Thyridium revised: Synonymisation of Phialemoniopsis under Thyridium and establishment of a new order, Thyridiales

Abstract

The genus Thyridium , previously known as a saprobic or hemibiotrophic ascomycete on various plants, was revised taxonomically and phylogenetically. Sequences of the following six regions, that is, the nuclear ribosomal internal transcribed spacer (ITS) region, the large subunit (LSU) of rDNA, the second largest RNA polymerase II subunit ( rpb2 ) gene, translation elongation factor 1-alpha ( tef1 ) gene, the actin ( act ) gene, and the beta-tubulin ( tub2 ) gene, were generated for molecular phylogenetic analyses of species of this genus. Phialemoniopsis , a genus encompassing medically important species, is synonymised with Thyridium based on molecular evidence and morphological similarities in their asexual characters. The generic concept for Thyridium is expanded to include species possessing both coelomycetous and hyphomycetous complex asexual morphs. In addition to type species of Thyridium , T. vestitum , nine species were accepted in Thyridium upon morphological comparison and molecular phylogenetic analyses in this study. All seven species of Phialemoniopsis were treated as members of the genus Thyridium and new combinations were proposed. A bambusicolous fungus, Pleospora punctulata , was transferred to Thyridium , and an epitype is designated for this species. A new species, T. flavostromatum , was described from Phyllostachys pubescens . The family Phialemoniopsidaceae, proposed as a familial placement for Phialemoniopsis , was regarded as a synonym of Thyridiaceae. A new order, Thyridiales, was established to accommodate Thyridiaceae; it forms a well-supported, monophyletic clade in Sordariomycetes.

Full text

Thyridium revised: Synonymisation of Phialemoniopsis
under Thyridium and establishment of a new order,
Thyridiales
Ryosuke Sugita1,2, Kazuaki Tanaka1
1Faculty of Agriculture and Life Science, Hirosaki University, 3 Bunkyo-cho, Hirosaki, Aomori 036-8561,
Japan 2e United Graduate School of Agricultural Sciences, Iwate University, 18-8 Ueda 3 chome, Morioka,
Iwate 020-8550, Japan
Corresponding author: Kazuaki Tanaka (k-tanaka@hirosaki-u.ac.jp)
Academic editor: Huzefa Raja|Received 7 December 2021|Accepted 11 January 2022|Published 1 February 2022
Citation: Sugita R, Tanaka K (2022) yridium revised: Synonymisation of Phialemoniopsis under yridium and
establishment of a new order, yridiales. MycoKeys 86: 147–176. https://doi.org/10.3897/mycokeys.86.78989
Abstract
e genus yridium, previously known as a saprobic or hemibiotrophic ascomycete on various plants,
was revised taxonomically and phylogenetically. Sequences of the following six regions, that is, the nuclear
ribosomal internal transcribed spacer (ITS) region, the large subunit (LSU) of rDNA, the second largest
RNA polymerase II subunit (rpb2) gene, translation elongation factor 1-alpha (tef1) gene, the actin (act)
gene, and the beta-tubulin (tub2) gene, were generated for molecular phylogenetic analyses of species of
this genus. Phialemoniopsis, a genus encompassing medically important species, is synonymised with y-
ridium based on molecular evidence and morphological similarities in their asexual characters. e generic
concept for yridium is expanded to include species possessing both coelomycetous and hyphomycetous
complex asexual morphs. In addition to type species of yridium, T. vestitum, nine species were accepted
in yridium upon morphological comparison and molecular phylogenetic analyses in this study. All sev-
en species of Phialemoniopsis were treated as members of the genus yridium and new combinations were
proposed. A bambusicolous fungus, Pleospora punctulata, was transferred to yridium, and an epitype is
designated for this species. A new species, T. avostromatum, was described from Phyllostachys pubescens.
e family Phialemoniopsidaceae, proposed as a familial placement for Phialemoniopsis, was regarded as
a synonym of yridiaceae. A new order, yridiales, was established to accommodate yridiaceae; it
forms a well-supported, monophyletic clade in Sordariomycetes.
Keywords
Ascomycota, Phialemoniopsidaceae, phylogeny, Sordariomycetes, taxonomy, yridiaceae
MycoKeys 86: 147–176 (2022)
doi: 10.3897/mycokeys.86.78989
https://mycokeys.pensoft.net
Copyright Ryosuke Sugita & Kazuaki Tanaka. This is an open access article distributed under the terms of the Creative Commons Attribution
License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source
are credited.
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Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
148
Introduction
yridium was originally established to accommodate species with cylindrical, uniseri-
ate, 8-spored asci and muriform, dark-coloured, ascospores (Nitschke 1867). Species
of this genus occur on various plants as saprobic or hemibiotrophic fungi (Eriksson
and Yue 1989; Taylor et al. 1997; Checa et al. 2013). Currently, yridium includes 33
species and is placed in yridiaceae (family incertae sedis, Sordariomycetes; Yue and
Eriksson 1987; Index Fungorum, http://www.indexfungorum.org, 2021). e type
species T. vestitum has been veried to produce both coelomycetous and hyphomycet-
ous asexual morphs, which have phialidic conidiogenous cells with collarette and el-
lipsoidal to allantoid hyaline conidia (Leuchtmann and Müller 1986).
Molecular information on yridium species is available only for two non-type
strains (CBS 113027, CBS 125582) of the type species T. vestitum (Lutzoni et al.
2004; Spatafora et al. 2006; Vu et al. 2019); however, the phylogenetic relationships
among species of this genus are unclear. A recent study on the phylogeny of Sord-
ariomycetes has shown that T. vestitum is closely related to two Phialemoniopsis spp.
(P. endophytica and P. ocularis), but their phylogenetic and taxonomic relationships
have not been claried (Dong et al. 2021; Hyde et al. 2021).
e genus Phialemoniopsis was placed in Phialemoniopsidaceae (Diaporthomyceti-
dae family incertae sedis, Sordariomycetes; Hyde et al. 2021). Species of this genus are
widely distributed in various environments and substrates, including industrial water,
plant materials, raw sewage, and soil (Gams and McGinnis 1983; Halleen et al. 2007;
Su et al. 2016). Several species have been reported from parts of the human body, such
as blood, eye, toenail, skin, and sinus (Perdomo et al. 2013; Tsang et al. 2014), and
some of them have also been isolated from patients with keratomycosis and phaeohy-
phomycosis (Perdomo et al. 2013; Desoubeaux et al. 2014). All species in this genus
are known to be asexual.
In our ongoing taxonomic study of sordariomycetous fungi in Japan, several new
specimens of yridium-like sexual morphs were collected. Single ascospore isolates
from these specimens formed typical Phialemoniopsis-like asexual morphs in culture,
suggesting that both genera are closely related. is study aims to reveal the taxonomic
and phylogenetic relationships between yridium and Phialemoniopsis, and to clarify
their ordinal position in Sordariomycetes.
Material and method
Isolation and morphological observation
All materials were obtained from Japan. Morphological characteristics were observed
in preparations mounted in distilled water by dierential interference and phase con-
trast microscopy (Olympus BX53) using images captured with an Olympus digital

