregated clusters. The asco- spores are hyaline to brown, exhibiting diverse morphologies such as scole- cospores, didymospores, phragmospores, or dictyospores, and with bipolar symmetry (Boehm et al. 2009a).

Mytilinidion was established to accommodate the type species M. aggre- gatum. This genus is characterized by globoid to obovoid, erect, conchate, or dolabrate ascomata, and a thin-walled and peridium, bitunicate, 8-spored asci. The ascospores are hyaline to brown and transversely 3-5(—7)-septate (Boehm et al. 2009b; Jayasiri et al. 2018). Ostreola was introduced to accom- modate two species, O. consociate and O. sessilis, characterized by conchiform to hatchet-shaped hysterothecia with a longitudinal slit that opens narrowly. The asci are cylindrical, containing eight uniseriate, brown and muriform as- cospores (Darker 1963). Ostreola shares similarities with Mytilinidion and Lo- phium in the ascomatal morphology but is distinguished by the presence of muriform ascospores. Lumbsch and Huhndorf (2007) placed Ostreola within Mytilinidiaceae. The members of Mytilinidion are primarily saprobic and plant pathogens, while Ostreola species are primarily saprobic.

Sclerococcum belongs to Dactylosporaceae, Sclerococcales, Eurotiomy- cetes (Hyde et al. 2024), and was described by Fries (1819, 1825) to accom- modate the parasitic hyphomycetous fungus S. sphaerale, which was previ- ously classified as Spiloma sphaerale. Phylogenetic analysis by Diederich et al. (2013) revealed that S. sphaerale clusters with two species of Dactylo- spora. In a subsequent study incorporating expanded molecular data, Die- derich et al. (2018) proposed adopting the family name Dactylosporaceae, with Sclerococcaceae as its synonym. They further recommended retain- ing Sclerococcum as the valid genus name and synonymizing Dactylospo- ra under it, as Sclerococcum has nomenclatural priority over Dactylospora. Consequently, 46 Dactylospora species were transferred to Sclerococcum (Diederich et al. 2018). Dactylosporaceae comprises six genera: Cylindro- conidiis, Fusichalara, Gamsomyces, Pseudosclerococcum, Rhopalophora and Sclerococcum (Hyde et al. 2024).

During a survey of microfungi in Yunnan Province (Thiyagaraja et al. 2024), two unidentified fungal specimens were collected. The first spec- imen was found on the bark of Abies fabri, and was often surrounded by the thallus of Pertusaria species while the second was observed growing on the thallus of a foliose lichen belonging to Sticta. Phylogenetic analyses and morphological comparison reveal that the first specimen represents a novel genus closely related to Mytilinidion within Mytilinidiaceae. The sec- ond specimen is proposed as a new lichenicolous species within the genus Sclerococcum based on morpho-molecular analyses.

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Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Materials and methods Sample collection and morphological examination

The specimens were collected from Yunnan Province, China and the im- portant collection information was noted (Rathnayaka et al. 2024). Mac- ro-morphological characteristics were observed using a stereomicroscope (Olympus ZX-16) and photographed with a fitted digital camera (Olympus SC180). The ascomata were sliced, temporarily mounted, and observed un- der a compound microscope (Nikon Y-TV55), with images captured using a fitted digital camera (Nikon DS-Ri2). Distilled water was used as a mounting slide solution and 10% potassium hydroxide (K) and Lugol's iodine solution (I) were used to stain and examine the hymenium. Photographic plates were assembled using Adobe Photoshop CC 2019 software (Adobe Systems, USA). Measurements were conducted using Image Framework software (Tarosoft, Version 0.9.7). The length, width, and length/width ratio (I/w) of asci and ascospores are provided (where n = 10) as: (min—) [X-SD]-[X+SD] (—max), where “min” and “max” represent the extreme observed values, X is the arithmetic mean, and SD is the standard deviation. The number of measurements (n) is indicated. Unless otherwise specified, measurements were taken from water mounts, and procedures followed those specified by Senanayake et al. (2020). The holotype specimens are deposited in the Li- chen Herbarium of Kunming Institute of Botany (KUN-L), Chinese Academy of Science, Yunnan, China.

DNA extraction, PCR amplification, and sequencing

Apothecia were carefully removed with a razor blade under a dissecting mi- croscope, and the lichen thallus was thoroughly cleaned. The sample was then transferred to a 200 uL centrifuge tube. Total genomic DNA was ex- tracted using a Forensic DNA Kit (Omega Bio-Tek, Norcross, Georgia), fol- lowing the manufacturer's instructions. The primer pairs ITS1f/ITS4, LROR/ LR5, 983F/2218R and mrSSU1/mrSSU3R were used respectively, to amplify the internal transcribed spacer region of rDNA (ITS), the 28S large-subunit of rDNA (LSU), the translation elongation factor 1-alpha (tef1-a), and the mitochondrial small subunit ribosomal RNA gene (mtSSU) (Vilgalys and Hester 1990; White et al. 1990; Gardes and Bruns 1993; Zoller et al. 1999; Rehner and Buckley 2005).

The polymerase chain reaction (PCR) was performed using a Mastercycler (Bio-RAD T-100) in a 25-uL reaction volume consisting of 12.5 uL of 2 x Mix (Solarbio, dNTPs Mix), 9.5 uL of double-distilled water (ddH,0), 1.0 pL of each primer (10 mM), and 1.0 uL of the DNA template. The PCR conditions were as follows: an initial denaturation at 95 °C for 3 minutes, followed by 35 cycles of denaturation at 95 °C for 45 seconds, annealing at 53 °C for ITS, LSU, and mtS- SU or at 58 °C for tef1-a, for 90 seconds, and elongation at 72 °C for 1 minute. A final extension was performed at 72 °C for 10 minutes, and the reaction was then held at 4 °C indefinitely. PCR products were sequenced by Shanghai San- gon Biotech (Chengdu, Sichuan Province, China).

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 279

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Sequence alignment and phylogenetic analyses

The quality of chromatogram sequences was verified using BioEdit Sequence Alignment software (Version 7.0.9.0). Forward and reverse sequences were assembled with ContigExpress software (New York, USA). The newly gener- ated sequences were subjected to BLASTn searches (https://blast.ncbi.nim. nih.gov/Blast.cgi) and deposited in the GenBank database. BLAST analysis of three genes (LSU, ITS, and tef1-a) from specimen KUN-L 88703 indicated that this species might belong to Gloniales, Hysteriales, or Mytilinidiales. To confirm its phylogenetic placement, all available sequence data from these orders were downloaded from GenBank (Table 1). Additionally, representative species from six adjacent orders, viz. Botryosphaeriales, Capnodiales, Do- thideales, Jahnulales, Patellariales, and Pleosporales were included based on references from Boehm et al. (2009b). Two species of Orbilia (Orbiliomycetes, Orbiliales, Orbiliaceae) were selected as the outgroup taxa following Hong- sanan et al. (2020) for phylogenetic analysis.

