Uncovering the hidden taxonomic diversity of fungi in Oman

Abstract

Hot desert regions are undoubtedly challenging to fungal survival on the Arabian Peninsula. Fungi are, however, recognized as the most stress-resistant organisms among all eukaryotes, which could be a result of the rapid evolution of distinct species. Our current understanding of these microorganisms is derived from studies examining only a fraction of the overall fungal diversity. Therefore, further studies are needed to understand the diversity of fungi in desert regions. This paper highlights the taxonomy of several unusual fungal genera collected in a range of aquatic and terrestrial environments in Oman. These taxa were identified based on phylogenetic analyses of nuclear ribosomal DNA (rDNA) (LSU, SSU and ITS) and protein-coding genes (TEF, RPB2 and TUB), plus morphological comparisons. Phylogenetic analyses, including the presently recognized genera in Patellariales resulted in splitting the order into four clades in Dothideomycetes. The new family Holmiellaceae and new order Holmiellales are introduced to include Holmiella. Species of Homortomycetaceae form a well-supported and distinct clade and raise it to Homortomycetales ordo novus. Omania (Halojulellaceae), Desertiserpentica (Lophiostomataceae) and Montanitestudina (Testudinaceae) are described as novel genera in Dothideomycetes. A synnematous hyphomycete with basidiomycetous affinity (Corticiales) was also identified and described as Basidiodesertica. Additionally, an asexual morph was observed for Holmiella junipericola; Patellaria quercus is synonymized under P. atrata, and seven new species are described.

Full text

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Fungal Diversity
https://doi.org/10.1007/s13225-020-00467-1
Uncovering thehidden taxonomic diversity offungi inOman
SajeewaS.N.Maharachchikumbura1,3· DhanushkaN.Wanasinghe2· RatchadawanCheewangkoon3·
AbdullahM.Al‑Sadi4
Received: 5 October 2020 / Accepted: 21 December 2020
© MUSHROOM RESEARCH FOUNDATION 2021
Abstract
Hot desert regions are undoubtedly challenging to fungal survival on the Arabian Peninsula. Fungi are, however, recognized
as the most stress-resistant organisms among all eukaryotes, which could be a result of the rapid evolution of distinct species.
Our current understanding of these microorganisms is derived from studies examining only a fraction of the overall fungal
diversity. Therefore, further studies are needed to understand the diversity of fungi in desert regions. This paper highlights the
taxonomy of several unusual fungal genera collected in a range of aquatic and terrestrial environments in Oman. These taxa
were identified based on phylogenetic analyses of nuclear ribosomal DNA (rDNA) (LSU, SSU and ITS) and protein-coding
genes (TEF, RPB2 and TUB), plus morphological comparisons. Phylogenetic analyses, including the presently recognized
genera in Patellariales resulted in splitting the order into four clades in Dothideomycetes. The new family Holmiellaceae
and new order Holmiellales are introduced to include Holmiella. Species of Homortomycetaceae form a well-supported and
distinct clade and raise it to Homortomycetales ordo novus. Omania (Halojulellaceae), Desertiserpentica (Lophiostomata-
ceae) and Montanitestudina (Testudinaceae) are described as novel genera in Dothideomycetes. A synnematous hyphomycete
with basidiomycetous affinity (Corticiales) was also identified and described as Basidiodesertica. Additionally, an asexual
morph was observed for Holmiella junipericola; Patellaria quercus is synonymized under P. atrata, and seven new species
are described.
Keywords 14 new taxa· Ascomycota· Basidiomycota· Desert· Dothideomycetes· Patellariales· Saprobes
Introduction
Fungi are a diverse, economically and ecologically important
group of microorganisms (Hunter-Cevera 1998). They are
second only to insects in the number of species thought to
exist (Hawksworth 2001). Based on various algorithms 2.2
and 3.8 million fungal species are estimated (Hawksworth
and Lücking 2017) with the latest estimate being 11.7–13.2
million species (Wu etal. 2019; Hyde etal. 2020a). How-
ever, less than 100,000 species or only between 2.6 and
4.5% of the estimated species of fungi have been described
(Hawksworth and Lücking 2017; Hyde etal. 2020b). This
indicates that there are numerous gaps in describing fun-
gal diversity and the exact number of global species remain
to be described. These gaps in the understanding of fungal
communities limit our ability to recognize fungal diversity
(Crous etal. 2015). The advent of modern molecular tech-
niques has facilitated the rapid discovery of numerous taxo-
nomic novelties in fungi from various habitats (Tekpinar
and Kalmer 2019). However, these studies have examined
This article is dedicated to Prof. Kevin David Hyde on the
occasion of his 65th birthday.
Supplementary Information The online version contains
supplementary material available at https ://doi.org/10.1007/s1322
5-020-00467 -1.
* Sajeewa S. N. Maharachchikumbura
sajeewa83@yahoo.com
* Abdullah M. Al-Sadi
alsadi@squ.edu.om
1 School ofLife Science andTechnology, University
ofElectronic Science andTechnology ofChina,
Chengdu611731, People’sRepublicofChina
2 CAS Key Laboratory forPlant Diversity andBiogeography
ofEast Asia, Kunming Institute ofBotany, Chinese
Academy ofScience, Kunming650201, Yunnan,
People’sRepublicofChina
3 Entomology andPlant Pathology Department, Faculty
ofAgriculture, Chiang Mai University, ChiangMai, Thailand
4 Department ofCrop Sciences, College ofAgricultural
andMarine Sciences, Sultan Qaboos University,
Al-Khod123, Oman