yridium revised 149
camera (DP21). All specimens were deposited in the herbarium at Hirosaki University
(HHUF), Hirosaki, Japan. Single spore isolations were performed from all specimens.
Colony characteristics were recorded from growth on potato dextrose agar (PDA),
malt extract agar (MEA), and oatmeal agar (OA) from Becton, Dickinson and Com-
pany (MD, USA), after a week at 25 °C in the dark. Colony colours were recorded
according to Rayner (1970). To observe the asexual morphs in culture, 5 mm squares
of mycelial agar were placed on water agar containing sterilised plant substrates such as
rice straws and banana leaves. en these plates were incubated at 25 °C for 2 weeks in
the dark. When the substrates were colonised, the plates were incubated at 25 °C under
black light blue illumination for 1–2 weeks to observe sporulation.
Phylogenetic analyses
DNA was extracted from four isolates using the ISOPLANT II kit (Nippon Gene, To-
kyo, Japan) following the manufacturer’s instructions. e following loci were ampli-
ed and sequenced: the internal transcribed spacer (ITS) region with primers ITS1 and
ITS4 (White et al. 1990), the large subunit nuclear ribosomal DNA (LSU) with prim-
ers LR0R (Rehner and Samuels 1994) and LR5 or LR7 (Vilgalys and Hester 1990),
the second largest RNA polymerase II subunit (rpb2) gene with primers fRPB2-5F and
fRPB2-7cR (Liu et al. 1999), the translation elongation factor 1-alpha (tef1) gene with
primers 983F and 2218R (Rehner and Buckley 2005), the actin (act) gene with prim-
ers Act-1 and Act-5ra (Voigt and Wöstemeyer 2000) and the beta-tubulin (tub2) gene
with primers TUB-F and TUB-R (Cruse et al. 2002). PCR products were puried us-
ing the FastGene Gel/PCR Extraction Kit (Nippon Gene, Tokyo, Japan) following the
manufacturer’s instructions and sequenced at SolGent (South Korea). Newly generated
sequences were deposited in GenBank (Table 1).
Primary analysis of LSU-rpb2-tef1 sequences from 88 strains of Sordariomycetes
(Table 1) was conducted to clarify the ordinal/familial placement of yridium (or
Phialemoniopsis) species. Barrmaelia rhamnicola and Entosordaria perdiosa (Xylari-
omycetidae) were used as outgroups. As a secondary analysis, single gene trees of ITS,
act and tub2, and a combined tree of these three loci were generated to assess the spe-
cies boundaries of 17 strains within yridium/Phialemoniopsis (Table 2). All sequence
alignments (LSU, ITS, rpb2, tef1, act and tub2) were produced using the server ver-
sion of MAFFT (http://www.ebi.ac.uk/Tools/msa/mat), checked and rened using
MEGA v. 7.0 (Kumar et al. 2016).
Phylogenetic analyses were conducted using maximum-likelihood (ML) and
Bayesian methods. e optimum substitution models for each dataset were estimated
using Kakusan4 software (Tanabe 2011) based on the Akaike information criterion
(AIC; Akaike 1974) for ML analysis and the Bayesian information criterion (BIC;
Schwarz 1978) for Bayesian analysis. ML analyses were performed using the TreeF-
inder Mar 2011 program (http://www.treender.de) based on the models selected with
the AICc4 parameter (used sequence length as sample size). ML bootstrap support

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
150
Table 1. Isolates and GenBank accessions of sequences used in the phylogenetic
analyses of Sordariomycetes (Fig. 1).
Taxon Isolatea Statusb GenBank accession numbersaRef.c
LSU rpb2 tef1
Acrodictys aquatica MFLUCC 18-0356 HT MG835712 – – 47
Acrodictys bambusicola HSAUP myr9510 KX033564 – – 44
Annulatascus velatisporus A70 18 AY316354 – – 3
Annulusmagnus triseptatus CBS 128831 GQ996540 JQ429258 –25, 29
Ascitendus austriascus CBS 131685 GQ996539 JQ429257 –25, 29
Atractospora reticulata CBS 127884 HT KT991660 KT991649 –41
Atractospora thailandensis KUMCC 16-0067 HT MF374362 MF370951 MF370962 45
Barbatosphaeria arboricola CBS 127689 HT KM492862 KM492901 –38
Barbatosphaeria barbirostris CBS 121149 EF577059 KM492903 –18, 38
Barbatosphaeria varioseptata CBS 137797 HT KM492869 KM492907 –38
Barrmaelia rhamnicola CBS 142772 ET MF488990 MF488999 MF489009 52
Bombardia bombarda AFTOL-ID 967 DQ470970 DQ470923 DQ471095 14
Calosphaeria pulchella CBS 115999 IT AY761075 GU180661 FJ238421 8, 27
Camarops microspora CBS 649.92 AY083821 DQ470937 –13, 14
Camarotella costaricensis MM-149 KX430484 KX451954 KX451982 43
Cancellidium cinereum MFLUCC 18-0424 HT MT370363 MT370486 MT370488 57
Cancellidium griseonigrum MFLUCC 17-2117 HT MT370364 MT370487 –57
Ceratolenta caudata CBS 125234 HT JX066704 JX066699 –33
PRM 899855 JX066705 – – 33
Chaetosphaeria ciliata ICMP 18253 GU180637 GU180659 –27
Chaetosphaeria curvispora ICMP 18255 GU180636 GU180655 –27
Cryptadelphia groenendalensis SH12 EU528007 – – 20
SMH3767 EU528001 – – 20
Diaporthe phaseolorum NRRL 13736 U47830 – – 1
Distoseptispora obpyriformis MFLUCC 17-1694 HT MG979764 MG988415 MG988422 48
Distoseptispora rostrata MFLUCC 16-096 HT MG979766 MG988417 MG988424 48
Endoxyla operculata UAMH 11085 JX460992 KY931927 –34, 49
Entosordaria perdiosa CBS 142773 ET MF488993 MF489003 MF489012 52
Fluminicola aquatica MFLUCC 15-0962 HT MF374366 –MF370960 45
Fluminicola saprotrophitica MFLUCC 15-0976 HT MF374367 MF370954 MF370956 45
Gnomonia gnomon CBS 199.53 AF408361 DQ470922 DQ471094 2, 14
Jobellisia fraterna SMH2863 AY346285 – – 4
Jobellisia luteola SMH2753 AY346286 – – 4
Lanspora coronata AFTOL-ID 736 U46889 DQ470899 –14
Lasiosphaeria ovina SMH4605 AY436413 AY600284 DQ836908 6, 7, 16
Lentomitella cirrhosa ICMP 15131 ET AY761085 KM492911 –11, 38
Lentomitella crinigera CBS 138678 KY931811 – – 49
Linocarpon livistonae HKUM 6520 DQ810205 DQ810248 –10
Magnaporthe salvinii M 21 JF414887 –JF710406 28
Magnaporthiopsis agrostidis CBS 142740 HT KT364754 –KT364756 37
Melanconis stilbostoma CBS 109778 AF408374 EU219299 EU221886 2
Myrmecridium montsegurinum JF 13180 HT KT991664 KT991654 –41
Myrmecridium schulzeri CBS 100.54 EU041826 – – 17
Myrmecridium thailandicum CBS 136551 HT KF777222 – – 30
Neolinocarpon enshiense HKUCC 2983 DQ810221 DQ810244 –10
Neolinocarpon globosicarpum HKUCC 1959 DQ810224 DQ810245 –10
Ophiostoma piliferum CBS 158.74 DQ470955 DQ470905 DQ471074 14
Ophiostoma stenoceras CBS 139.51 DQ836904 DQ836891 DQ836912 16
Papulosa amerospora AFTOL-ID 748 DQ470950 DQ470901 DQ471069 14
Pararamichloridium caricicola CBS 145069 HT MK047488 – – 46
Pararamichloridium livistonae CBS 143166 HT MG386084 – – 54
Pararamichloridium verrucosum CBS 128.86 HT MH873621 – – 56
Phaeoacremonium
fraxinopennsylvanica
M.R. 3064 HQ878595 HQ878609 –26