The BLAST analysis of three genes (LSU, ITS, and mtSSU) from the specimen KUN-L 88687 indicated that this species belongs to Sclerococcales. Sequence data for all available taxa within this order were retrieved from GenBank and are listed in Table 2. Two species of Caliciopsis (Coryneliaceae, Coryneliales, Coryneliomycetidae, Eurotiomycetes) were included as the outgroup following Olariaga et al. (2019).

Sequence alignment, concatenation, model selection, and format con- version were performed using the OFPT program (Zeng et al. 2023). Each gene region dataset was aligned using the ‘auto’ strategy in MAFFT (Katoh and Standley 2013) and trimmed with the ‘gappyout’ command in TrimAl (Capella-Gutierrez et al. 2009). The best-fit nucleotide substitution models for each dataset were selected using the Bayesian Information Criterion (BIC) from twenty-two common DNA substitution models with rate hetero- geneity, as implemented in ModelFinder (Kalyaanamoorthy et al. 2017). The datasets were then concatenated with partition data for subsequent phylogenetic analyses.

Maximum likelihood (ML) analysis was conducted on the IQ-TREE web server applying the ultrafast bootstrap approximation with 1,000 repli- cates (Hoang et al. 2018), and the SH-like approximate likelihood ratio test (SH-aLRT) (Guindon et al. 2010; Nguyen et al. 2015). The consensus tree was summarized using the extended majority rule. For further verification, an additional ML analysis was performed with RAxML-HPC2 on ACCESS (v8.2.12), using the GTRGAMMA model with a rapid bootstrap analysis of 1000 replicates (Miller et al. 2010; Stamatakis 2014).

Bayesian inference was carried out using two parallel Metropolis-coupled Markov Chain Monte Carlo (MCMC) runs, each consisting of one ‘cold’ chain and three heated chains, in MrBayes (Ronquist et al. 2012). Trees were sam- pled every 1,000 generations, and the run was terminated when the average standard deviation of split frequencies fell below 0.01. The final tree was sum- marized after discarding the first 25% of samples as burn-in and visualized in FigTree v1.4.4 (Rambaut 2016). The newly identified taxon was registered in Index Fungorum and Faces of Fungi database (Jayasiri et al. 2015).

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Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Table 1. Sequences used in phylogenetic analysis of Mytilinidiales and adjacent orders, with specimens or strains information

and GenBank accession numbers. Newly obtained sequences are in bold font. “NA” indicates the sequence is unavailable.

Species name

Aigialus grandis A. mangrovis

A. parvus

A. rhizophorae

Aliquandostipite crystallinus

A. khaoyaiensis Ascagilis guttulaspora

A. submersa

A. thailandensis Botryosphaeria dothidea Brachiosphaera tropicalis Bullatosporium taxicola B. taxicola

Capnodium aciculiforme C. alfenasii

C. coartatum Cenococcum geophilum C. geophilum Chaetocapnodium insulare C. philippinense

C. placitae Conidiocarpus asiaticus C. caucasicus

C. fici-septicae

C. siamensis

Delitschia chaetomioides D. winteri

Dothidea insculpta D. sambuci Ericboehmia centramura

EF. curtisii

E. doimaeensis Fusculina eucalypti Gloniopsis arciformis G. calami

G. leucaenae

G. percutanea Glonium circumserpens G. circumserpens

G. stellatum

G. stellatum Glyphium elatum

Gordonomyces mucovaginatus

Graphyllium caracolinense Guignardia gaultheriae

Voucher/strains

BCC 20000 BCC 33563 PUFD45 BCC 33572 AFO07 MFLUCC 21-0106 MFLUCC 17-0244 MFLUCC 18-1143 MFLUCC 18-1149 CBS 115476 SS 2523 a21-004; CBS 151403 a21-005; CBS 151402 CBS 892.73 CBS 146151 CPC 17779 CG5 CG54 CBS 146159 MFLUCC 12-0110 CBS 124758 MFLUCC10-0062 GUMH937 MFLUCC 19-0072 SICAUCC 23-0010 DSE871 AFTOL-ID 1599 CBS 225.62 CBS 189.58 DAOM 231303 chuni 70 MFLUCC 12-0808 CBS 198.34 MFLUCC 16-0329 CBS 120083 GKM L166A MFLUCC 15-0739 MFLU 21-0201 FMR 8713 CBS 123342 CBS 123343 ANM 32; A. Miller 32, F CBS 207.34 EB 0365; BPI 892671 CBS 127273 HUEFS 42838 CBS 447.70

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031

GenBank accession numbers

ITS NA NA MK028710 NA NA MT864350 NA NR_171970 NR_171969 NA FJ887923 PP516536 PPat6525 NA MN749233 MN749236 KC967409 KC967410 NR_168830 NR_168831 MH863403 NA NA MW063143 OR405901 MW209042 NA

NA NA KM272258

NA MH535872 DQ923531

NA NR_164398 OL782134 AM286786

NA

NA MZ570257 KM220945 NR_157428

NA MH859790

LSU GU479775 GU479776 MK026761 GU479780 EF175652 MT860428

NG_064432 MN888485 MN913693 DQ678051 JN819284 PPS16533 PP516534 GU301847 MN749165 MN749167

NA NA

NG_068681 KP744503 MH874920 JN832612 KC833050 MW063206 OR405912 MW209067 DQ6/8077

DQ247802 AY544681 KM272256

MH866967 MH9535894 DQ923531 GU323211 NG_059715 OL782050 LS997561 FJ161208 FJ161200 GQ221887 FJ161179 KM220939 NG_057941 NG_060651 DQ678089