Fungal Diversity
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a minute fraction of the overall diversity, and the identifi-
cations are mostly made at the generic level, meaning the
great majority of the data are uninformative. Accounting for
biodiversity from various habitats is the basis for sustain-
able management of natural resources in the face of ongoing
global change (Reid and Miller 1989). The susceptibility of
fungal communities to natural perturbations remains rela-
tively poorly studied, particularly in extreme environments
such as deserts (Magan 2007; Maharachchikumbura etal.
2016). Therefore, further studies are needed to understand
the morpho-phylo diversity of fungi and their ecological
background in desert regions.
Oman is one of the oldest independent Arabian states in
the world. It occupies the south-eastern coast of the Arabian
Peninsula at the joining of the Persian Gulf and Arabian Sea
(Henderson 2015). It has a land area of 309,500 square km
with the coastline extending approximately 3000km from
the Strait of Hormuz in the north to the borders of Yemen
in the south (Maharachchikumbura etal. 2016). Oman has
the highest biological diversity in the entire Arabian Penin-
sula and the country includes 82% valleys and desert, 15%
mountain ranges and 3% coastal areas (Fouda etal. 1997;
Carranza etal. 2018). It has a subtropical dry, hot climate
with low rainfall. The temperature is very high in the sum-
mer and there are large differences between the maximum
and minimum temperatures, especially in the inland areas
(Al-Charaabi and Al-Yahyai 2013). Oman is a large country
and its climate conditions change according to geography.
October to April is warm (37°C), sunny and pleasant dur-
ing the day and about 17°C at night (Alsharhan etal. 2001).
May to August is dry and hot. The majority of inner Oman
falls within the sandy, treeless, and largely waterless region
of the Arabian Peninsula known as the Rub’ al-Khali (Edgell
2006).
The Rub’ al-Khali is the world’s largest area of continu-
ous sand and one of the driest regions in the world (Almaz-
roui etal. 2012). Therefore, plant diversity is minimal and
consists of very diffuse shrub communities. Compared to the
interior of Oman, the coastline has much more plant life (El-
Sheikh 2013). The Dhofar region in the southern coastline
of Oman is the most fertile area in the country and during
May to September has its own microclimate with monsoons
from the Indian ocean with rain and lower temperatures than
the rest of the country (Nizamuddin and Campbell 1995).
The monsoon climate brings up constantly flowing streams
and the Dhofar region is rich with temperate vegetation and
crops such as coconuts, alfalfa, sorghum, bananas, and veg-
etables (Sale 1980; Jayasuriya etal. 2017). The area in the
Batinah coastal plain has fertile land with alluvial deposits
coming from the Al Hajar mountains (Abdelrahman etal.
1993). The soil has high organic matter content, high nutri-
ent levels and a slow formation rate, providing good condi-
tions for plant growth and is thus the principal agricultural
region in Oman, producing approx. 70% of the country’s
agricultural crops (Abdelrahman etal. 1993).
In addition to Batinah, the Al Hajar Mountain range
(“The Stone Mountains”, “Oman Mountains”) in north-
eastern Oman are the highest mountain range in the eastern
Arabian Peninsula. These mountains are rich in biodiversity
and have higher numbers of endemic flora and fauna com-
pared to most parts of the Arabian Peninsula (Gebauer etal.
2009). The cool mountain air and more significant rainfall
encourage abundant plant growth resulting in woodlands
and agricultural products including pomegranates, walnuts,
apricots, black grapes and peaches (Gebauer etal. 2009).
Moreover, these mountain ranges are sources for the Omani
Falaj systems, which are a sophisticated irrigation system
in the country dating from 500 AD (Cremaschi etal. 2018).
These irrigation systems nourish almost every village, town
and city in Oman and five of the Omani Falaj systems are
listed as UNESCO World Heritage Sites (Sulaimani etal.
2007). Extensive sand dunes are associated with the coastal
areas and mangrove forests that encompass the country’s
wetlands (Scott etal. 1995).
Topographic habitat heterogeneity and various factors
increase and influence the biodiversity and number of taxo-
nomic groups in Oman, compared to other parts of the Ara-
bian Peninsula (Ahmed and Choudri 2012). Further, Oman
has a unique plant community due to being separated from
the continents of Africa and Asia for over 1000years. Biota
in southern Oman are African in origin and in northern
Oman they are of Iranian and southwest Pakistan origin,
and these two regions are separated from the central desert
of Oman (Pickering and Patzelt 2008). Oman accounts for
> 1240 vascular plant species, which includes high lev-
els of endemism (Patzelt etal. 2014). Based on this high
level of plant diversity, it is estimated that approximately
6,000–7,200 fungal species may exist in Oman (Maharach-
chikumbura etal. 2016) based on Hawksworth (1991) who
suggested a ratio of six fungal taxa to each plant host. Nev-
ertheless, according to the checklist provided by Maharach-
chikumbura etal. (2016), there are only about 320 known
species from 173 genera in Ascomycota, Basidiomycota,
Glomeromycota and Zygomycota. This indicates that only
4–5% of fungal species have been discovered and 95–96%
are potentially unexplored. This has been demonstrated in
northern Thailand, where extensive studies showed that up
to 96% of species in common genera such as Agaricus L. and
Amanita Pers. are new to science (Hyde etal. 2018).
Soil and phytopathogenic fungi are presently the best-
known groups in Oman and otherwise little is known
about fungal diversity, ecology and their place in the
evolutionary history of fungal taxa (Maharachchikum-
bura etal. 2016). Therefore, further studies are needed to
establish the actual number of fungal species in Oman and
understand their ecological background. We are engaged

Fungal Diversity
1 3
in research to characterise and conserve the fungal biodi-
versity and therefore initiated studies on saprobic fungi
occurring throughout the different ecological niches in
Oman. To elucidate the phylogenetic relationships of
these isolates, we analysed DNA sequences representing
rRNA and protein-coding genes. Herein we provide for-
mal descriptions for the new taxa, with diagnostic mor-
phological characters and genetic information.
Materials andmethods
Isolation andidentification
Dead leaves, wood and bark samples were collected during
2016 to 2018 in Al Jabal al-Akhdar (Fig.1g, h), Muscat,
Nizwa, Salalah and Sur in Oman. The samples were col-
lected in paper bags and transported to the laboratory, and
macroscopic and microscopic traits were observed. Macro-
morphological characters were photographed using a Carl
Zeiss Stemi 508 stereo microscope. Freehand sections of
Fig. 1 Habitats types in Oman. a, b Desert area. c, d Desert plants e Date palm oasis. f–h Mountain range. i–k Salalah region during monsoon. l
Coastal

Fungal Diversity
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fungal structures were made wherever necessary and
mounted in water for microscopic study. Morphological
characters were visualized with a Nikon Eclipse Ni-U micro-
scope. All microphotographs were arranged using Adobe
Photoshop CC 2018, and all measurements were made
with NIS-element D software. Specimens were preserved
and deposited at the Sultan Qaboos University Herbarium.
Single spore isolations were made from spore suspensions
on 2% potato dextrose agar (PDA) (Chomnunti etal. 2014)
and incubated at 25°C. Living cultures are maintained in
the Sultan Qaboos University culture collection (SQUCC).
Molecular laboratory work
Genomic DNA was extracted from fungal colonies growing
on PDA at 25°C for 1–2 week following a modification
of the method of Al-Sadi etal. (2012). We attempted to
obtain partial sequences of five loci for all the novel Doth-
ideomycetes species reported in this study. This included
nuclear large subunit ribosomal RNA (LSU), small subunit
RNA (SSU), the internal transcribed spacer region includ-
ing the 5.8S rDNA (ITS), the translation elongation factor
1-α (TEF), the second largest subunit of RNA polymerase II
(RPB2) and β-tubulin II (TUB). Regions of the LSU, SSU,
ITS, TEF, RPB2 and TUB were respectively amplified using
the primer pairs LR0R and LR5, ITS5 and ITS4, NS1 and
NS4, EF1 983F and EF1-2218R, RPB2-5f and RPB2-7cr
and BT2a and BT2b (Vilgalys and Hester 1990; White etal.
1990; Glass and Donaldson 1995; O’Donnell and Cigelnik
1997; Liu etal. 1999; Rehner 2001). PCR for each gene
was performed with 25μl PCR mixture using PuRe-Taq™
Ready-To-Go™ PCR beads (GE Healthcare, Buckingham-
shire, UK), 1μl of each primer (0.4mM), 1μl DNA and
22μl sterile distilled water. Amplifications were carried
out in an Applied Biosystems ProFlex PCR System (Life
Technologies, USA) with the profile detailed in previous
studies (Wanasinghe etal. 2017; Halo etal. 2019; Al-Jaradi
etal. 2020). The PCR products were visualized on a 1%
agarose electrophoresis gel stained with ethidium bromide.
PCR products were purified and sequenced at Macrogen Inc.
(South Korea).
Sequence alignment andphylogenetic analysis
Sequences were assembled with DNASTAR Lasergene
SeqMan Pro v. 8.1.3 and deposited at GenBank under the
accession numbers listed in Supplementary Tables1–9.
The sequences generated in this study were supplemented
by additional sequences obtained from GenBank (based on
BLAST searches) and the literature, including the latest out-
line of Dothideomycetes by Hongsanan etal. (2020a). Mul-
tiple sequence alignments were produced in MEGA v.7.0.26
(Kumar etal. 2016) and edited manually.
Phylogenetic analyses were carried out using maximum
likelihood (ML) and Bayesian inference (BI). The RAxML
analyses were run on the CIPRES Science Gateway portal
(Miller etal. 2012). ML analyses for the datasets were per-
formed with RAxML-HPC2 on XSEDE v. 8.2.10 (Stama-
takis 2014) using a GTR + GAMMA substitution model
with 1,000 bootstrap iterations. Branches with bootstrap
(MLB) values of 60–74% were considered to have low sup-
port, branches with MLB values of 75–89% were regarded
to have moderate support, and branches with MLB values
≥ 90% were regarded as strongly supported. Suitable models
for the Bayesian analysis were first selected using models of
nucleotide substitution for each gene, as determined using
MrModeltest v. 2.2 (Nylander 2004) and included for each
gene partition. The BI (MrBayes v. 3.2.1; Nylander, 2004)
was carried out as detailed in Wanasinghe etal. (2020) and
Samarakoon etal. (2020a). Branches with posterior prob-
ability (PP) values of 0.95–0.99 were regarded to have
moderate support, and branches with pp values of 1.00 were
considered as strongly supported. The resulting trees were
visualized with MEGA v.7.0.26 (Kumar etal. 2016) and the
layout was made with Adobe Illustrator CC 2018. The maxi-
mum clade credibility tree of orders in Dothideomycetes was
obtained from a Bayesian approach using BEAST v 1.8.4
(Drummond etal. 2012) to provide additional evidence to
support the new orders (Supplementary Fig.1).
Results
Phylogenetic relationships
Dothideomycetes SSU, 5.8S, LSU, RPB2 andTEF phylogeny
The alignment contained 142 isolates, and the tree was
rooted to Graphostroma platystomum (Schwein.) Piroz.
(CBS 270.87), Diatrype disciformis (Hoffm.) Fr. (AFTOL
-ID 927) and Sordaria fimicola (Roberge ex Desm.) Ces. &
De Not. (AFTOL -ID 216). The final alignment contained a
total of 4,022 characters used for the phylogenetic analyses,
including alignment gaps. The RAxML analysis of the com-
bined dataset yielded the best scoring tree with a final ML
optimization likelihood value of − 80,561.727970. Based on
the results of MrModelTest, dirichlet base frequencies and
the GTR+I+G model were used for the Bayesian analysis.
The Bayesian analyses generated 5,899 trees (saved every
1,000 generations) from which 4,199 were sampled after
25% of the trees were discarded as burn-in.
Notes—Sequences of SSU, 5.8S, LSU, RPB2 and TEF
were used to show the phylogenetic placement of currently
recognized genera of Patellariales within Dothideomy-
cetes (Fig.2). The sequence data of available genera of
Patellariaceae were included in the phylogenetic analysis.