yridium revised 151
Taxon Isolatea Statusb GenBank accession numbersaRef.c
LSU rpb2 tef1
Phaeoacremonium novae-
zealandiae
CBS 110156 HT AY761081 – – 8
Phomatospora bellaminuta AFTOL-ID 766 FJ176857 FJ238345 –23
Phomatospora biseriata MFLUCC 14-0832A KX549448 – – 51
Phyllachora graminis TH-544 KX430508 – – 43
Pleurostoma ootheca CBS 115329 IT AY761079 HQ878606 FJ238420 8, 23,
26
Pseudostanjehughesia
aquitropica
MFLUCC 16-0569 HT MF077559 –MF135655 53
Pseudostanjehughesia lignicola MFLUCC 15-0352 HT MK849787 MN124534 MN194047 55
Pyricularia borealis CBS 461.65 DQ341511 – – 24
Pyricularia bothriochloae CBS 136427 HT KF777238 – – 30
Rhamphoria delicatula CBS 132724 FJ617561 JX066702 –22, 33
Rhamphoria pyriformis CBS 139024 MG600397 MG600401 –50
Rubellisphaeria abscondita CBS 132078 HT KT991666 KT991657 –41
Sordaria micola CBS 723.96 AY780079 DQ368647 –9, 19
Spadicoides bina CBS 137794 KY931824 KY931851 –49
Sporidesmium minigelatinosa NN 47497 DQ408567 DQ435090 –12
Sporidesmium parvum HKUCC 10836 DQ408558 – – 12
yridium cornearis CBS 131711 HT KJ573450 –LC382144 36
yridium curvatum CBS 490.82 HT AB189156 –LC382142 15
yridium endophyticum ACCC 38980 HT KT799560 – – 42
yridium avostromatum KT 3891 =
MAFF 247509
HT LC655963 LC655967 LC655971 is
study
yridium hongkongense HKU39 HT KJ573447 – – 36
yridium limonesiae CBS 146752 HT MW050976 – – 58
yridium oculorum CBS 110031 HT KJ573449 –LC382145 36
yridium pluriloculosum CBS 131712 HT HE599271 –LC382141 32
KT 3803 =
MAFF 247508
LC655964 LC655968 LC655972 is
study
yridium punctulatum KT 1015 =
MAFF 239669
LC655965 LC655969 LC655973 is
study
KT 3905 =
MAFF 247510
ET LC655966 LC655970 LC655974 is
study
yridium vestitum CBS 113027 AY544671 DQ470890 DQ471058d5, 14
CBS 125582 MH875182 – – 56
Tirisporella beccariana BCC 36737 JQ655450 – – 39
Tirisporella bisetulosus BCC 00018 EF622230 – – 21
Wongia grinii BRIP 60377 KU850470 –KU850466 40
Woswasia atropurpurea CBS 133167 HT JX233658 JX233659 –31
Xylochrysis lucida CBS 135996 HT KF539911 KF539913 –35
Xylolentia brunneola PRA-13611 HT MG600398 MG600402 –50
a Strains and sequences generated in this study are shown in bold.
b ET = epitype; HT = holotype; IT = isotype
c 1: Viljoen et al. 1999; 2: Castlebury et al. 2002; 3: Raja et al. 2003; 4: Huhndorf et al. 2004; 5: Lutzoni et al. 2004;
6: Miller and Huhndorf 2004a; 7: Miller and Huhndorf 2004b; 8: Réblová et al. 2004; 9: Miller and Huhndorf 2005;
10: Bahl 2006; 11: Réblová 2006; 12: Shenoy et al. 2006; 13: Smith et al. 2006; 14: Spatafora et al. 2006; 15: Yaguchi
et al. 2006; 16: Zhang et al. 2006; 17: Arzanlou et al. 2007; 18: Réblová 2007; 19: Tang et al. 2007; 20: Huhndorf et
al. 2008; 21: Pinruan et al. 2008; 22: Réblová 2009; 23: Schoch et al. 2009; 24: ongkantha et al. 2009; 25: Réblová
et al. 2010; 26: Réblová 2011; 27: Réblová et al. 2011; 28: Zhang et al. 2011; 29: Réblová et al. 2012; 30: Crous et al.
2013; 31: Jaklitsch et al. 2013; 32: Perdomo et al. 2013; 33: Réblová 2013; 34: Untereiner et al. 2013; 35: Réblová et al.
2014; 36: Tsang et al. 2014; 37: Crous et al. 2015b; 38: Réblová et al. 2015; 39: Suetrong et al. 2015; 40: Khemmuk et
al. 2016; 41: Réblová et al. 2016; 42: Su et al. 2016; 43: Mardones et al. 2017; 44: Xia et al. 2017; 45: Zhang et al. 2017;
46: Crous et al. 2018; 47: Hyde et al. 2018; 48: Luo et al. 2018; 49: Réblová et al. 2018; 50: Réblová and Štěpánek 2018;
51: Senanayake et al. 2018; 52: Voglmayr et al. 2018; 53: Yang et al. 2018; 54: Crous et al. 2019; 55: Luo et al. 2019; 56:
Vu et al. 2019; 57: Hyde et al. 2021; 58: Martinez et al. 2021.
d is tef1 sequence (DQ471058) of yridium vestitum was excluded from this analysis. A Blast search using this sequence
suggested that it is close to Phialemonium obovatum (Cephalothecales) rather than yridium/Phialemoniopsis (yridiales).

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
152
(ML BS) values were obtained using 1000 bootstrap replicates. Bayesian analyses were
performed using MrBayes v. 3.2.6 (Ronquist et al. 2012), with substitution models se-
lected based on the BIC4 parameter (used sequence length as sample size). Two simul-
taneous and independent Metropolis-coupled Markov chain Monte Carlo (MCMC)
runs were performed for 9,000,000 generations for primary analysis and 1,000,000
generations for secondary analyses (except for the ITS dataset for 1,500,000 genera-
tions) with the tree sampled every 1,000 generations. Convergence of the MCMC
procedure was assessed from the eective sample size scores (all > 100) using MrBayes
and Tracer v. 1.6 (Rambaut et al. 2014). First 25% of the trees were discarded as
burn-in, and the remainder were used to calculate the 50% majority-rule trees and to
determine the posterior probabilities (PPs) for individual branches. ese alignments
were submitted to TreeBASE under study number S28934.
Result
Phylogeny
For primary analysis, ML and Bayesian phylogenetic trees were generated using an
aligned sequence dataset comprising of LSU (1,205 base pairs), rpb2 (1,059 bp) and
tef1 (954 bp). Of the 3,218 characters included in the alignment, 1,478 were variable
and 1,686 were conserved. is combined dataset provided higher condence values
for ordinal and familial classication than those of individual gene trees, with 25 orders
and three families (order unknown) being reconstructed in Sordariomycetes (Fig. 1).
ML analysis of the combined dataset was conducted based on the selected substitution
model for each partition (GTR+G for LSU, J2+G for the rst and third codon posi-
tions of rpb2, J1+G for the second codon positions of rpb2, F81+G for the rst codon
positions of tef1, JC69+G for the second codon positions of tef1, and J2+G for the third
codon position of tef1). e ML tree with the highest log likelihood (–43687.562) is
shown in Fig. 1. Topology recovered by Bayesian analysis was almost identical to that of
the ML tree. All species previously described as Phialemoniopsis (marked with blue circle
in Fig. 1), one species of “Linocarpon”, two species of “Neolinocarpon” and four strains
newly obtained in this study formed a monophyletic clade with the type species of y-
ridium (T. vestitum). eir monophyly was completely supported (100% ML BS/1.0
Bayesian PP; Fig.1). e family yridiaceae was found to be related to Annulatascales
and Myrmecridiales but did not cluster with any existing order in Sordariomycetes.
For secondary analysis, ML and Bayesian phylogenetic trees were generated us-
ing sequences of ITS (483 bp), act (646 bp), tub2 (375 bp), and a combined dataset
of these three regions (1,504 bp). e selected substitution models for each region
were as follows: J2ef+G for ITS, F81+H for the rst and second codon positions of
act, J2+G for the third codon position of act, K80+H for the rst codon positions