tefl1-a

NA GU479840 MN520611 GU479844

NA MT873577

NA

NA DQ767637 JN819298 PP514386 PP514385 GU349045 MN829346 MN829348

NA

NA MN829359 MN829362 MN829363

NA

NA OR671432 MW238837 DQ677922

DQ471081 DQ497606 KM277819

FJ161093

NA

NA KX671965 OL875100 LS997569

NA

NA GQ221926 FJ161095 KM220933

NA

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Species name

Halokirschsteiniothelia maritima H. maritima

H. maritima

Hysterium angustatum

H. pulicare

H. rhizophorae

Hysterobrevium baoshanense

H. constrictum

H. rosae

Hysterodifractum partisporum Hysterographium didymosporum H. fraxini

H. minus

Hysteropatella elliptica

H. prostii

Jahnula appendiculata J. dianchia

J. rostrata

Lophium arboricola

L. arboricola

L. mytilinum

L. zalerioides Massaria inquinas

M. vomitoria Mytilinidion acicola

M. acicola

M. andinense

M. australe

M. californicum

M. didymospora

M. mytilinellum

M. resinicola

M. rhenanum

M. scolecosporum

M. thujarum

M. tortile

M. tortile Neocamarosporium goegapense N. phragmitis Neomassaria fabacearum N. formosana Oedohysterium insidens

O. insidens O. sinense

Voucher/strains

3124D CBS 221.60 NWHC 45703-222 KUMCC 21-0213 EB 0238; CBS 123377 MFLUCC 15-0950 MFLUCC 16-2162 KUN-HKAS102101 CBS 149699 HUEFS 42865 MFLUCC 10-0101 CBS 109.43 JCM 2758 AFTOL-ID 1790 CBS 935.97 G.M. 2016-02-20.2 BCC11400 KUMCC 17-0039 MFLU 20-0435 CBS 758.71 FMR 3868 P99; KRAM F-59986 CBS 114111 CBS 269.34 MFLUCC 14-0417 WU 30527 WU 30606 EB 0349; BPI 879794 EB 0379; BPI 879793 EB 0330; CBS 123562 CBS 301.34 EB 0385; BPI 879795 MFLUCC 16-0619 CBS 303.34 EB 0386; BPI 879796 CBS 303.34 CBS 304.34 CBS 135.34 EB 0341; CBS 135.45 CBS 305.34 EB 0268; BPI 879797 CBS 306.34 EB 0377; BPI 879798 CBS 138008 MFLUCC 17-0756 MFLUCC 16-1875 NTUCC 17-007 ANM 1443 A. Miller 1443, F CBS 238.34 EB 0339; BPI 879800

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031

GenBank accession numbers

ITS KM272366 NA MK782369 OK482567 NA NR_189349 MZ467049 MN429070 0Q990113 NA NA NA NA NA

MT341324 JN819280 KY928456 MT627720 NA KU705825 OR754902 EF596819 OM337540 MF621583 HQ599402 HQ599437 NA NA NA NR_160067 NA NA NA NA MH855535 NA NA MH855536 NA MH855537 NA KJ869163 MG844345 NA NA NA

NA NA

LSU NA AY849943 NA OK482568 FJ161201 NG_241879 KX772765 MN429073 0Q990064 NG_060652 NG_064526 FJ161171 NG_059814 DQ7/6/657

MT341324 FJ743446 KY928457 MT627657 MH872091 KU705842 OR/54924 EF596819 MH867013 MF621587 HQ599402 HQ599437 GU323209 GU397346 FJ161199 MH867035 GU323208 MH535902 MH867037 GU397347 MH867038 NA GU323207 MH867039 GU323206 MH867040 GU323205 KJ869220 NG_070431 KX524145 MH714756 GQ221882

FJ161182 GU397348

tefl-a NA GU349001 NA NA FJ161109 MF615401 KX772769 MN442088 0Q989245 NA NA FJ161088 NA DQ767640

NA JN819299 NA NA NA NA NA NA NA NA HQ599342 HQ599375 NA NA FJ161107 NA NA NA FJ161100 NA FJ161101 FJ161092 NA FJ161102 NA NA NA NA MG844351 KX524149 MH714762 NA

FJ161097 GU397339

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Species name Voucher/strains

Orbilia auricolor AFTOL-ID 906 CBS 547.63 O. vinosa AFTOL-ID 905 GBS:917.72 Ostreichnion sassafras CBS 322.34 Lijiangomyces laojunensis KUN-L 88703 Patellaria apiculatae MCD 096; MFLU 19-1236 P atrata CBS 958.97 P. chromolaenae MFLUCC 17-1482 Pseudocamaropycnis pini CBS 115589

Culture BA4b001 NBRC 111599 FLAS-F-59166

CBS 112412 MFLU 11-0214 MFLU 18-2218

MFLUCC 16-0611

MFLUCC 17-0319

EB QR; RLG 14189

Pseudocenococcum floridanum

Psiloglonium araucanum

P colihuae

P macrosporum Purpurepithecium murisporum P murisporum

Quasiconcha reticulata

Quasiconcha sp. ZY 22.011 CGMCC 3.25498 Quasiconcha sp. LY 27-0: 2 CGMCC 3.25498 Quasiconcha sp. LVN 22.013 CGMCC 3.25498 Rhytidhysteron bannaense KUMCC 21-0483 R. bruguierae SDBR-CMU 473 R. camporesii KUNCC 22-12388 Yuccamyces citri CBS 143161 Y. pilosus CBS 579.92

Table 2. Sequences used in phylogenetic analysis of Dactylosporaceae with specimens or strains’ information and Gen-

GenBank accession numbers

ITS DQ491512

DQ491511

MH855548

PQ049177

MN047094 NA

MT214381

KU728518 NA

NA KP744466 OR225075

NA

NA OR680490

OR680491

OR680492

OP526399 0Q943970 OR807853 MG386043 MG386044

LSU DQ470953

DQ470952

FJ161188 PQ047633 MNO17860 GU301855 MT214475 KU728557 LC095431

FJ161172 KP744511 OP612525 NG_059797 KY799174 GU397349 OR680557

OR680558

OR680559

OP526409 0Q940376 OR801302 MG386096 MG386097

tef1-a DQ471072

DQ471071

NA PQ267963 NA GU349038 MT235796

LC095383

FJ161089 NA OR140436 KY887666 KY799177 NA OR865892

OR865893

OR865894

OP572200 0Q973477 OR832866 NA NA

Bank accession numbers. Newly obtained sequences are in bold font. “NA” indicates the sequence is unavailable.