Fungal Diversity
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Jahnula aquatica R68-1
Banhegyia cf. setispora G.M. 2015-04-29
Mytilinidion resinicola CBS 304.34
Flavobathelium epiphyllum MPN67
Muyocopron dipterocarpi MFLUCC 14-1103
Valsaria spartii CBS 139070
Abrothallus buellianus SPO303
Asterodiscus tamaricis CBS 136919
Eremithallus costaricensis F Luecking 15683
Rhexothecium globosum CBS 955.73
Aliquandostipite khaoyaiensis CBS 118232
Dyfrolomyces thamplaensis MFLUCC 15-0635
Inocyclus angularis VIC 39747
Parmularia styracis VIC 42587
Muyocopron laterale FMR13797
Strigula jamesii MPN548
Pirozynskiella laurisilvatica CBS 138109 (Asterinales)
Myrmaecium fulvopruinatum CBS 139058
Minutisphaera aspera DSM 29478
Dyfrolomyces maolanensis GZCC 16-0102
Hysterobrevium smilacis CBS 114601
Abrothallus acetabuli SPO308
Minutisphaera japonica KTC 2738
Mytilinidion scolecosporum CBS 305.34
Abrothallus hypotrachynae SPO302
Trypethelium virens AFTOL-ID 1774
Muyocopron atromaculans BPI GB1369
Lichenoconium aeruginosum CBS 129239
Muyocopron lithocarpi MFLUCC 14-1106
Clypeococcum psoromatis Ertz 19259
Parmularia styracis VIC 42450
Acrospermum longisporium MFLU 17-2849
Asterina cynometrae MFLU 13-0373
Stigmatodiscus enigmaticus CBS 132036
Trypetheliopsis kalbii MPN243
Arthogrphis longispora CBS 135935
Rhytidhysteron thailandicum MFLUCC 14-0503
Banhegyia setispora G.M. 2014-05-25.1
Cladoriella eucalypti CPC 10953
Rhytidhysteron neorufulum MFLUCC 13-0216
Phyllobathelium anomalum MPN242
Acrospermum gramineum M152
Encephalographa elisae EB 0347
Brachiosphaera tropicalis SS2523
Lichenoconium erodens CBS 128704
Rhizodiscina lignyota G.M. 2016-10-12.No1631
Acrospermum adeanum M133
Muritestudina chiangraiensis MFLUCC 17-2551
Acrospermum compressum M151
Hysterium angustatum CBS 236.34
Megalotremis verrucosa MPN104
Polycoccum vermicularium Diederich 17545
Rhizodiscina lignyota G.M. 2017-01-12.1
Phaeoseptum terricola MFLUCC 10-0102
Murilentithecium clematidis MFLUCC 14-0562
Leptosphaeria doliolum CBS 505.75
Pleospora herbarum CBS 191.86
Myrmaecium rubrum CBS 109505
Parmularia styracis VIC 42447
Asterina phenacis TH 589
Dyfrolomyces sinensis MFLUCC 17-1344
Cladoriella kinglakensis CPC 32730
Asterina weinmanniae TH 592
Valsaria insitiva CBS 127882
Arthogrphis kalrae CBS 693.77
Alloarthopyrenia italica MFLU 15-0399
Asterotexis cucurbitacearum VIC 42814
93/-
*
*
*
97/0.95
*
*
60/0.98
98/0.90
95/1.0
*
91/0.96
73/-
*
*
*
*
99/1.0
61/-
*
*
*
*
74/0.99
*
99/1.0
95/-
99/0.96
*
68/0.99
86/0.99
*
91/1.0
*
*
*
95/-
91/-
*
98/1.0
*
88/1.0
91/-
*
98/0.95
98/1.0
*
Asterinales
Parmulariales
Clade 1
Clade 2
-/0.95
-/1.0
*
-/1.0
Cladoriellales
Abrothallales
Lichenoconiales
Jahnulales
Eremithallales
Pleosporales
Hysteriales
Mytilinidiales
Strigulales
Monoblastiales
Dyfrolomycetales
Stigmatodiscales
Muyocopronales
Acrospermales
Eremomycetales
Trypetheliales
Valsariales
Minutisphaerales
2x
2x
2x
3x
2x
2x
3x
2x
2x
Fig. 2 Phylogenetic tree inferred using the combined sequences of
SSU, 5.8S, LSU, RPB2 and TEF of the analysed Dothideomycetes
orders. The tree backbone was constructed using maximum likeli-
hood (ML) analysis. Orders are indicated in coloured blocks. RAxML
bootstrap support values (MLB) and Bayesian posterior probabilities
(PP) above 60% and 0.95, respectively, are given at the nodes (MLB/
PP). An asterisk (*) indicates branches with MLB = 100% and PP val-
ues = 1.0. Species are followed by the strain accession numbers. The
order name is indicated to the right of the clade. The new strains are
indicated in blue bold. The scale bar represents the expected number
of changes per site for ML