yridium revised 153
of tub2, JC69+H for the second codon position of tub2 and TN93+H for the third
codon position of tub2. e ML trees with the highest log likelihood (–1172.0198 in
ITS, –1196.6012 in act, –859.37115 in tub2 and –3315.7254 in ITS-act-tub2) are
shown in Fig. 2. Our results conrmed close phylogenetic relationships between y-
ridium and Phialemoniopsis (Fig. 2A–D). Except for act (Fig. 2B) and tub2 (Fig. 2C),
where sequence data of T. vestitum were unavailable, the existence of ten distinct
species was suggested (Fig. 2A, D). e following three lineages were found in our
four strains (Fig. 2A–D): 1) a bambusicolous lineage (KT 3891) close to T. curvatum
and T. limonesiae, 2) a fungus on Betula maximowicziana (KT 3803) nested with T.
pluriloculosum, which was previously reported from clinical sources (Perdomo et al.
2013), and 3) another bambusicolous lineage represented by two strains (KT 1015
and KT 3905).
Table 2. Isolates and GenBank accessions of sequences used in the phylogenetic analyses of yridium
species (Fig. 2).
Taxon IsolateaSubstrate/Host StatusbGenBank accession numbersaRef.c
ITS act tub2
yridium cornearis CBS 131711 human corneal uid HT KJ573445 HE599252 HE599301 1, 2
UTHSC 06-
1465
shin aspirate HE599285 HE599253 HE599302 2
yridium
curvatum
CBS 490.82 skin lesion HT AB278180 HE599258 HE599307 2
UTHSC R-3447 human eye HE599291 HE599259 HE599308 2
yridium
endophyticum
ACCC 38979 lower stem of Lua
cylindrica (endophyte)
KT799556 KT799553 KT799562 4
ACCC 38980 lower stem of Lua
cylindrica (endophyte)
HT KT799557 KT799554 KT799563 4
yridium
avostromatum
KT 3891 =
MAFF 247509
dead twigs of Phyllostachys
pubescens
HT LC655959 LC655979 LC655975 is
study
yridium
hongkongense
HKU39 the right forearm nodule
biopsy of a human
HT KJ573442 KJ573452 KJ573457 3
yridium
limonesiae
CBS 146752 Skin nodule HT MW050977 MW349126 MW048608 6
yridium oculorum CBS 110031 human keratitis HT KJ573444 HE599247 HE599296 2, 3
UTHSC 05-
2527
peritoneal dialysis catheter HE599281 HE599249 HE599298 2
yridium
pluriloculosum
CBS 131712 human toe nail HT HE599286 HE599254 HE599303 2
KT 3803 =
MAFF 247508
dead wood of Betula
maximowicziana
HT LC655960 LC655980 LC655976 is
study
UTHSC 09-
3589
synovial uid HE599287 HE599255 HE599304 2
yridium
punctulatum
KT 1015 =
MAFF 239669
dead culms of Phyllostachys
pubescens
LC655961 LC655981 LC655977 is
study
KT 3905 =
MAFF 247510
dead twigs of Phyllostachys
nigra var. nigra
ET LC655962 LC655982 LC655978 is
study
yridium vestitum CBS 125582 MH863721 – – 5
a Strains and sequences generated in this study are shown in bold.
b ET = epitype; HT = holotype
c 1: Tang et al. 2007; 2: Perdomo et al. 2013; 3: Tsang et al. 2014; 4: Su et al. 2016; 5: Vu et al. 2019; 6: Martinez et
al. 2021.

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
154
Figure 1. Maximum-likelihood tree of Sordariomycetes based on combined LSU, rpb2 and tef1 se-
quence. ML bootstrap proportion (BP) greater than 70% and Bayesian posterior probabilities (PP) above
0.95 are presented at the nodes as ML BP/Bayesian PP and a node not present in the Bayesian analysis
is shown with ‘x’. A hyphen (‘-’) indicates values lower than 70% BP or 0.95 PP. Ex-holotype, isotype,
paratype and epitype strains are shown in bold and the newly obtained sequences are shown in red. Strains
previously described as Phialemoniopsis species are marked with a blue circle. e scale bar represents
nucleotide substitutions per site.

yridium revised 155
Figure 2. Maximum-likelihood tree of yridium species based on each ITS (A), act (B), tub2 (C) and
combined sequences (ITS-act-tub2; D). ML bootstrap proportion (BP) greater than 70% and Bayesian
posterior probabilities (PP) above 0.95 are presented at the nodes as ML BP/Bayesian PP. A hyphen (‘-’)
indicates values lower than 70% BP or 0.95 PP and a node not present in the Bayesian analysis is shown
with ‘x’. Ex-holotype and epitype strains are shown in bold and the newly obtained sequences are shown
in red. Strains previously as Phialemoniopsis species are marked with a blue circle. e scale bars represent
nucleotide substitutions per site.

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156
Taxonomy
A new order, yridiales, is introduced to accommodate yridiaceae because its line-
age is phylogenetically and morphologically distinct from any known orders in Sord-
ariomycetes. We concluded yridium and Phialemoniopsis to be congeneric based on
their morphological similarities and phylogenetic relatedness. An expanded generic cir-
cumscription of yridium that integrates the generic concept of Phialemoniopsis is pro-
vided below. One new species and eight new combinations of yridium are proposed.
yridiales R. Sugita & Kaz. Tanaka, ord. nov.
MycoBank No: 841916
Type family. yridiaceae J.Z. Yue & O.E. Erikss., Syst. Ascom. 6(2): 233 (1987).
Sexual morph. Stromata scattered to grouped. Ascomata perithecial, subglobose
to ampulliform. Ostiolar neck cylindrical, periphysate. Paraphyses numerous, un-
branched, cylindrical, hyaline. Asci unitunicate, cylindrical, with an apical annulus,
pedicellate. Ascospores obovoid to ellipsoid, muriform, hyaline to brown.
Asexual morph. Coelomycetous asexual morph: Conidiomata pycnidial, globose
to subglobose. Conidiogenous cells phialidic. Conidia ellipsoidal to obovoid, aseptate,
hyaline. Hyphomycetous synasexual morph: Colonies euse or sporodochial. Con-
idiophores micronematous, mononematous, simple or branched, hyaline, thin-walled.
Conidiogenous cells phialidic. Conidia ellipsoidal to allantoid, aseptate, hyaline.
Notes. yridiaceae has been treated as incertae sedis in Sordariomycetes (Yue
and Eriksson 1987). Members of yridiaceae dier from Myrmecridiales by hav-
ing pycnidial conidiomata, becoming cup-shaped in the coelomycetous state and
micronematous conidiophores with monophialidic conidiogenous cells in the hy-
phomycetous state. Myrmecridiales have brown thick-walled conidiophores with
polyblastic conidiogenous cells (Crous et al. 2015a). Annulatascales have relatively
massive refractive, well-developed, conspicuous apical annulus in asci (Wong et al.
1999; Campbell and Shearer 2004; Dong et al. 2021). In contrast, those of members
of yridiaceae are compact and inconspicuous. erefore, a new order, yridiales,
is introduced for this lineage.
yridiaceae J.Z. Yue & O.E. Erikss., Syst. Ascom. 6(2): 233 (1987).
Phialemoniopsidaceae K.D. Hyde & Hongsanan, [as Phialemoniopsaceae] Fungal Di-
vers. 107: 95 (2021).
Type genus. yridium Nitschke, Pyrenomyc. Germ. 1: 110 (1867).
Notes. Phialemoniopsidaceae is considered a synonym of yridiaceae because
Phialemoniopsis, the type genus of Phialemoniopsidaceae, was revealed congeneric with
yridium and is placed in the synonymy of the latter genus in this study. e type