Species name Voucher/strains

Umbilicaria sp. INB_i04503Q Umbilicaria sp. INB_i04513J Umbilicaria sp. INB_i04513L Caliciopsis orientalis CBS 138.64 C. pinea AFTOL-ID 1869 CBS 139.64 Cylindroconidiis aquaticus MFLUCC 11-0294 Fusichalara minuta CBS 709.88

MFLUCC 18-1015 MFLUCC 18-0982

Gamsomyces aquaticum

G. chiangmaiensis

G. longisporus CBS 118.86 G. longisporus CBS 240.89 G. stilboideus CBS 146494

Pseudosclerococcum golindoi ARAN-Fungi 6619

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031

GenBank accession numbers

LSU

ITS

KM242300 KM242356 KM242358 NG_058741 DQ6/8097

MH236579 KX537758 MN335230 MN335229 MT020877 MT020878 MT020879 NG_073673

KM242300 KM242356 KM242358 NA NA

MH236576 KX537754 MN335228 MN335227 MT020865 MT020866 MT020867 NR_171236

mtSSU NA NA NA FJ190654 FJ190653

NA KX537762 NA NA NA NA NA MK759897

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GenBank accession numbers

Species name Voucher/strains

Sclerococcum ahtii RP 23 KY661659 KY661630 F. Hognabba 1325a (H)

S. ahtii RP182 NA KY661622 CHI17-37a (H)

S. chiangraiensis MFLU 16-0570 NG_066422 NR_163755 A

S. deminutum RP235 N KY661629 J. Pykala 39390 (H)

NA

S. haliotrephum AFTOL-ID 758 FJ176855 ATCC MYA-3590 S. haliotrephum FJ713617 NA AFTOL-ID 798

S. lobariellum ARAN-Fungi 10091 NA

S. martynii MZ221620 MZ221612 ie Bell

S. martynii MZ221623 MZ221616 — ae

S. martynii D. Haelew. F_1577a MZ221619 MZ221610 PUL F27741 S. parasiticum ARAN-Fungi 2724 MK759892

S. parasiticum RP422 KY661666 KY661646 LE 260868

S. pseudobactrodesmium OR514704 S. pseudobactrodesmium OR514705 S. sphaerale JX081674 NA

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031

mtSSU KX537760 KX537759 KX537761 KY661686

NA

NA NA

NA

NA KY661683 ORO035764 KJ766382

NA

MH698503

MH698502

MK759898

KJ766383 NA

NA

MK759899 KY661690

MK759901 MK759902 MK759900 OR588037 OR588038 OR588039 OR588036 MT153924 MT153923 MT153925 MT153922 MZ676669 MF085485 KT263115 KY661689 JX081678 JX081677 JX081676

284

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

GenBank accession numbers

Species name Voucher/strains LSU ITS mtSSU S. stictae KUN-L 88687 PQ407923 PQ408029 PQ415057 S. stictae KUN-L 88687-1 NA PQ408030 NA S. tardum ICMP 24355 NA NR_176187 NA S. tardum PDD 91756 NA OL709435 NA S. tardum PDD 105454 NA MK432753 NA S. stygium ARAN-Fungi 00823 NA MK759886 MK759904 S. stygium ARAN-Fungi 3395 MK759896 NA MK759903 S. stygium BHI-F312 (FH) NA MF161218 NA S. vrijmoediae NTOU 4002 KC692153 NR_138396 NA

Abbreviations: ARAN-Fungi: The ARAN-Fungi Fungarium; BCC: BIOTEC Culture Collection, Thailand; BPI: United States USDA ARS National Fungus Collections, Beltsville, MD; CBS: CBS Fungal Biodiversity Center, Utrecht, The Netherlands; C6MCC: China General Mi- crobiological Culture Collection Center, Institute of Microbiology, Chinese Academy of Sciences, Beijing, China; CPC: Collection of PW. Crous; DAOM: Canadian Collection of Fungal Cultures, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada; FH: FH Fungarium of Helsinki University Museum, Finland; FLAS: University of Florida Herbarium, located at the Florida Museum of Natural History in Gaines- ville, Florida, USA; FMR: Fungal Biodiversity Centre of the University of Valencia, Spain; GZCC: Guizhou Culture Collection, China; GUMH: Guangxi University Microbial Herbarium, China; GZU: Herbarium of the Institute of Botany, University of Graz, Austria; HUEFS: Herbario da Universidade Estadual de Feira de Santana, Brazil; ICMP: International Collection of Microorganisms from Plants, University of Auck- land, New Zealand; JAC: University of Johannesburg Herbarium, USA; JCM: Japan Collection of Microorganisms; KRAM: Wtadystaw Szafer Institute of Botany, Polish Academy of Sciences Herbarium; KUMCC: Kunming Institute of Botany Culture Collection; KUN-HKAS: Herbarium of Kunming Institute of Botany, Chinese Academy of Sciences, Yunnan, China; KUN-L: Lichen Herbarium of Kunming Insti- tute of Botany, Chinese Academy of Science, Yunnan, China; MFLU: the herbarium of Mae Fah Luang University, Chiang Rai, Thailand; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand; NBRC: NITE Biological Resource Center, Japan; NTOU: National Taiwan Ocean University; NWHC: National Wildlife Health Center of U.S. Geological Survey; PDD: Plant Disease Database at the Auckland Museum, New Zealand; PUFD: Purdue University Forestry Department; PUL: Purdue University Herbarium; RLG: The Robert L. Gilbertson Mycological Herbarium at the University of Arizona; SDBR-CMU: Sustainable Development of Biodiversity Resources, Chiang Mai University, Thailand; SICAUC:Sichuan Agricultural University Culture Collection;WU: Herbarium of the University of Vienna.

Results

In the analysis of Mytilinidiales, the final dataset comprised 116 taxa (Table 1) with 2246 aligned characters, including gaps (ITS 1-494 bp, LSU 495- 1346 bp, and tefl-a 1347-2246 bp). The best-fit models for each gene, de- termined using the Bayesian information criterion (BIC), were as follows: ITS: SYM+I+G4, LSU: TN+F+R4, and tefl-a: TN+F+l+G4. For the combined data- set (LSU, ITS, and tef1-a), the parameters of the GTRGAMMA model were as follows: estimated base frequencies: A = 0.24, C = 0.25, G = 0.28, T = 0.23; substitution rate: AC = 1.12, AG = 3.35, AT = 1.77, CG = 0.88, CT = 8.54, GT = 1.00; gamma distribution shape parameter (a) = 0.325118; and tree- length = 6.868758. The best-scoring RAxML tree was constructed with a final maximum likelihood (ML) optimization likelihood value of -34,803.39. Bayesian posterior probabilities (BYPP) were calculated using MCMC anal- ysis, achieving a final average standard deviation of split frequencies of 0.009998. The final tree topologies of ML and BYPP analyses were consis- tent. The best-scoring RAxML tree, based on combined LSU, ITS, and tef1-a sequence datasets, is presented in Fig. 1.

The resulting phylogram distinguishes eight order-level clades, most of which are well-supported, except forthe Patellariales clade. Sevengenera forma strong- ly supported Mytilinidiales clade, with Mytilinidion appearing polyphyletic. The newly identified species clusters within the Mytilinidiales clade are closely re- lated to a Mytilinidion subclade containing eight species. The remaining Mytilin- idion species form a separate clade alongside Lophium. Halokirschsteiniothe- lia and Quasiconcha form a sister clade to the Ostreola-Mytilinidion grouping.