Fungal Diversity
1 3
0.2
Aureobasidium lini CBS 125.21
Neodactylaria obpyriformis CBS 142668
Glyphium elatum EB 0365
Kirschsteiniothelia rostrata MFLUCC 15-0619
Fusicladium africanum CPC 12828
Botryobambusa fusicoccum MFLUCC 11-0143
Collemopsidium cf. foveolatum RO27
Sordaria fimicola AFTOL-ID 216
Superstratomyces albomucosus CBS 140270
Septorioides pini-thunbergii CBS 473.91
Botryosphaeria qingyuanensis CGMCC 3.18742
Aplosporella prunicola CBS 121167
Botryosphaeria pseudoramosa CGMCC 3.18739
Glyphium elatum EB 0342
Aplosporella papillata CBS 121780
Kellermania micranthae CBS 131724
Trichodelitschia bisporula CBS 262.69
Kirschsteiniothelia phoenicis MFLUCC 18-0216
Zwackhiomyces coepulonus RO31
Lembosina aulographoides CPC 33049
Patellaria atrata CBS 958.97
Bezerromyces brasiliensis CBS 141545
Botryosphaeria wangensis CGMCC 3.18744
Venturia catenospora CBS 447.91
Patellaria atrata SQUCC 15290
Dissoconium aciculare CBS 204.89
Pseudofusicoccum stromaticum CBS 117449
Dothiora cannabinae CBS 737.71
Kellermania confusa CBS 131723
Collemopsidium cf. halodytes RO26
Murramarangomyces corymbiae CPC 33000
Zeloasperisporium cliviae CBS 139915
Lichenothelia calcarea L1842
Diatrype disciformis AFTOL-ID 927
Phyllosticta ericarum CBS 132534
Phyllosticta maculata CBS 132581
Homortomyces tamaricis MFLUCC 13-0441
Saccharata intermedia CBS 125546
Sympoventuria capensis CBS 120136
Patellaria quercus CPC 27232
Zeloasperisporium siamense IFRDCC 2194
Cercospora beticola CBS 116456
Catinella olivacea UAMH 10679
Muripulchra aquatica KUMCC 15-0276
Dothidea insculpta CBS 189.58
Lichenothelia convexa LMCC0484
Holmiella junipericola MFLUCC 18-0503
Tubeufia guangxiensis MFLUCC 17-0045
Myriangium hispanicum CBS 247.33
Holmiella junipericola SQUCC 15186
Hysteropatella elliptica CBS 935.97
Zeloasperisporium pterocarpi MFLUCC 17-0910
Tubeufia paludosa CBS 120503
Zalaria obscura DAOM 250849
Neodactylaria simaoensis YMF 1.3984
Holmiella sabina G.M. 2015-04-29.3
Collemopsidium angermannicum s1473
Holmiella sabina G.M. 2015-04-29.2
Superstratomyces atroviridis CBS 140272
Capnodium salicinum CBS 131.34
Kirschsteiniothelia tectonae MFLUCC 12-0050
Zeloasperisporium eucalyptorum CBS 124809
Trichodelitschia munkii Kruys 201
Botryosphaeria dothidea CBS 115476
Endomelanconiopsis endophytica CBS 120397
Phaeotrichum benjaminii CBS 541.72
Capnodium coffeae CBS 147.52
Hysteropatella clavispora CBS 247.34
Holmiella juniperi-semiglobosae MFLUCC 17-1955
Superstratomyces flavomucosus CBS 353.84
Graphostroma platystoma CBS 270.87
Tumidispora shoreae MFLUCC 14-0574
Elsinoe centrolobii CBS 222.50
Homortomyces combreti CPC 19808
Melanops tulasnei CBS 116805
Elsinoe phaseoli CBS 165.31
Aplosporella africana CBS 121777
Kellermania dasylirionicola CBS 131720
*
*
*
*
**
*
*
*
83/-
69/-
97/1.0
*
*
*
*
*
94/1.0
*
79/1.0
*
60/-
*
*
*
*
*
88/-
62/0.98
86/-
*
94/1.0
*
83/-
*
*
*
98/-
*
*
89/-
88/1.0
73/-
*
98/1.0
95/0.96
69/1.0
84/-
84/-
99/1.0
*
90/1.0
*
71/1.0
Natipusilla decorospora ILL S AF236
Natipusilla limonensis ILL S AF286
Natipusilla naponensis ILL S AF217
*
*
Microthyrium buxicola MFLUCC 15-0213
Microthyrium propagulensis CBS 115976
*
Botryosphaeriales
Holmiellales ordo novus
Homortomycetales ordo novus
Catinellales
Patellariales
Neodactylariales
Tubeufiales
Kirschsteiniotheliales
Zeloasperisporiales
Natipusillales
Phaeotrichales
Microthyriales
Murramarangomycetales
Venturiales
Collemopsidiales
Lembosinales
Lichenoconiales
Capnodiales
Myriangiales
Dothideales
Superstratomycetales
Clade 3
Clade 4
-/1.0
-/0.99
*
-/0.95
2x
3x
2x
2x
2x
2x
Fig. 2 (continued)

Fungal Diversity
1 3
Patellariaceae typified with Patellaria Fr. and two other
genera – Hysteropatella Rehm and Glyphium Nitschke ex
F. Lehm. – form a clade sister to the order Neodactylari-
ales (Clade 4, Fig.2) in Dothideomycetes and recognized
as Patellariales sensu stricto. Two other genera previously
placed in Patellariaceae viz. Banhegyia L. Zeller & Tóth
(Clade 1, Fig.2) and Rhizodiscina Hafellner (Clade 2,
Fig.2) form a sister clade neighbouring the order Asteri-
nales. However, the sequence data of these Banhegyia cf.
setispora and Rhizodiscina lignyota isolates are from an
unpublished study on DNA barcoding of ascomycetes from
the LUX herbarium by Hermant & Marson in 2017. The
species of the genus Holmiella Petrini, Samuels & E. Müll.
form a separate clade (Clade 3, Fig.2) sister to Catinellales,
Homortomycetaceae and Botryosphaeriales and are recog-
nized as a new family and a new order and described herein.
The orders arrangement in the present study is mostly in
agreement with the Hongsanan etal. (2020a).
Patellariales sensu stricto LSU, ITS andTEF phylogeny
The combined LSU, ITS and TEF alignment was used to
understand the species relationship in sequenced genera
in Patellariales sensu stricto. The alignment comprised
28 strains (including the outgroup taxa Neohelicosporium
acrogenisporum (MFLUCC 17-2019) and N. thailandicum
(MFLUCC 16-0221)) and the manually adjusted dataset
comprised 2366 characters including gaps. The RAxML
analysis of the combined dataset yielded the best scor-
ing tree with a final ML optimization likelihood value
of − 8,017.48094. Based on the results of MrModelTest,
dirichlet base frequencies and the GTR+I+G model were
used for the Bayesian analysis. The Bayesian analyses gen-
erated 1951 trees (saved every 100 generations) from which
1464 were sampled after 25% of the trees were discarded
as burn-in.
Notes—The ML and BI analyses showed similar tree
topologies and were congruent; the ML tree is shown in
Fig.3. Phylogenetic results showed that the genera Glyph-
ium, Hysteropatella and Patellaria nested in Patellariales
sensu stricto. Several Patellaria atrata sequences are avail-
able in GenBank, however, to our knowledge, none of these
are linked to a morphological description. The morpho-
logical characters examined in our isolate (SQUCC 15117)
largely overlap with those of the type and those illustrated
in Yacharoen etal. (2015). Therefore, we identified our col-
lection as P. atrata.
Didymellaceae LSU, ITS, TUB andRPB2 phylogeny
The alignment contained 52 isolates representing
Ascochyta Lib.; Briansuttonomyces Crous; Calophoma
Q. Chen & L. Cai; Neodidymelliopsis Q. Chen & L. Cai;
Neomicrosphaeropsis Thambug. etal.; Phoma Sacc.; Pho-
matodes Q. Chen & L. Cai; and Pseudoascochyta Valenz.-
Lopez etal. Leptosphaeria conoidea (De Not.) Sacc. (CBS
616.75) and L. doliolum (Pers.) Ces. & De Not. (CBS
505.75) were used as outgroups to root the tree. The final
alignment contained a total of 2,717 characters used for
the phylogenetic analyses, including alignment gaps. The
RAxML analysis of the combined dataset yielded a best
scoring tree with a final ML optimization likelihood value
of − 12,669.918089. Based on the results of MrModelTest,
dirichlet base frequencies and the GTR+I+G model were
used for the Bayesian analysis. The Bayesian analyses gen-
erated 2,751 trees (saved every 100 generations) of which
2,064 were sampled after 10% of the trees were discarded
as burn-in.
Notes—There were no topological conflicts between
trees generated from ML and BI analyses; the ML tree is
shown in Fig.4. The combination of these four nuclear
loci provided robust phylogenetic support for all genera as
monophyletic lineages. In individual gene analyses, RPB2
was more stable compared to other regions. Newly gener-
ated strain SQUCC 13750 clustered with the remaining
species in Calophoma with strong support.
Didymosphaeriaceae SSU, LSU, TEF andITS phylogeny
The alignment contained 74 isolates, and the tree was
rooted to Bambusistroma didymosporum D.Q. Dai & K.D.
Hyde (MFLU 15-0057 and MFLU 15-0058). The final
alignment contained a total of 3,427 characters used for
the phylogenetic analyses, including alignment gaps. The
RAxML analysis of the combined dataset yielded a best
scoring tree with a final ML optimization likelihood value
of − 21,337.099859. Based on the results of MrModelTest,
dirichlet base frequencies and the GTR+I+G model were
used for the Bayesian analysis. The Bayesian analyses
generated 8751 trees (saved every 100 generations) from
which 7876 were sampled after 10% of the trees were dis-
carded as burn-in.
Notes—Representative strains of all the available gen-
era in Didymosphaeriaceae were included to examine the
phylogenetic placement of SQUCC 13296. The combi-
nation of SSU, LSU, TEF and ITS nuclear loci provided
strong phylogenetic support for the majority as mono-
phyletic lineages except for the monotypic taxa (Fig.5).
SQUCC 13,296 clustered with species of Dictyoarthrin-
ium S. Hughes, had a sister relationship with D. sacchari