yridium revised 157
genera of both families, that is, yridium and Phialemoniopsis, share many morpho-
logical features in their asexual states, as noted below.
yridium Nitschke, Pyrenomyc. Germ. 1: 110 (1867).
Melanospora subgen. Bivonella Sacc., Syll. fung. (Abellini) 2: 464 (1883).
Bivonella (Sacc.) Sacc., Syll. fung. (Abellini) 9: 989 (1891).
Pleurocytospora Petr., Annls mycol. 21: 256 (1923).
Sinosphaeria J.Z. Yue & O.E. Erikss., Syst. Ascom. 6: 231 (1987).
Phialemoniopsis Perdomo, Dania García, Gené, Cano & Guarro, Mycologia 105: 408
(2013).
Type species. yridium vestitum (Fr.) Fuckel, Jb. nassau.Ver. Naturk. 23–24: 195
(1870) [1869–70].
Sexual morph. Stromata scattered to grouped, subepidermal to erumpent, yel-
lowish to dark brown, red in KOH or not changing. Ascomata perithecial, subglobose
to ampulliform, single to grouped, immersed in stromata to erumpent through host
surface. Ascomatal wall composed of several layers of polygonal, dark brown cells.
Ostiolar neck cylindrical, short or long, separated or convergent in upper stromata,
periphysate. Paraphyses numerous, septate, unbranched, cylindrical, hyaline. Asci uni-
tunicate, cylindrical, broadly rounded at the apex, with a pronounced non-amyloid
apical annulus, pedicellate. Ascospores obovoid or ellipsoid, smooth, pale brown to
brown, with several transverse and 0–3 longitudinal or oblique septa.
Asexual morph. Coelomycetous and/or hyphomycetous morphs formed. Coelo-
mycetous asexual morph: Conidiomata pycnidial, single to grouped, supercial or im-
mersed in stromata, globose to subglobose, composed of polygonal to prismatic cells,
often becoming cup-shaped when mature, surrounded by setose hyphae. Conidiomatal
wall composed of several layers of polygonal, dark brown cells. Ostiolar neck cylindri-
cal, central, periphysate. Setose hyphae erect, usually unbranched, septate, cylindrical,
with slightly pointed or blunt tips, hyaline to pale brown, smooth-walled. Conidi-
ophores hyaline, thin-walled, simple or irregularly branched, with branches bearing a
small group of phialides terminally. Phialides swollen at the base, tapering at the tip,
hyaline. Conidia obovoid to oblong, with a slightly apiculate base, hyaline, smooth-
walled, in slimy masses. Hyphomycetous synasexual morph: Colonies euse or sporo-
dochial. Conidiophores micronematous, mononematous, hyaline, thin-walled, simple
or irregularly branched, with branches bearing a small group of phialides terminally.
Phialides swollen at the base, tapering at the tip, hyaline. Adelophialides absent or
rarely present. Conidia ellipsoidal to allantoid, with a slightly apiculate base, hyaline,
smooth-walled, in slimy head. Chlamydospores absent or rarely present, hyaline to
pale brown, thick- and rough-walled.
Notes. e newly obtained yridium collections formed synasexual morphs, coe-
lomycetous and hyphomycetous, in culture that were similar to those of Phialemoniop-
sis, having coelomycetous and/or hyphomycetous conidial states in culture (Perdomo

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
158
et al. 2013). In this study, Phialemoniopsis is treated as a synonym of yridium because
of their morphological similarities in asexual morphs and phylogenetic relatedness.
e genus Pleurocytospora has been proposed as a synonym of yridium by culture
studies (Leuchtmann and Müller 1986). We agree that the morphological features of
Pleurocytospora, such as phialidic conidiogenous cells and hyaline, ellipsoidal conidia
formed from both coelomycetous and hyphomycetous states (Leuchtmann and Müller
1986), are almost identical to those of the generic concept of yridium emended here.
We accept both Bivonella and Sinosphaeria as synonyms of yridium, as proposed
in previous studies (Eriksson and Yue 1989; Checa et al. 2013). Sinosphaeria (typied by
S. bambusicola = yridium chrysomallum; Yue and Eriksson 1987) was established as a
new genus without knowing the existence of Bivonella (typied by B. lycopersici; Saccardo
1891). Both genera are characterised by yellowish stromata. e validity of these genera
being synonymised under yridium is conrmed by the presence of T. avostromatum,
which has yellowish stromata, in the strongly supported yridium clade (Fig. 1).
yridium avostromatum R. Sugita & Kaz. Tanaka, sp. nov.
MycoBank No: 841917
Figs 3, 6A
Holotype. J, Yamaguchi, Nagato, Misumikami, near Kusaritoge, on dead twigs
of Phyllostachys pubescens, 26 March 2018, K. Tanaka, K. Arayama and R. Siguta, KT
3891 (HHUF 30647, holotype designated here), living culture MAFF 247509.
Etymology. e name refers to yellowish stromata.
Sexual morph. Stromata scattered to grouped, subepidermal, becoming erumpent
to supercial, 0.7–1.4 mm long, 0.4–0.7 mm wide, yellowish to dark brown, red in
2% KOH. Ascomata perithecial, subglobose to ampulliform, mostly 2–6 grouped,
190–240 µm high, 200–220 µm diam., immersed in stromata to erumpent through
host surface. Ascomatal wall 15–23 µm thick, composed of 5–8 layers of polygonal,
2.5–7 × 1.5–3.5 µm, dark brown cells. Ostiolar neck central, cylindrical, 80–140 µm
long, 55–90 µm wide, periphysate. Paraphyses numerous, septate, unbranched, cy-
lindrical, 50–105 µm long. Asci unitunicate, cylindrical, 62.5–90 × 6.5–10 µm (av.
78.7×7.8 µm, n = 30), broadly rounded at the apex, with a pronounced non-amyloid
apical annulus, short-stalked (5–17.5 µm long), with 8 ascospores. Ascospores obovoid
to ellipsoid, smooth, hyaline to pale brown, with 3 transverse and 0–2 vertical septa,
9.5–14 × 5–7.5 µm (av. 11.3 × 5.8 µm, n = 50), l/w 1.4–2.5 (av. 2.0, n = 50).
Asexual morph (nature). Not observed.
Asexual morph (culture). Hyphomycetous asexual morph formed. Conidi-
ophores micronematous, mononematous, hyaline, thin-walled, simple or irregularly
branched, with branches bearing a group of 2–3 phialides terminally. Phialides swollen
at the base, tapering at the tip, hyaline, 3–6 × 1–1.5 µm. Adelophialides rarely present.
Conidia ellipsoidal to allantoid, with a slightly apiculate base, hyaline, smooth-walled,
2–7 × 1–2.5 µm (av. 4.1 × 1.6 µm, n = 50). Chlamydospores rarely present, solitary,
3.5–6.5 µm diam., hyaline to pale brown, thick- and rough-walled.

yridium revised 159
Figure 3. yridium avostromatum (A–S KT 3891 = HHUF 30647 T–AC culture KT 3891 = MAFF
247509) A–S sexual morph A–C appearance of stromata on substrate D, E ascomata in longitudinal sec-
tion (D in 2% KOH) F ostiolar neck of ascoma G paraphyses H ascomatal wall I–K asci L apex of the as-
cus M stipe of the ascus N–R ascospores S germinating ascospore T–AC hyphomycetous asexual morph
T sporulation in culture U phialides V slimy conidial heads W conidiophores X phialide Y adelophialide
Z–AB conidia AC chlamydospores and conidia. Scale bars: 1 mm (A); 500 µm (B, C); 100 µm (D, E);
50 µm (F); 10 µm (G–K, M, S, U, V); 5 µm (L, N–R, W–AC); 250 µm (T).
Culture characteristics. Colonies on MEA at 25 °C attained 28–29 mm diam.
after a week in the dark, whitish. On OA attained 35–37 mm diam., whitish. On PDA
attained 28–31 mm diam., whitish to bu (45; Rayner 1970) (Fig. 6A).