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 285

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

79/1.00

| Hysteriales 100/1.00 | Mytilinidion acicola EB 0349 Mytilinidion acicola EB 0379 400/1.00 99/1.00 Mytilinidion tortile EB 0377 \—_ Mytilinidion thujarum EB 0268 | Mpytilinidion andinense EB 0330 Mytilinidion Mytilinidion australe CBS 301.34 Mytilinidion mytilinellum CBS 303.34 | 400/1.007 Mytilinidion mytilinellum EB 0386

89/1.00

98/1.00

MN Ssess Mytilinidion didymospora MFLUCC 16-0619 (<P) Mytilinidion californicum EB 0385 — Lijiangomyces laojunensis KUN-L 88703 Lijiangomyces om Quasiconcha sp. ZY 22-013 SS SE Ue 99/1.00| Quasiconcha sp. ZY 22-011 F | Quasiconcha sp. ZY 22-012 Quasiconcha om Quasiconcha reticulata EB QR = ee Halokirschsteiniothelia maritima NWHC 45703-222 — : Halokirschsteiniothelia maritima CBS 221.60 Halokirschsteiniothelia

Halokirschsteiniothelia maritima 3124D Lophium arboricola CBS 758.71 93/1.00 Lophium arboricola FMR 3868

i SAG Lophium zalerioides MFLUCC 14-0417 Lophium

Myt

Lophium arboricola P99 88/1.00

8210-28 OBST Lophium mytilinum CBS 269.34

jon a3 wes Lophium mytilinum CBS 114111 SSS Mytilinidion resinicola CBS 304.34 100/1.00 Kl .| Mytilinidion rhenanum CBS 135.34 BS Mytilinidion rhenanum EB 0341

Mytilinidion scolecosporum CBS 305.34 My tilinidion Mytilinidion tortile CBS 306.34 Pseudocamaropycnis pini CBS 115589 Pseudocamaropycnis 00, Bullatosporium taxicola a21-004 Bullatosporium taxicola a21-005

Bullatosporium 82/1.00} | | 22/1-00

98/1.00 100/1.00 ~~ | Capnodiales

100/1.00 — Dothideales

100/1.00 eer. ee DI Botryosphaeriales

Patellariales

] Pleosporales

100/1.00

| Jahnulales SE a Orbilia vinosa AFTOL_ID 905 Out Orbilia auricolor AFTOL_ID 906 bl aoa

0.05 Figure 1. RAXML analysis of Mytilinidiales based on the combined LSU, ITS, and tef1-a sequence data. Bootstrap sup- port values for maximum likelihood (ML = 70%), and the Bayesian Posterior Probabilities (PP = 0.90) are shown near the nodes as ML/PP. Orbilia auricolor (AFTOL-ID 906) and O. vinosa (AFTOL-ID 905) were used as outgroups. The newly generated sequence is in red bold font.

Additionally, two Bullatosporium strains form a distinct clade closely related to other genera within the Mytilinidiales order.

In the analysis of Dactylosporaceae, the final dataset comprised 62 taxa (Table 2) with 2041 aligned characters, including gaps (ITS 1-455 bp, LSU 456-1310 bp, and mtSSU 1311-2041 bp). The best-fit nucleotide substitution models, selected based on the Bayesian information criterion (BIC), were as follows: ITS: TIM2e+1+G4, LSU: TNe+Il+G4, mtSSU: TVM+F+l+G4. The param- eters for the GTRGAMMA model of the combined LSU, ITS, and mtSSU were as follows: estimated base frequencies: A = 0.29, C = 0.19, G = 0.25, T = 0.26, and substitution rate AC = 1.01, AG = 2.68, AT = 1.67, CG = 0.85, CT = 5.43, and GT = 1.00. Gamma distribution shape parameter a = 0.256278, and the tree length = 2.405501. The best-scoring RAxML tree was constructed with a final ML optimization likelihood value of - 14773.67. The final tree topologies of ML and BYPP analyses were consistent. The best-scoring RAXxML tree, based on combined LSU, ITS, and mtSSU sequence datasets, is presented in Fig. 2. The newly collected specimen clusters within the Sclerococcum clade and is closely related to S. ricasoliae.

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 286

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

95//0.91; Sclerococcum stygium ARAN-Fungi 3395 -100/1.00" 1 Sclerococcum stygium ARAN-Fungi 00823 90/1.00 Sclerococcum stygium BHI-F312 (FH)

100/1.00, Sclerococcum martynii D. Haelew. F-1570b 100/1.00 | 1001.60"! Sclerococcum martynii D. Haelew. F-1567b : Sclerococcum martynii D. Haelew. F-1577a

100/1.00) Sclerococcum fusiformis MFLU 18-0678 0.97 -9511,00[ Selerocaceum fusiformis MFLU 16-0593 Sclerococcum chiangraiensis MFLU 16-0570 Sclerococcum mangrovei AFTOL_ID 2108 ; Selerococcum haliotrephum AFTOL-ID 798 Sclerococcum haliotrephum AFTOL-ID 758 Sclerococcum vrijmoediae NTOU 4002 100/0,93, Sclerococcum parasiticum F283586 Sclerococcum parasiticum F283587 Sclerococcum parasiticum RP422 Sclerococcum parasiticum ARAN-Fungi 2724 Sclerecoccum parasiticum ARAN-Fungi 43044025 100/1.00) Sclerococcum tardum PDD 91756 99/1.00 | | Sclerococcum tardum ICMP 24355 Sclerococcum tardum JAC13187 Sclerococcum ricasoliae A. F. 25611 Sclerococcum ricasoliae A. F. 25967 Sclerococcum ricasoliae A. F, Fla6b Sclerococcum ricasoliae A. F. 29132 100/1.00) Sclerococcum stictae KUN-L 88687 Sclerococcum Sclerococcum stictae KUN-L 88687-1 Sclerococcum lobariellum Diederich 17708 Sclerococcum lobariellum Diederich 18109 Sclerococcum lobariellum ARAN-Fungi 10091 Sclerococcum sp. A1153 Sclerococcum sp. A1016 Sclerococcum sphaerale Diederich 17279 Sclerococcum sphaerale Diederich 17283 Sclerococcum sphaerale Ertz 17425 (BR) Sclerococcum simplex MFLU 21-0117

4 Sclerococcum pseudobactrodesmium GZCC 23-0056 ' Sclerococcum pseudobactrodesmium GZCC 23-0057

89/1.00

100/1.00

Dactylosporaceae

00/1.00! s-ferococcum pseudobactrodesmium GZCC 23-0549 Sclerococcum pseudobactrodesmium CGMCC 3.25577T