Fungal Diversity
1 3
(J.A. Stev.) Damon (MFLUCC 20-0005, CBS 529.73) and
was recognised as a new species.
Phaeosphaeriaceae SSU, LSU, ITS andTEF phylogeny
The alignment contained 24 isolates, and the tree was rooted
to Staurosphaeria rhamnicola Wanas., Gafforov & K.D.
Hyde (MFLUCC 17-0813 and MFLUCC 17-0814). The
final alignment contained a total of 3387 characters used
for the phylogenetic analyses, including alignment gaps.
The RAxML analysis of the combined dataset yielded a best
scoring tree with a final ML optimization likelihood value of
-10,677.693091. Based on the results of MrModelTest, dir-
ichlet base frequencies and the GTR+I+G model were used
for the Bayesian analysis. The Bayesian analyses generated
401 trees (saved every 100 generations) of which 301 were
sampled after 25% of the trees were discarded as burn-in.
Notes —The ML and BI trees generated based on
sequence analysis of the combined dataset indicate that
our new isolate, SQUCC 15290 belongs in Phaeospha-
eriaceae and clusters with Hydeomyces desertipleosporoides
(SQUCC 15259 and SQUCC 15260) and H. pinicola (GZ-
06) with 100% ML and 1.00 PP statistical support (Fig.6).
Pleosporineae SSU, LSU, RPB2 andTEF phylogeny
The alignment contained 78 isolates, and the tree was rooted
to Massarina cisti S.K. Bose (CBS 266.62) and M. eburnea
(Tul. & C. Tul.) Sacc. (CBS 473.64). The final alignment
contained a total of 3,662 characters used for the phylo-
genetic analyses, including alignment gaps. The RAxML
analysis of the combined dataset yielded a best scor-
ing tree with a final ML optimization likelihood value of
− 27,502.223035. The Bayesian analyses generated 49,701
0.04
Patellaria andina G.M. 2003-05-28.712-5
Glyphium elatum EB 0329
Patellaria microspora MFLU 16-0567
Hysteropatella prostii G.M. 2016-02-20.2
Neohelicosporium thailandicum MFLUCC 16-0221
Patellaria atrata SQUCC 15117
Patellaria chromolaenae MFLUCC 17-1479
Patellaria andina G.M. 2003-05-30.716-5
Neohelicosporium acrogenisporum MFLUCC 17-2019
Patellaria cf. atrata G.M. 2001-11-26
Patellaria cf. atrata BCC 28877
Patellaria apiculatae MFLU 19-1236
Hysteropatella prostii G.M. 2014-05-20
Patellaria cf. atrata BCC 28876
Glyphium elatum EB 0342
Patellaria sp. G.M. 2003-06-10.741
Patellaria andina G.M. 2003-05-30.715.195-2
Patellaria atrata G.M. 2013-03-19.1
Hysteropatella clavispora CBS 247.34
Patellaria andina G.M. 2003-05-28.710-5
Patellaria atrata CBS 958.97
Glyphium elatum EB 0388
Hysteropatella elliptica G.M. 2013-05-06
Patellaria chromolaenae MFLU 16-0578
Patellaria atrata JAC11495
Patellaria chromolaenae MFLUCC 17-1482
Patellaria atrata (=P. quercus) CPC 27232
Hysteropatella prostii H.B. 9934b
*
96/1.0
85/0.95
*
*
83/-
*
*
*
93/0.99
*
98/1.0
84/-
99/1.0
97/1.0
*
82/-
*
Patellaria
Fig. 3 Phylogenetic tree inferred using the combined sequences of
LSU, ITS and TEF of the analysed Patellariales sensu stricto. The
tree backbone was constructed using maximum likelihood (ML)
analysis. Species are indicated in coloured blocks. The MLB and
PP above 60% and 0.95, respectively, are given at the nodes (MLB/
PP). An asterisk (*) indicates branches with MLB = 100% and PP
values = 1.0. Species are followed by the strain accession numbers.
The new strain is indicated in blue bold. The scale bar represents the
expected number of changes per site for ML