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
160
Notes. Phylogenetic analyses based on ITS, act, and tub2 sequences suggested that
T. avostromatum was closely related to T. curvatum, T. hongokgense and T. limonesiae
(Fig. 2), of which only T. hongokgense has unknown conidial state. Although T. curva-
tum forms sporodochial conidiomata (Perdomo et al. 2013), those are not found in T.
avostromatum. Conidia of T. limonesiae (2.3–4.9 × 1.4–2 µm; Martinez et al. 2021)
are smaller than those of T. avostromatum (2–7 × 1–2.5 µm). yridium avostroma-
tum is similar to T. lasiacidis on Lasiacis ligulata (Samuels and Rogerson 1989) in 1)
having yellowish stromata becoming red in KOH, and 2) ellipsoidal ascospores with
three transverse septa, with or without one longitudinal septum in 1–2 median cells.
However, T. lasiacidis diers from T. avostromatum by ascomata with a longer ostiolar
neck (90–170 µm long) and dark brown ascospores with terminal pale brown cells
(Samuels and Rogerson 1989).
yridium pluriloculosum (Perdomo, Dania García, Gené, Cano & Guarro) R.
Sugita & Kaz. Tanaka, comb. nov.
MycoBank No: 841918
Figs 4, 6B
Basionym. Phialemoniopsis pluriloculosa Perdomo, Dania García, Gené, Cano &
Guarro, Mycologia 105: 412 (2013).
Holotype. USA, Nevada, human toe nail, D.A. Sutton, CBS H-20782, living
culture CBS 131712 = UTHSC 04–7 = FMR 11070 (not seen).
Sexual morph. Stromata scattered to grouped, pulvinate, circular to elliptical in
outline, elevated beyond bark surface forming pustules, 0.6–0.7 mm high, 0.9–1.0 mm
diam., dark brown to black. Ascomata perithecial, subglobose to ampulliform, 4–8
grouped, 700–780 µm high, 220–280 µm diam., immersed in stromata. Ascomatal
wall 17–25 µm thick, composed of 7–10 layers of polygonal, 4–6.5 × 2–4 µm, dark
brown cells. Ostiolar neck central, cylindrical, 400–430 µm long, 100–110 µm wide,
periphysate. Paraphyses septate, unbranched, cylindrical, 92.5–110 µm long, 3.5–5.5
µm wide. Asci unitunicate, cylindrical, 110–175 × 9–12.5 µm (av. 145.6 × 10.3 µm,
n = 15), broadly rounded at the apex, with a pronounced non-amyloid apical annulus,
pedicellate (12.5–27.5 µm long), with 8 ascospores. Ascospores fusiform to ellipsoid,
smooth, brown, with 3 transverse and 0–2 oblique or vertical septa, 13.5–18 × 6–8 µm
(av. 15.5 × 7.3 µm, n = 50), l/w 1.7–2.6 (av. 2.1, n = 50).
Asexual morph (nature). Conidiomata pycnidial, globose to subglobose, grouped,
220–300 µm high, 90–150 µm diam., immersed in stromata. Conidiomatal wall 8–18
µm thick, composed of 3–5 layers of polygonal, 3–4.5 × 2.5–4 µm, dark brown cells.
Ostiolar neck central, cylindrical, 80–110 µm long, 90–110 µm wide, composed of
polygonal cells, periphysate. Conidiophores hyaline, thin-walled, with branches bear-
ing a group of 2–5 phialides terminally. Phialides tapering toward the tip, hyaline,
11–16 × 1–2 µm. Conidia ellipsoidal, with a slightly apiculate base, hyaline, smooth-
walled, 3–4.5 × 1–2 µm (av. 3.7 × 1.5 µm, n = 50). Chlamydospores not observed.

yridium revised 161
Figure 4. yridium pluriloculosum (A–Y KT 3803 = HHUF 30648 Z–AL culture KT 3803 = MAFF
247508) A–R sexual morph A, B appearance of stromata on substrate (B transverse sections) C ascomata
in longitudinal section D ostiolar neck of ascoma E paraphyses F ascomatal wall G pseudostromatic tissue
H–J asci K apex of ascus L–Q ascospores R germinating ascospore S–AF coelomycetous asexual morph
(S–Y nature Z–AF culture) S appearance of conidiomata on substrate T conidiomata in longitudinal
section U conidiomatal wall V conidiophores W phialide X , Y conidia Z–AB conidiomata in culture
(AB multiloculate conidiomata) AC setose hypha of conidiomata AD conidiophores with groups of
phialides AE, AF conidia AG–AL hyphomycetous synasexual morph AG, AH sporulation in culture
AI phialide AJ, AK conidia AL chlamydospores. Scale bars: 1 mm (A, B , S , AB ); 500 µm (C , Z , AA );
100 µm (D, T ); 20 µm (AG, AH); 10 µm (E–J, L–R, U, V); 5 µm (K, W–Y, AC–AF, AI–AL).

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
162
Asexual morph (culture). Coelomycetous asexual morph: Conidiomata pyc-
nidial, scattered, single to grouped, supercial, globose to subglobose, 180–380 µm
high, mostly 80–580 µm diam., up to 1170 µm diam. when grouped, often becoming
cup-shaped when mature, surrounded by setose hyphae. Conidiomatal wall composed
of polygonal to prismatic, 3–4.5 × 2.5–4 µm, dark brown cells. Setose hyphae erect,
usually unbranched, septate, up to 360 µm long, 2–3 µm wide, pale brown. Conidi-
ophores hyaline, thin-walled, simple or irregularly branched, with branches bearing
a group of 2–5 phialides terminally. Phialides tapering toward the tip, hyaline, 10–
25×1–2.5 µm. Conidia ellipsoidal, with a slightly apiculate base, hyaline, smooth-
walled, in slimy masses, 3–4.5 × 1–2 µm (av. 3.8 × 1.4 µm, n = 50). Hyphomycetous
synasexual morph: Conidiophores micronematous, mononematous, hyaline, simple
or rarely branched. Phialides slightly tapering toward the tip, 4–11 × 1–2.5 µm, hya-
line. Adelophialide absent. Conidia allantoid, hyaline, smooth-walled, in slimy heads,
3–9 × 1–2.5 µm (av. 6.2 × 1.7 µm, n = 50). Chlamydospores rarely present, solitary,
3.5–6.5 µm diam., hyaline to pale brown, thick- and rough-walled.
Culture characteristics. Colonies on MEA at 25 °C attained 31–33 mm diam.
after a week in the dark, whitish. On OA attained 32–36 mm diam., whitish to grey
olivaceous (107). On PDA attained 32–33 mm diam., whitish to bu (45) (Fig. 6B).
Specimen examined. J, Aomori, Hirakawa, Hirofune, Shigabo Forest Park,
on dead twigs of Betula maximowicziana, 10 October 2017, K. Tanaka, KT 3803
(HHUF 30648), living culture MAFF 247508.
Notes. e conidia from aerial hyphae of strain KT 3803 were larger (3–9 × 1–2.5
µm) in culture than those of the original description of yridium pluriloculosum
(3–5×1–2.5 µm; Perdomo et al. 2013). However, we identied this new collection
on Betula maximowicziana as T. pluriloculosum, based on the high sequence homology
of three loci with ex-type culture of this species (CBS 131712; 99.6% in ITS, 99.2%
in act, and 99.5% in tub2). e sexual-asexual relationship of T. pluriloculosum was
veried in this study. Although this species has been reported from clinical sources as
an asexual morph (Perdomo et al. 2013), the recently collected material represents a
sexual morph on plant material.
In yridium, T. betulae has also been recorded on Betula sp. in France (Roume-
guère 1891). Although sequences of T. betulae are unavailable for molecular compari-
son, it is clearly dierent from T. pluriloculosum in having ascospores with 5–7 trans-
verse and one longitudinal septum.
yridium punctulatum (I. Hino & Katum.) R. Sugita & Kaz. Tanaka, comb. nov.
MycoBank No: 841919
Figs 5, 6C
Basionym. Pleospora punctulata I. Hino & Katum., Icones Fungorum Bamb. Jpn.: 181
(1961).
Holotype. J, Shizuoka, Fuji Bamboo Garden, on dead twigs of Phyllostachys
nigra var. henonis, 1 April 1958, K. Katumoto, YAM 21851.