100/1.00;— Sclerococcum glaucomarioides RP275 Sclerococcum glaucomarioides KUN-L 88756 Sclerococcum deminutum RP235 Sclerococcum sp. RP391

.00 -— Sclerococcum ahtii RP182 — Sclerococcum ahtti RP23

73/1.00- Gamsomyces longisporus CBS 240.89

86/1.00|! Gamsomyces chiangmaiensis MFLUCC 18-0982

72/- Gamsomyces longisporus CBS 118.86 Gamsomyces

: Gamsomyces stilboideus CBS 146494 Gamsomyces aquaticum MFLUCC18-1015 Rhopalophora clavispora CBS 281.75 Rhopalophora clavispora CBS 129.74 Rhopalophora Rhopalophora clavispora CBS 637.73 Cylindroconidiis aquaticus MFLUCC 11-0294 Cylindroconidiis —— Fusichalara minuta CBS 709.88 Fusichalara

72/-

100/1.00

beetle associate disolate INB 104513L beetle associate fungi beetle associate disolate INB i04503Q 100/1.00 -— Caliciopsis pinea AFTOL-ID 1869 '—_ Caliciopsis orientalis CBS 138.64

Outgroup

0.05 Figure 2. RAxML analysis of Dactylosporaceae based on the combined LSU, ITS and mtSSU sequence data. Bootstrap support values for maximum likelihood (ML = 70%), and the Bayesian Posterior Probabilities (PP = 0.90) are shown near the nodes as ML/PP. Caliciopsis orientalis (CBS 138.64) and Caliciopsis pinea (AFTOL-ID 1869) were used as outgroups. The newly generated sequence is in red bold font.

Taxonomy

Lijiangomyces Meng & Jayaward., gen. nov. Index Fungorum: IF903174 Facesoffungi Number: FoF 17069

Etymology. The genus name “Lijiangomyces’ refers to “Lijiang”, the city in Yun- nan Province of China from where the holotype was collected.

Description. Sexual morph: Ascomata hysterothecia, superficial, solitary, dis- persed, sessile, obovoid to broadly shell-shaped or irregularly rounded, with a

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 287

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

broadly open or slit-like disc. Margin black, vertically erect, fragile, with the disc surface appearing yellowish-brown. Peridium carbonaceous, black in the lateral and upper regions. Hymenium hyaline to slightly yellowish, with a densely packed hamathecium. Paraphyses filiform, hyaline, non-anastomosed, and non-septate. Hypothecium slightly yellowish. Asci bitunicate, 8-spored, elongated to clavate, with a rounded apex lacking ascal wall thickening, I-, K-. Ascospores uniseriate, arranged obliquely and parallelly, hyaline, thin-walled, smooth, fusiform to ellip- soidal, K-, 1+ dark blue, aseptate at immature, becoming muriform at maturity with 4-7 transverse septa and 1-2 longitudinal septa. Asexual morph: Not observed.

Type species. Lijiangomyces laojunensis Meng & Jayaward.

Notes. The genus is distinguished by black, broadly shell-shaped to irregular- ly rounded hysterothecial ascomata, with a broadly open or occasionally closed disc, typically light brown to flesh-yellow. Asci are elongated to clavate, contain- ing eight uniseriate, hyaline, and muriform ascospores. Phylogenetic analysis places this genus within the family Mytilinidiaceae (Mytilinidiales, Dothideomy- cetes) closely related to Mytilinidion. In the single-gene phylogenies, the new species is positioned outside the Mytilinidia sensu stricto clade in the ITS and LSU trees but clusters within it in the tef1-a tree. However, in the concatenated analysis combining all three genes, it is again placed outside the Mytilinidia sen- su Stricto clade. These results support its recognition as a distinct lineage within Mytilinidiaceae. Morphologically, this genus differs significantly from Mytilinidi- on in having obovoid to broadly shell-shaped or irregularly rounded, with broadly open ascomata (vs. globoid to obovoid, conchate, or dolabrate ascomata with narrow slit-like openings) and hyaline and muriform ascospores (vs. hyaline to brown and transverse septa). In addition, this genus shares similar morphology with Ostreola in having muriform ascospores but differs in the broadly shell- shaped or irregularly rounded ascomata (vs. conchiform to hatchet-shaped), a broad disc opening (vs. narrowly slit-like), and hyaline ascospores (vs. brown).

Lijiangomyces laojunensis Meng & Jayaward., sp. nov. Index Fungorum: IF902457

Facesoffungi Number: FoF 16265

Fig; 3

Etymology. The species epithet “/aojunensis” refers to the type locality “Laojun Mountain National Nature Reserve” in Yunnan Province of China.

Holotype. KUN-L 88703.

Description. Sexual morph: Ascomata hysterothecia, (0.8—)0.88-1.05(-1.1) x (0.4-)0.47-0.78(—-0.8) mm (X = 0.97 x 0.63, n = 10), superficial, solitary, dis- persed, sessile, non-stromatic, obovoid to broadly shell-shaped or irregularly rounded, with a broadly open or slit-like disc. Margin black, vertically erect, frag- ile, with a yellowish-brown, slightly depressed disc surface appearing below the rim of the lateral wall. Peridium 70-100 um thick, carbonaceous, black laterally and apically, transitioning to grayish near the base. Hymenium 350-400 um high, hyaline to slightly yellowish, densely packed with hamathecium. Paraphyses 1-2 um wide, unbranched, hyaline, non-anastomosed, non-septate. Hypothe- cium 35-50 um thick, slightly yellowish. Asei (120-)122.6-168.7(-190) x (13- )13.4-16.5(-18) um (xX = 145.7 x 14.9, n = 10), bitunicate, 8-spored, elongated to

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 288

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Figure 3. Lijiangomyces laojunensis sp. nov., (KUN-L 88703) a, b habitat c-e ascomata (arrows) f-h section of ascomata in water i-k asci in water I, m asci in IKI n paraphyses in water o-u ascospores in water. Scale bars: 500 um (e); 200 um (f, g); 50 um (h); 25 ym (i-n); 10 ym (o-u).

clavate, rounded apex, without apical thickening of ascal wall, K-, |-. Ascospores (20—)20.9-28.8(-37.5) x (9-)10.1-15.4(-17.5) (x = 24.8 x 12.8, n = 30) um, K-, I+ reddish brown, then turning to dark blue, uniseriate, arranged obliquely and par- allelly, hyaline, thin-walled, smooth, fusiform to ellipsoidal, aseptate at immature,

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 999

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

becoming muriform at maturity with 4-7 transverse septa and 1-2 longitudinal septa, sometimes slightly constricted at the median septum, rounded at the ends in aged ascospores. Asexual morph: Not observed.