Fungal Diversity
1 3
trees (saved every 1000 generations) of which 44,731 were
sampled after 10% of the trees were discarded as burn-in.
Notes—The single-locus datasets were examined for
topological incongruence among members of the Ple-
osporineae. The conflict-free alignments of each gene region
were concatenated into a multi-locus alignment that was
subsequently subjected to phylogenetic analyses of the sub
order. Based on results of the four-gene phylogeny, SQUCC
3026 was part of a monophyletic Halojulellaceae with abso-
lute support. There were no topological conflicts between
trees generated from ML and BI analyses and the ML tree
is shown in Fig.7.
0.07
Calophoma vincetoxici CBS 185.55
Calophoma complanata CBS100311
Calophoma petasitis MFLUCC15-0076
Ascochyta rabiei CBS206.30
Phoma herbarum CBS 502.91
Phomatodes nebulosa CBS 100191
Phoma herbarum CBS 274.37
Calophoma aquilegiicola CBS 107.31
Neomicrosphaeropsis elaeagni MFLUCC17-0740
Phoma herbarum CBS 615.75
Neomicrosphaeropsis cytisi MFLUCC130396
Neomicrosphaeropsis italica MFLUCC15-0485
Calophoma glaucii CBS114.96
Ascochyta rabiei CBS237.37
Pseudoascochyta novae-zelandiae CBS141689
Leptosphaeria doliolum CBS505.75
Neomicrosphaeropsis tamaricicola MFLUCC14-0602
Neodidymelliopsis xanthina CBS383.68
Calophoma clematidis-rectae CBS507.63
Calophoma complanata CBS268.92
Calophoma aquilegiicola CBS 107.96
Ascochyta coronillae-emeri MFLUCC13-0820
Ascochyta phacae CBS184.55
Briansuttonomyces eucalypti CBS114879
Calophoma glaucii CBS112.96
Calophoma rosae LC 8119
Phoma herbarum CBS 134.96
Calophoma humuli MFLUCC18-0101
Phomatodes aubrietiae CBS627.97
Neomicrosphaeropsis alhagi-pseudalhagi MFLUCC17-0825
Calophoma clematidina CBS 108.79
Calophoma hydei SQUCC 13750
Calophoma rosae CGMCC3.18347
Briansuttonomyces eucalypti CBS114887
Ascochyta herbicola CBS 629.97
Leptosphaeria conoidea CBS616.75
Calophoma vodakii CBS173.53
Neodidymelliopsis achlydis CBS 256.77
Ascochyta pisi CBS122751
Calophoma parvula CBS 620.68
Calophoma aquilegiicola CBS108.96
Calophoma clematidina CBS 102.66
Phomatodes nebulosa CBS 117.93
Neodidymelliopsis cannabis CBS234.37
Calophoma aquilegiicola CBS 116402
Calophoma aquilegiicola CBS109.96
Calophoma sandfjordenica CBS145571
Phomatodes nebulosa CBS 740.96
Neomicrosphaeropsis cytisicola MFLU 16-0114
Neodidymelliopsis polemonii CBS109181
Phomatodes aubrietiae CBS383.67
Phoma herbarum CBS 377.92
97/-
*
60/0.95
*
*
89/-
*
*
88/1.0
63/-
*
82/-
*
99/1.0
99/1.0
*
97/1.0
*
*
*
67/0.96
76/0.96
87/0.98
69/0.98
*
*
74/0.99
*
*
62/-
93/1.0
96/1.0
*
99/1.0
97/1.0
68/1
94/0.97
*
95/0.99
89/0.97
79/-
64/-
*
Calophoma
Fig. 4 Phylogenetic tree inferred using the combined sequences of
LSU, ITS, TUB and RPB2 of the analysed Didymellaceae genera.
The tree backbone was constructed using maximum likelihood (ML)
analysis. The MLB and PP above 60% and 0.95, respectively, are
given at the nodes (MLB/PP). An asterisk (*) indicates branches with
MLB = 100% and PP values = 1.0. Species are followed by the strain
accession numbers. The new strain is indicated in blue bold. The
scale bar represents the expected number of changes per site for ML

Fungal Diversity
1 3
0.03
Alloconiothyrium aptrootiiCBS 981.95
Tremateiaarundicola MFLU 16-1275
Letendraeacordylinicola MFLUCC 11-0148
Spegazziniaradermacherae MFLUCC 17-2285
Letendraea padouk CBS 485.70
MontagnulakrabiensisMF LUCC 16-0250
Deniquelatabarringtoniae MF LUCC 11-0422
Kalmusia ebuli CBS 123120
Paramassariosphaeria anthostomoides CBS 615.86
LaburnicolahawksworthiiMFLUCC 13-0602
MontagnulacirsiiMF LUCC 13 0680
Alloconiothyrium aptrootiiCBS 980.95
Paracamarosporium fagi CPC 24890
Kalmusibambusa triseptata MFLUCC 13-0232
Karstenula rhodostoma CBS 691.94
Kalmusia variisporumCBS 121517
Paracamarosporium fagi CPC 24892
Paraphaeosphaeria rosae MFLUCC 17-2547
Spegazzinia tessarthra SH 287
NeokalmusiaarundinisMFLUCC 16-0405
NeokalmusiabrevisporaKT 1466
Dictyoarthrinium sacchari MFLUCC 20-0005
Chromolaenicolalampangensis MFLUCC 17-1462
Phaeodothiswinteri CBS 182.58
Xenocamarosporium acaciae MFLUCC 17-2432
Didymosphaeria rubi -ulmifolii MFLUCC 14-0023
ChromolaenicolathailandensisMFLUCC 17-1475
Dictyoarthrinium hydei SQUCC 13296
Neokalmusiascabrispora KT 1023
Xenocamarosporium acaciae CPC 24755
Letendraeacordylinicola MFLUCC 11-0150
Austropleosporaarchidendri CBS 168.77
Montagnulabellevaliae MF LUCC 14-0924
Montagnulasaik huensis MFLUCC 16-0315
Karstenula rhodostoma CBS 690.94
Didymosphaeria rubi -ulmifolii MFLUCC 14-0024
Didymocrea sadasivanii CBS 438.65
Bimuria novae -zelandiae CBS 107.79
Didymosphaeria rubi -ulmifolii CBS 100299
Bimuria omanensis SQUCC 15280
Dictyoarthrinium musaeMFLUCC 20-0004
Laburnicola muriformis MFLUCC 14-0921
Pseudocamarosporium pteleaeMFLUCC 17-0724
Spegazzinia deightonii yone 212
Letendraea helminthicola CBS 884.85
Kalmusia italicaMFLUCC 14-0560
Paramassariosphaeria clematidicola MFLU 16-0172
Paraconiothyrium cyclothyrioidesCBS 972.95
Montagnulascabiosae MF LUCC 14-0954
Paraconiothyrium estuarinum CBS 109850
Pseudopithomycesentadae MFLUCC 17-0917
ChromolaenicolathailandensisMFLUCC 17-1510
Verrucoconiothyrium nitidaeCBS 119209
PseudopithomyceskunmingensisMFLUCC 17-0314
Tremateiaguiyangensis GZAAS01
Paraphaeosphaeria rosaeMFLUCC 17-2549
Paracamarosporium hawaiiense CBS 120025
Laburnicola muriformis MFLUCC 16-0290
Austropleosporaosteospermi MFLUCC 17-2429
Deniquelataquercina ABRIICC 10068
Paraphaeosphaeria rosicola MFLUCC 15-0042
Deniquelatabarringtoniae MFLUCC 16-0271
Dictyoarthrinium sacchari CBS 529.73
Neptunomyces aureus CMG13
Cylindroaseptospora leucaenae MFLUCC 17-2424
Dictyoarthrinium musaeMFLUCC 20-0003
BambusistromadidymosporumMFLU 15-0057
BambusistromadidymosporumMFLU 15-0058
Neptunomyces aureus CMG12
Pseudocamarosporium ulmi-minoris MFLUCC 17-0671
NeokalmusiabrevisporaKT 2313
Pseudocamarosporium propinquum MFLUCC 13-0544
Pseudopithomycesrosae MF LUCC 15-0035
Neptunomyces aureus CMG14
*
*
*
*
*
*
*
100/0.95
99/1.0
*
*
78/1.0
*
*
*
*
75/1.0
77/1.0
75/-
*
*
65/-
68/-
76/-
*
98/0.96
93/1.0
*
99/1.0
71/-
80/1.0
*
*
*
72/-
99/1.0
*
74/0.99
*
89/1.0
99/1.00
*
98/-
90/1.0
98/0.95
*
74/-
62/-
*
*
100
78
75/1.0
100
90
69
-/0.99
*
*
*