yridium revised 163
Epitype. J, Yamaguchi, Hagi, Akiragi, near Chikurindoro-park, on dead
twigs of Phyllostachys nigra var. nigra, 26 March 2018, K. Tanaka, K. Arayama and
R. Sugita, KT 3905 (HHUF 30649 epitype designated here; MBT 10004137), ex-
epitype culture MAFF 247510.
Sexual morph. Stromata scattered to grouped, subepidermal, becoming erumpent
to supercial, 0.5–1.2 mm long, 0.2–0.4 mm wide, dark brown. Ascomata perithecial,
subglobose to conical, single to 2–3 grouped, 130–190 µm high, 140–230 µm diam.,
immersed in stromata to erumpent through host surface. Ascomatal wall 7–15 µm thick,
composed of 3–5 layers of polygonal, 3–6.5 × 1–4.5 µm, dark brown cells. Ostiolar neck
central, cylindrical, 37–85 µm long, 37–63 µm wide, periphysate. Paraphyses numerous,
septate, unbranched, cylindrical, hyaline, 77–103 µm long. Asci unitunicate, cylindrical,
67.5–105 × 7.5–11.5 µm (av. 82.9 × 9.4 µm, n = 60), broadly rounded at the apex, with
a pronounced non-amyloid apical annulus, short-stalked (3.5–11.5 µm long), with 8 as-
cospores. Ascospores ellipsoid to oblong, smooth, pale brown, with 3 transverse and 1–2
vertical septa, 10–15 × 5–9 µm (av. 12.8 × 7.0 µm, n = 60), l/w 1.4–2.4 (av. 1.8, n = 60).
Asexual morph (nature). Not observed.
Asexual morph (culture). Coelomycetous asexual morph: Conidiomata pycnidi-
al, single to grouped, supercial, globose to subglobose, 100–250 µm high, 170–620
µm diam., composed of polygonal to prismatic, 3.5–7.5 × 2.5–4 µm cells, often be-
coming cup-shaped when mature, surrounded by setose hyphae. Setose hyphae erect,
usually unbranched, septate, up to 225 µm long, 1.5–2.5 µm wide, pale brown. Con-
idiophores hyaline, thin-walled, simple or irregularly branched, with branches bearing
a group of 2–5 phialides terminally. Phialides swollen at the base, tapering at the tip,
7–20 × 1–3 µm, hyaline. Conidia ellipsoidal to obovoid, with a slightly apiculate base,
hyaline, smooth-walled, in slimy masses, 2–3.5 × 1–2 µm (av. 2.9 × 1.4 µm, n = 50).
Hyphomycetous synasexual morph: Conidiophores micronematous, mononematous,
hyaline, thin-walled, simple or irregularly branched, with branches bearing a group of
2–3 phialides terminally. Phialides swollen at the base, tapering at the tip, hyaline, 3–9
× 1–2 µm. Adelophialide absent. Conidia ellipsoidal to allantoid, hyaline, smooth-
walled, in slimy heads, 2.5–8 × 1–3 µm (av. 4.3 × 1.6 µm, n = 87). Chlamydospores
rarely present, solitary or chained, 4–5.5µm diam., hyaline to pale brown.
Culture characteristics. Colonies on MEA at 25 °C attained 31–32 mm diam.
after a week in the dark, granulose, whitish. On OA attained 38–39 mm diam., granu-
lose, whitish. On PDA attained 35–36 mm diam., whitish to bu (45) (Fig. 6C).
Other specimen examined. J, Iwate, Morioka, Ueda, Campus of Iwate Uni-
versity, on dead culms of Phyllostachys pubescens, 17 February 2003, K. Tanaka and Y.
Harada, KT 1015 (HHUF 29350), living culture JCM 13159=MAFF 239669.
Notes. is species has been described from Phyllostachys nigra var. henonis, as a spe-
cies of Pleospora (Dothideomycetes; Hino 1961). Our phylogenetic analysis (Fig. 1) shows
that this species is a member of the genus yridium (Sordariomycetes). e morphologi-
cal features of this species are consistent with those of the genus yridium, including
immersed to erumpent, single to grouped, perithecial ascomata with a cylindrical ostiolar
neck, unitunicate asci and muriform, pigmented ascospores (Eriksson and Yue 1989).
erefore, we propose a new combination, T. punctulatum, for Pleospora punctulata.

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
164
Figure 5. yridium punctulatum (A–N, Q, R KT 3905 = HHUF 30649 O, P YAM 21851
S, T, W–AB culture KT 1015 = JCM 13159 = MAFF 239669 U, V, AC–AK culture KT 3905 = MAFF
247510) A–R sexual morph A, B appearance of stromata on substrate C, D ascomata in longitudi-
nal section E ostiolar neck of ascoma F paraphyses G ascomatal wall H–J asci K apex of ascus L stipe
of ascus M–Q ascospores R germinating ascospore S–AD coelomycetous asexual morph S–V conidi-
omata in culture W conidioma in longitudinal section X conidiomatal wall Y setose hyphae of conidi-
omata Z, AA conidiophores AB phialides AC , A D conidia AE–AK hyphomycetous synasexual morph
AE conidiophore AF slimy head AG phialide AH–AJ conidia AK chlamydospores. Scale bars: 1 mm
(A, S); 500 µm (B); 100 µm (C , W ); 50 µm (D); 10 µm (E–J, L, R, X–AA, AE, AF); 5 µm (K, M–Q,
AB–AD, AG–AK); 200 µm (T–V).