Material examined. CHINA * Yunnan Province, Lijiang City, Laojun Mountain National Nature Reserve, 26°39'N, 99°43'E, 3900 m elev., on the bark of Abies fabri (Pinaceae), 10 Apr 2022, Qing-feng Meng, Ijs-52 (holotype KUN-L 88703).

Notes. L ijiangomyces laojunensis closely resembles Ostreola consociata (the type species of Ostreola) and O. sessilis, in having cylindrical asci and uniseri- ate muriform ascospores. However, the new species is distinguished by its as- comatal morphology, which is broadly shell-shaped or irregularly rounded with a widely opened disc, in contrast to Ostreola species which have conchiform to hatchet-shaped ascomata with a narrow slit-like opening. Furthermore, the as- cospores of the new species are hyaline and larger in size (20.9-28.8 x 10.1- 15.4 um), in contrast, brown and smaller ascospores (14-22 x 6-8 um) are the characteristic feature of Ostreola (Darker 1963).

Phylogenetic analysis places this species as a sister clade to Mytilinidion, however, it can be distinguished by its obovoid to broadly shell-shaped or irreg- ularly rounded ascomata (vs. globoid to obovoid, erect, conchate, or dolabrate), and hyaline and muriform ascospores (vs. hyaline to dark brown and trans- versely septate) (Boehm et al. 2009b; Jayasiri et al. 2018).

Sclerococcum stictae Meng, Diederich & Thiyagaraja, sp. nov. Index Fungorum: IF903175

Facesoffungi Number: FoF17070

Fig. 4

Etymology. The species epithet “stictae” refers to “Sticta”, the host lichen on which the holotype was found.

Holotype. KUN-L 88687.

Description. Sexual morph: Ascomata apothecioid, 200-300 um in diam., rounded, cup-shaped, sessile, erumpent from the host thallus with a narrow base, 170-200 um in diam., either dispersed or occurring in small groups, black, matte. Dise flat, black. Margin distinct, persistent, and concolorous with the disc. Exciple brown, paraplectenchymatous, laterally 40-70 um wide. Epithecium (12—)14.3-21.0(-24) um thick (X = 17.7, n = 30), with dark brown- ish granules. Hymenium (46—)65.2-94.8(—96) um high (xX = 80, n = 30), light brown, and distinctly gelatinized, K/I+ reddish with bluish in epihymenium. Paraphyses 2-3 um wide, anastomosing, branched, septate, with swollen, pig- mented apices. Hypothecium (86-)89.6—95.2(—97) um thick (x = 92.4, n = 30), dark orange-brown, with irregularly shaped hypothecial cells. Asci (40—)44.8- 67.9(-72) x (7-)7.8-11.9(-12) um (x = 56.3 x 9.9, n = 10), bitunicate, nar- rowly clavate to cylindrical, ascus wall thickened at the apex, ocular chamber absent, 8-spored, K/I-, except for the K/I+ blue outer gelatinous coat, most intensely colored around the ascus apex. Ascospores (8-)9.2-11.1(-12) x (5-)5.1-6.2(-7) um (X = 10.1 x 5.6, n = 30), I/w ratio = (1.5-)1.7-1.9(-2) (X = 1.8, n = 30), brown when mature, 1-septate, slightly constricted at the septum, verrucose, slightly asymmetric with a larger upper cell, ellipsoidal, sometimes soleiform. Asexual morph: Not observed.

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 290

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

,, - ghom aes

Sun ib gh ce

R ¥. a 2 . " ero J q 3 : Ls _——— Bes Ep q

Figure 4. Sclerococcum stictae sp. nov., growing on the thallus of Sticta sp. (KUN-L 88687) a, b habitat and appearance of the host lichen c, d appearance of the apothecia e, f section of apothecium in water g immature ascus in water h mature ascus with ascospores in water i asci in | j-o ascospores in water. Scale bars: 1 mm (c); 200 um (d); 100 um (e); 50 um

(f); 10 um (g-i); 5 um (§j-o).

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 291

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Material examined. CHINA + Yunnan Province, Diging Autonomous Prefec- ture, Meili Mountain National Nature Reserve, 28°24'N, 98°48'E, 3300 m elev., on the thallus of Sticta sp., on the bark of Rhododendron lapponicum (Ericace- ae), 21 Apr 2023, Qing-feng Meng, ml-68 (holotype KUN-L 88687).

Notes. Sclerococcum stictae clusters within a well-supported subclade along with S. ricasoliae and S. lobariellum. The basepair comparison with S. ricasoliae revealed 7.5% differences (34/455 bp) in ITS, 2.2% (19/855 bp) differences in LSU, and 1% (7/730 bp) differences in mtSSU sequences (Flakus et al. 2019). Compared to S. Jobariellum, it exhibits 4.21% differences (36/855 bp) in LSU and 1.1% (8/730) in mtSSU. Morphologically, S. stictae resembles S. ricasoliae in its ascomatal and ascospore appearance but can be distinguished by its longer, narrower asci (c. 45-68 x 8-12 um vs. 35-50 x 10-15 um) and broader ascospores (c. 9-11 um vs 4-6 um), with a smaller length/width ratio (1.7-1.9 vs. 1.5-3.5) (Flakus et al. 2019). Phylogenetically, S. stictae is also related to S. lobariellum, but the basepair comparison revealed 4.3% (37/855 bp) differ- ences in LSU and 1.1% (8/730 bp) differences in mtSSU sequences (Hafellner 1979). Hence based on recommendations outlined by Jeewon and Hyde (2016), the establishment of the new species is supported. Diederich et al. (2024) have shown that both Sclerococcum lobariellum and S. ricasoliae possess an asexu- al stage producing dark brown, dispersed, muriform conidia, often co-occurring with the sexual stage. In all other known lichenicolous Sclerococcum species with an asexual stage, conidia are produced within compact sporodochia, and this stage is never accompanied by a sexual stage. We anticipate, therefore, that an asexual stage with dispersed, muriform conidia also exists in the new S. stictae and should be searched for when more specimens become available.

Another species, Sclerococcum dendriscostictae, also found on Sticta, shares morphological traits with S. stictae. However, the new species can be distinguished by its longer asci (c. 45-68 x 8-12 um vs. 33-44 x 9.5-13.5 um) and verrucose ascospore ornamentation, in contrast to smooth-walled asco- spores reported in S. dendriscostictae (Joshi 2021).

Discussion

Hysteriaceous fungi are distinguished by their persistent, carbonaceous, na- vicular pseudothecia with a longitudinal slit opening. Historically, the mytilinid- iaceous fungi, which possess fragile, shell-shaped pseudothecia that dehisce through a longitudinal cristate apex, were considered part of the hysteriaceous. However, Boehm et al. (2009a, 2009b) reclassified these fungi based on phylo- genetic analyses, leading to the establishment of the order Mytilinidiales.