Fungal Diversity
1 3
Lophiostomataceae LSU, SSU, ITS, RPB2 andTEF phylogeny
The alignment contained 56 isolates and the tree was rooted
to Teichospora rubriostiolata Jaklitsch & Voglmayr (TR 7)
and T. trabicola Fuckel (C 134). The final alignment con-
tained a total of 4419 characters used for the phylogenetic
analyses, including alignment gaps. The RAxML analysis
of the combined dataset yielded a best scoring tree with a
final ML optimization likelihood value of − 25,819.148291.
Based on the results of MrModelTest, dirichlet base frequen-
cies and the GTR+I+G model were used for the Bayesian
analysis. The Bayesian analyses generated 506 trees (saved
every 100 generations) from which 456 were sampled after
10% of the trees were discarded as burn-in.
Notes—We included representative taxa from the gen-
era that have sequence data for Lophiostomataceae in Gen-
Bank (Hongsanan etal. 2020b). We did not resolve strongly
supported monophyletic lineages for these members in the
single gene phylogenetic analyses. The concatenated data
analysis provided a robust and similar topology to recently
published data (Wanasinghe etal. 2016; Hashimoto etal.
2018; Bao etal. 2019). In the ML and Bayesian analyses,
our isolate (SQUCC 15092) grouped with Dimorphiopsis
brachystegiae Crous (CPC 22679) and Pseudopaucispora
brunneospora A. Hashim. etal. (KH 227) at the basal posi-
tion in Lophiostomataceae.
Testudinaceae SSU, LSU, 5.8S andTEF phylogeny
The alignment contained 56 isolates and the tree was rooted
to Lophiostoma arundinis (Pers.) Ces. & De Not. (CBS
621.86) and L. crenatum (Pers.) Fuckel (CBS 629.86). The
final alignment contained a total of 3,494 characters used for
the phylogenetic analyses, including alignment gaps. The
RAxML analysis of the combined dataset yielded a best
scoring tree with a final ML optimization likelihood value
of − 16,931.116907. Based on the results of MrModelTest,
dirichlet base frequencies and the GTR+I+G model were
used for the Bayesian analysis. The Bayesian analyses gen-
erated 18,201 trees (saved every 100 generations) of which
13,651 were sampled after 25% of the trees were discarded
as burn-in.
Notes—The genera of Testudinaceae (Verruculina
Kohlm. & Volkm.-Kohlm., Ulospora D. Hawksw.,
Muritestudina Wanas. etal., Neotestudina Segretain &
Destombes, Lepidosphaeria Parg.-Leduc, Xenolophium
Syd., Halotestudina Dayar. & K.D. Hyde, Trematosphaeria
Fuckel and Angustospora Abdel-Aziz) and related families
were included in the present analysis (Fig.8). The overall
topology of the phylogenetic tree was in accordance with
Wanasinghe etal. (2017). Overall the terminal clades are
well-supported in the analysis. The isolate from our col-
lection formed a moderately supported clade sister to the
monotypic genus Lepidosphaeria represented by L. nicotiae
Parg.-Leduc (CBS 101341).
Corticiales SSU, LSU andITS phylogeny
The alignment contained 31 isolates and this tree was rooted
to Gloeophyllum sepiarium (Wulfen) P. Karst. (Wilcox
3BB). The final alignment contained a total of 4,419 charac-
ters used for the phylogenetic analyses, including alignment
gaps. The RAxML analysis of the combined dataset yielded
a best scoring tree with a final ML optimization likelihood
value of − 16,539.058910. Based on the results of MrMod-
elTest, dirichlet base frequencies and the GTR+I+G model
were used for the Bayesian analysis.
Notes—Within Corticiales, a total of five main clades
were resolved, which showed a moderately supported back-
bone and was congruent with the results of Li etal. (2016)
(Fig.9). These five clades represent four families viz. Cor-
ticiaceae, Dendrominiaceae, Punctulariaceae, Vuillemi-
niaceae and Leptocorticium tenellum Nakasone placed in
Corticiales genera incertae sedis. The hyphomycetes fungus
from the present study (SQUCC 15289) was part of a clade
that included the hyphomycete Tretopileus sphaerophorus
(Berk. & M.A. Curtis) S. Hughes & Deighton (JCM10092),
coelomycete Giulia tenuis (Sacc.) Tassi ex Sacc. & D.
Sacc. (BCC 13066), and cereal crops and turf grass patho-
gen Waitea circinata Warcup & P.H.B. Talbot (AFTOL-ID
1129) (Pirozynski and Shoemaker 1971; Okada 1998; de la
Cerda etal. 2007).
Taxonomy
Phylum Ascomycota Caval.-Sm., Biological Reviews Cam-
bridge 73: 247 (1998)
Notes—The Ascomycota are commonly known as sac
fungi or ascomycetes and the phylum consists of subphyla
Pezizomycotina, Saccharomycotina and Taphrinomycotina
(Lumbsch and Huhndorf 2010). Nearly 6600 genera have
been listed as ascomycetes and the placement of genera
has been rapidly modified in the recent past with the use of
sequence data. The subphylum Saccharomycotina includes
only a single class while Pezizomycotina and Taphrinomy-
cotina include five and 13 classes, respectively (Wijaya-
wardene etal. 2018, 2020).
Fig. 5 Phylogenetic tree inferred using the combined sequences of
SSU, LSU, TEF and ITS of the analysed Didymosphaeriaceae genera.
The tree backbone was constructed using maximum likelihood (ML)
analysis. The MLB and PP above 60% and 0.95, respectively, are
given at the nodes (MLB/PP). An asterisk (*) indicates branches with
MLB = 100% and PP values = 1.0. Species are followed by the strain
accession numbers. The new strain is indicated in blue bold. The
scale bar represents the expected number of changes per site for ML
◂

Fungal Diversity
1 3
Subphylum Pezizomycotina O.E. Erikss. & Winka,
Myconet 1: 9 (1997)
Notes—The Pezizomycotina is the largest subphylum of
Ascomycota that comprises 13 classes: Arthoniomycetes,
Coniocybomycetes, Dothideomycetes, Eurotiomycetes,
Geoglossomycetes, Laboulbeniomycetes, Lecanoromy-
cetes, Leotiomycetes, Lichinomycetes, Orbiliomycetes,
Pezizomycetes, Sordariomycetes, Xylonomycetes and
Xylobotryomycetes (Wijayawardene etal. 2020). Most of
these classes consist of filamentous, ascoma-producing
species with diverse ecologies (Spatafora etal. 2006).
Class Dothideomycetes O.E. Erikss. & Winka,
Myconet 1: 5 (1997)
Notes—The recent Outline of Fungi and fungus-like
taxa (Wijayawardene etal. 2020) provided the most
updated arrangement of Dothideomycetes, which included
38 orders. In this current study, we introduce two other
orders, Homortomycetales and Holmiellales. Divergence
time estimation for Dothideomycetes fell in the range of
400–492 Mya (crown age) (Liu etal. 2017).
Holmiellales Maharachch. & Wanas., ordo
novus
MycoBank: MB 837548
Saprobic especially on dead wood in terrestrial habitats.
Sexual morph: Ascomata apothecial, solitary or gregari-
ous, immersed to superficial, globose to sub-globose, uni-
to multi-locular, black. Exciple pseudoparenchymatous.
Hamathecium composed of filamentous, anastomosing pseu-
doparaphyses, forming a dark brown epithecium above the
asci. Asci 8-spored, bitunicate, clavate to cylindro-clavate,
short or long-pedicellate, apically rounded with an ocu-
lar chamber. Ascospores 2–3-seriate overlapping, septate,
clavate to ellipsoidal or ovoid, hyaline to brown, with or
without a mucoid sheath. Asexual morph: corniculari-
ella-like. Conidiomata pycnidial, superficial or immersed
in cultures. Conidiophores branched, hyaline, septate.
0.02
Dematiopleospora salsolae MFLUCC 17-0828
Dlhawksworthia clematidicola MFLUCC 14-0910
Ophiobolus ponticus MFLUCC 17-2273
Muriphaeosphaeria galatellae MFLUCC 14-0614
Phaeosphaeriopsis dracaenicola MFLUCC 11-0193
Dematiopleospora mariae MFLU 16-0121
Dlhawksworthia lonicerae MFLUCC 14-0955
Pseudoophiobolus rosae MFLUCC 17-1786
Pseudoophiobolus galii MFLUCC 17-2257
Hydeomyces hydei SQUCC 15290
Dlhawksworthia alliariae MFLUCC 13-0070
Pseudoophiobolus urticicola KUMCC 17-0168
Ophiobolopsis italica MFLUCC 17-1791
Paraophiobolus arundinis MFLUCC 17-1789
Hydeomyces desertipleosporoides SQUCC 15260
Hydeomyces desertipleosporoides SQUCC 15259
Dematiopleospora mariae MFLUCC 15-0612
Nodulosphaeria guttulatum MFLUCC 15-0069
Phaeosphaeriopsis glaucopunctata CBS 653.86
Staurosphaeria rhamnicola MFLUCC 17-0814
Hydeomyces pinicola GZ-06
Nodulosphaeria multiseptata MFLUCC 15-0078
Ophiobolus disseminans MFLUCC 17-1787
Staurosphaeria rhamnicola MFLUCC 17-0813
86/0.96
*
*
72/-
*
97/1.0
*
*
88/1.0
*
*
*
*
66/-
78/-
86/1.0
**
*
Fig. 6 Phylogenetic tree inferred using the combined sequences of
LSU, SSU, ITS and TEF of the analysed Hydeomyces and allied gen-
era in Phaeosphaeriaceae. The tree backbone was constructed using
maximum likelihood (ML) analysis. Genera are indicated in col-
oured blocks. The MLB and PP above 60% and 0.95, respectively, are
given at the nodes (MLB/PP). An asterisk (*) indicates branches with
MLB = 100% and PP values = 1.0. Species are followed by the strain
accession numbers. The new strain is indicated in blue bold. The
scale bar represents the expected number of changes per site for ML