yridium revised 165
yridium cornearis (Perdomo, Dania García, Gené, Cano & Guarro) R. Sugita
& Kaz. Tanaka, comb. nov.
MycoBank No: 841920
Basionym. Phialemoniopsis cornearis Perdomo, Dania García, Gené, Cano & Guarro,
Mycologia 105: 408 (2013).
yridium curvatum (W. Gams & W.B. Cooke) R. Sugita & Kaz. Tanaka, comb. nov.
MycoBank No: 841921
Phialemoniopsis curvata (W. Gams & W.B. Cooke) Perdomo, Dania García, Gené,
Cano & Guarro, Mycologia 105: 410 (2013).
Basionym. Phialemonium curvatum W. Gams & W.B. Cooke, Mycologia 75: 980 (1983).
yridium endophyticum (Lei Su & Y.C. Niu) R. Sugita & Kaz. Tanaka, comb. nov.
MycoBank No: 841922
Basionym. Phialemoniopsis endophytica Lei Su & Y.C. Niu, Mycol. Progr. 15: 3 (2016).
yridium hongkongense (Tsang, Chan, Ip, Ngan, Chen, Lau, Woo) R. Sugita &
Kaz. Tanaka, comb. nov.
MycoBank No: 841923
Basionym. Phialemoniopsis hongkongensis Tsang, Chan, Ip, Ngan, Chen, Lau, Woo, J.
Clin. Microbiol. 52: 3284 (2014).
Figure 6. Colony characters of yridium species used in this study on MEA (bottom right), OA
(bottom left) and PDA (upper) within 1 week at 25 °C in the dark A T. avostromatum (culture KT
3891=MAFF 247509) B T. pluriloculosum (culture KT 3803 = MAFF 247508) C T. punctulatum (cul-
ture KT 3905=MAFF 247510). Scale bars: 3 cm (A–C).

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
166
yridium limonesiae (A. Riat, L.W. Hou & Crous) R. Sugita & Kaz. Tanaka,
comb. nov.
MycoBank No: 841927
Basionym. Phialemoniopsis limonesiae A. Riat, L.W. Hou & Crous, Emerging Mi-
crobes & Infections 10: 403 (2021).
yridium oculorum (Gené & Guarro) R. Sugita & Kaz. Tanaka, comb. nov.
MycoBank No: 841924
Phialemoniopsis ocularis (Gené & Guarro) Perdomo, Dania García, Gené, Cano &
Guarro, Mycologia 105: 411 (2013).
Basionym. Sarcopodium oculorum Gené & Guarro, J. Clin. Microbiol. 40: 3074 (2002).
Discussion
We show that the asexual genus Phialemoniopsis (established by Perdomo et al. 2013)
is a synonym of the sexual genus yridium (established by Nitschke 1867). We found
a new species of yridium (T. avostromatum), transferred Pleospora punctulata into
yridium, and proposed seven new combinations in yridium for strains previously
treated in Phialemoniopsis. We provided a revised generic circumscription of yridium
based on both sexual and asexual characteristics and revealed the phylogenetic relation-
ships of species within this genus.
e genus yridium has been dened mainly on the basis of sexual characters
(Nitschke 1867; Eriksson and Yue 1989). Currently, 33 species are recorded in this
genus (http://www.indexfungorum.org, 2021). Asexual morphs are unknown in most
species of yridium, with the exceptions of T. avum and T. vestitum, in which asexual
morphs have been recorded based on sexual-asexual association on the same specimen
(Petch 1917) and on the basis of culture study (Leuchtmann and Müller 1986, this
study), respectively. In contrast, the genus Phialemoniopsis has been dened based only
on asexual characters (Perdomo et al. 2013). Its ordinal aliation within Sordariomy-
cetes has not been resolved, but recent phylogenetic analyses of this class suggest that
Phialemoniopsis is close to yridium (Hyde et al. 2021). In our phylogenetic analysis,
all species previously described as Phialemoniopsis (marked with blue circle; Fig. 1)
were clustered in a single clade, including the type species of yridium (T. vestitum),
as well as two new strains proposed here (T. avostromatum and T. punctulatum). Both
genera have similar asexual morphs, which have conidiophores bearing small groups
of phialides, hyaline phialidic conidiogenous cells, and ellipsoidal or allantoid, hyaline
conidia in both coelomycetous and hyphomycetous states (Petch 1917; Leuchtmann
and Müller 1986; Perdomo et al. 2013). Morphological and molecular phylogenetic
evidence clearly shows that Phialemoniopsis is congeneric with yridium.

yridium revised 167
Synonymising Phialemoniopsis under yridium expanded information about the
asexual morphs of yridium. In this genus, only T. vestitum has been demonstrated
to have asexual morphs by culture studies (Leuchtmann and Müller 1986). It has both
coelomycetous and hyphomycetous complex asexual morphs, which have phialidic
conidiogenous cells with collarette and ellipsoidal to allantoid hyaline conidia (Leucht-
mann and Müller 1986). Members of Phialemoniopsis also have coelomycetous and/or
hyphomycetous conidial states (Perdomo et al. 2013; Tsang et al. 2014; Su et al. 2016;
Martinez et al. 2021). e close relationship of Phialemoniopsis and yridium suggests
that such complex asexual morphs may be common within yridium species.
In yridium, T. endophyticum and T. curvatum have been isolated from both plants
and animals (Gam and McGinnis 1983; Halleen et al. 2007; Perdomo et al. 2013; Su
et al. 2016; Ito et al. 2017). ere are several examples of fungal species, including
human pathogens, detected from various substrates. For example, Phaeoacremonium
minimum is a pathogen on grapevines, where it forms both sexual and asexual morphs
(Crous et al. 1996; Pascoe et al. 2004), but it has also been reported as a causative agent
of subcutaneous phaeohyphomycosis in humans as asexual morph (Choi et al. 2011).
Other species of yridium may also have cryptic life cycles and can colonise each host
substrate at dierent reproductive stages. An example of this prediction can be found
in T. pluriloculosum. is species was originally found in human nails as an asexual
fungus (Perdomo et al. 2013), and its sexual state was rediscovered on twigs of Betula
maximowicziana in our study.
Epitypication of the type species of yridium (T. vestitum) will be a necessary
issue in the future. We used sequences from two non-type strains (CBS 113027, CBS
125582) of this species for phylogenetic analyses but they did not form a monophy-
letic clade (Fig. 1). Sequence dierences between these two strains were found at 34
positions with four gaps in the LSU. ese results indicate that the strains obtained
from Acer pseudoplatanus (CBS 113027) and no host information (CBS 125582) in
Austria are not conspecic. A fresh collection of T. vestitum on original host plant from
the type locality (Ribes rubrum, Sweden; Fries 1823) and its phylogenetic analysis are
required to x generic circumscription of yridium.
yridiales established here may encompass other genera and families with mor-
phologies distinct from the genus yridium (yridiaceae). Some species of “Linocar-
pon” and “Neolinocarpon” are nested within the yridiales (Fig. 1). Linocarpon and
Neolinocarpon sensu stricto belong to Linocarpaceae (Chaetosphaeriales) and are mor-
phologically distinct from yridium in having liform, straight or curved, unicellular,
hyaline, or pale-yellowish ascospores (Huhndorf and Miller 2011; Konta et al. 2017).
e “Linocarpon” and “Neolinocarpon” species phylogenetically unrelated to Linocarpon
and Neolinocarpon sensu stricto may be new lineages in yridiaceae or belong to its own
new undescribed family. However, we cannot clarify the phylogenetic/taxonomic relat-
edness of these atypical Linocarpon-like species because none of them are ex-types and
their morphological information are unavailable. Further molecular phylogenetic study
of these fungi based on protein-coding sequences and nding additional specimens/iso-
lates of “Linocarpon” and “Neolinocarpon” species related to yridium will be necessary
to clarify their taxonomic aliation and better understand the concept of yridiales.

Ryosuke Sugita & Kazuaki Tanaka / MycoKeys 86: 147–176 (2022)
168
Acknowledgments
We gratefully acknowledge Y. Harada and K. Arayama for their help with the collec-
tion of fungal specimens. We thank the curator of YAM, S. Ito, who permitted us to
examine type collection. is work was partially supported by grants from the Japan
Society for the Promotion of Science (JSPS 19K06802).
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