In our phylogenetic analysis, six genera within Mytilinidiales form a dis- tinct yet complex clade. Mytilinidion is split into two subclades, indicating its polyphyletic nature. This is consistent with the previous study by Boehm et al. (2009a). One subclade includes four species of Mytilinidion alongside all Lophium species, forming a well-supported Lophium-Mytilinidion clade. The other subclade contains eight Mytilinidion species with our new collection, al- though this grouping is weakly supported. Morphologically, Mytilinidion spe- cies are defined by globoid to obovoid, conchate, or dolabrate ascomata with narrow slit-like openings, bitunicate, 8-spored asci, and hyaline to dark brown ascospores with 3—5(—7) transverse septa (Boehm et al. 2009b; Jayasiri et al.

MycoKeys 114: 277-298 (2025), DOI: 10.3897/mycokeys.114.146031 292

Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

2018). However, our new species displays distinct morphological features that diverge significantly from those of Mytilinidion, supporting its exclusion from this genus based on both phylogenetic and morphological evidence.

Hyde et al. (2024) accepted one family and eleven genera within Mytilinid- iales, most of which lack muriform septation except Ostreola which was in- troduced by Darker (1963). Although molecular evidence for Ostreola remains unavailable, our new species shares features with it, such as cylindrical asci and uniseriate muriform ascospores but differs from Ostreola in having broadly opened hysterothecia, while Ostreola exhibits narrow, slit-like openings. Addi- tionally, Ostreola typically has dull-brown muriform spores, whereas our spe- cies is distinguished by its hyaline muriform ascospore (Harkness and Cooke 1878; Darker 1963; Tilak and Kale 1968; Rao and Modak 1972; Barr 1987).

Phylogenetic analysis further positions our new species ata significant distance from the recently described genus Bullatosporium (Andreasen et al. 2024). Apart from the six genera included in our phylogenetic analysis, four additional genera within the family, viz. Actidium, Camaroglobulus, Peyronelia, and Zoggium lack mo- lecular data but exhibit distinct morphological characteristics. Actidium is charac- terized by simple, rounded spores (Fries 1815). Camaroglobulus was introduced as the asexual morph of Mytilinidion resinae, though its taxonomic placement re- quires further molecular confirmation (Speer 1986). Peyronelia is defined by brown, fusiform conidia that form short chains of slender, septate, interconnected cells (Ciferri and Fragoso-Romualdo 1927) and Zoggium features broadly filiform or ver- miform spores that are transversely septate and pale-colored (Vasilyeva 2001).

Given these morphological and phylogenetic distinctions, the establishment of a new genus to accommodate our newly identified species is both neces- sary and justified. This classification will provide a clearer framework for under- standing diversity within the order Mytilinidiales.

The holotype of Lijiangomyces laojunensis exhibits another intriguing fea- ture: the base of its apothecium is surrounded by the thallus of Pertusaria sp. This phenomenon has led us to mistakenly identify it as a lichenicolous spe- cies. Although lichenicolous behavior was not confirmed in this study, the close physical association suggests that the two fungi coexist without conflict. It is also possible that lichenicolous species may be identified as more specimens are collected and studied in the future.

Before 2018, Dactylospora and Sclerococcum were considered distinct genera, with Sclerococcum containing 21 species, 19 of which were lichenicolous. Died- erich et al. (2018) synonymized Dactylospora with Sclerococcum and transferred 46 species from Dactylospora to Sclerococcum. Subsequently, Olariaga et al. (2019) transferred 14 non-lichenicolous species of Dactylospora to Sclerococcum. Johnston (2022) introduced six new saprophytic species of Sclerococcum. Since 2018, ten new lichenicolous species and three combinations have been added to the genus (Elix et al. 2019; Flakus et al. 2019; Fryday 2019; Navarro-Rosinés and Romero 2019; Spribille et al. 2020; Joshi 2021; Zhurbenko 2022; Paz-Bermudez et al. 2023; Diederich et al. 2024; Zhurbenko and Diederich 2024). Diederich et al. (2024) have accepted a total of 85 species in Sclerococcum, with 64 lichenico- lous species and the remainder being saprotrophs on liverworts, wood, and bark in both terrestrial and marine habitats (Dong et al. 2020; Thiyagaraja et al. 2022).

The first study of Sclerococcum in China was conducted by Thiyagaraja et al. (2022), who reported a new geographical record of S. simplex, collected from

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a corticolous Pertusaria thallus in Yunnan province. Ma et al. (2023) described the lignicolous asexual species, S. pseudobactrodesmium from Guizhou Prov- ince. Meng et al. (2024) reported a new geographical record of S. glaucom- arioides found on Ochrolechia akagiensis from China. This study provides an additional new Sclerococcum species from China.

Acknowledgments

Qingfeng Meng thanks Thesis Writing Grant of Mae Fah Luang University, Chi- ang Rai, Thailand. Vinodhini Thiyagaraja thanks Chinese Research Fund (proj- ect no E1644111K1) entitled “Flexible introduction of high-level expert program, Kunming Institute of Botany, Chinese Academy of Sciences”, Yunnan Province “Caiyun Postdoctoral Program” in 2023, Choi Wan Postdoctoral Program 2023 and National Postdoctoral funding, China. RS Jayawardena would like to thank the Eminent scholar offered by Kyun Hee University.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by Zunyi Scientific and Technological Innovation Talent Team Training Project (ZSK-RC-[2023]8), Zunyi, Guizhou, China; Zunyi Medical University Aca- demic new seedling cultivation and innovation project (Qian Ke He Fundamental [2017] 5733-063), Zunyi, Guizhou, China.

Author contributions

Qingfeng Meng and Ruvishika S. Jayawardena designed the experiments and struc- tured the manuscript. Qingfeng Meng conducted the experiments, analyzed the data, and drafted the manuscript. Shaobin Fu provided funding. Paul Diederich, Vinodhini Thiyagaraja, Ruvishika S. Jayawardena, and Damien Ertz contributed to data analysis and manuscript revision. Xinyu Wang, Natsaran Saichana, Kevin D. Hyde, and Shao- bin Fu revised the manuscript. All authors reviewed, edited, and approved the final version of the manuscript for publication.

Author ORCIDs

Qingfeng Meng ® https://orcid.org/0000-0001-9814-8238

Xinyu Wang ® hittps://orcid.org/0000-0003-21 66-6111

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Qingfeng Meng et al.: Lijiangomyces, novel genus & a new lichenicolous species of Sclerococcum (China)

Data availability

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

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