Fungal Diversity
1 3
Conidiogenous cells integrated, hyaline. Conidia straight to
curved, hyaline. Chlamydospores may present.
Type family: Holmiellaceae Maharachch. & Wanas.
Holmiellaceae Maharachch. & Wanas., fam.
nov.
MycoBank: MB 837549
Saprobic on dead wood in terrestrial habitats. Sexual
morph: Ascomata apothecial, solitary, superficial, expos-
ing a velvet hymenium when mature, globose, black. Exci-
ple pseudoparenchymatous. Hamathecium composed of
filamentous, septate, branched, hyaline, anastomosing
pseudoparaphyses, forming a dark brown epithecium above
the asci. Asci 8-spored, bitunicate, with an ocular chamber.
Ascospores 2–3-seriate overlapping, clavate to ellipsoidal,
1-septate, brown to dark brown. Asexual morph: cornicu-
lariella-like. Conidiomata pycnidial, globose, black, super-
ficial or immersed in cultures. Conidiophores branched,
hyaline, septate. Conidiogenous cells integrated, subulate,
hyaline. Conidia straight to curved, hyaline. Chlamydo-
spores may present.
Type genus: Holmiella Petrini, Samuels & E. Müll.
Notes—This new order Holmiellales and new family Hol-
miellaceae are introduced for a lineage of saprobic fungi that
were previously placed in the monotypic order Patellariales.
The family Patellariaceae was introduced by Corda (1839) to
accommodate the genera Cryptodiscus Corda and Mellitio-
sporium Corda (Kutorga and Hawksworth 1997; Yacharoen
etal. 2015). Hongsanan etal. (2020b) reported that Patellari-
ales was around evolved between 31 Mya (crown age) and
276 Mya (stem age). Patellariaceae is heterogenous family
characterised by apothecial fruiting bodies, and in the lit-
erature, taxa belonging to the group have been described in
various families and orders (Nannfeldt 1932; Luttrell 1951;
Reid and Pirozynski 1966; Barr 1979; Samuels and Müller
1979; Haffellner 1979; Petrini etal. 1979; Eriksson 1981;
Eriksson and Hawksworth 1993; Boehm etal. 2009; Schoch
etal. 2009; Jones and Pang 2012). The arrangement based
on Yacharoen etal. (2015) recognised 14 genera, while a
recent outline by Wijayawardene etal. (2020) accepted 21
genera within the Patellariaceae. However, most of these
genera lack sequence data, and therefore, natural classifica-
tion of Patellariaceae within the Dothideomycetes is still
incomplete. The phylogenetic analysis of combined SSU,
LSU, 5.8S, RPB2 and TEF sequences for species of Hol-
miella, revealed that the genus is closely related to Botry-
osphaeriales, Catinellales and Homortomycetaceae, refut-
ing earlier suggestions of its placement in Patellariaceae
(Yacharoen etal. 2015; Pem etal. 2018). Holmiella was
erected by Petrini etal. (1979) to accommodate Triblidium
sabinum De Not. (= Holmiella macrospora) designated by
De Notaris in 1867. The type species Holmiella sabina (De
Not.) Petrini, Samuels & E. Müll. inhabits as an endophyte
or saprobe within needles and wood of Juniperus species
(Holm and Holm 1977; Kutorga and Hawksworth, 1997;
Eriksson 2014). The new order Holmiellales has a stem age
at 97 million of years and has the same common ancestor
with new orders Homortomycetales at 120 million of years
(Supplementary Fig.1).
In the past Holmiella sabina has been assigned to diverse
genera (Kutorga and Hawksworth 1997). Later the second
species, H. macrospora (Bonar & E.K. Cash) Kutorga & D.
Hawksw., was added to the genus by Kutorga & Hawksworth
(1997); this species inhabits bark of Calocedrus decurrens
(Torr.) Florin in the USA that was introduced as Tryblidiella
macrospora Bonar & E.K. Cash by Cash (1945). Pem etal.
(2018) introduced Holmiella junipericola Pem etal. and H.
juniperi-semiglobosae Pem etal., thus the genus presently
comprises four species. Based on evidence from phylog-
eny and morphology in our assessments, we suggest that
the species of Holmiella do not belong to Patellariales, and
therefore the new order Holmiellales and the new family
Holmiellaceae are introduced herein. Species of Holmiella
have apothecial ascomata which are initially enclosed in host
parenchyma, and subsequently crack open to become finally
exposed through a wide, irregular hymenium. The apothecial
ascomata in Holmiellales, unlike most other Dothideomy-
cetes, are clearly distinct from Botryosphaeriales and the
other newly introduced order, Homortomycetales. Apart
from Holmiella, the corniculariella-like asexual morphs are
only known from the Leotiomycetes family Dermateaceae.
Pem etal. (2018) also suggested that H. juniperi-semiglo-
bosae sequences are related to unidentified endophytic iso-
lates in GenBank, which are designated as Botryosphaeriales
sp. Because morphological details cannot be compared, they
did not further explain the relationship but emphasized that
this relationship is interesting. Further, they did not go into
details as their phylogeny lacked other taxa, apart from the
genera placed in Patellariaceae.
Homortomycetales Maharachch. & Wanas.,
ordo novus
MycoBank: MB 837547
Foliicolous, associated with leaf spots or saprobic on dead
twigs or branches in terrestrial habitats. Sexual morph:
Ascomata scattered, immersed to erumpent, ostiolate.
Ostiole central, apapillate to papillate, filled with hya-
line to brown cells. Peridium comprising angularis cells.

Fungal Diversity
1 3
0.04
Pyrenophora phaeocomes DAOM 222769
Parapyrenochaeta protearumCBS 131315
Coniothyrium telephii CBS188.71
Neocamarosporium lamiacearum MFLUCC 17-0560
Neophaeosphaeriafilamentosa CBS102202
Massarinacisti CBS 266.62
Pyrenochaetopsis americanaUTHSCDI16225
Camarosporiumquaternatum CPC31081
Libertasomyces quercus CBS134.97
Thyrostromatiliae MFLUCC 16-1178
Acrocalymmaaquaticum MFLUCC 11-0208
Neocamarosporium korfii MFLUCC 17-0745
Pseudopyrenochaetaterrestris CBS282.72
Plenodomusartem isiae KUMCC18-0151
Acrocalymmafici CBS317.76
Camarosporiumquaternatum CPC23216
Coniothyrium dolichiCBS124140
Halojulella avicenniae JK 5326A
Tzeanania taiwanensis NTUCC17-005
Acrocalymmawalkeri UTHSCDI16-195
Pyrenochaetopsis leptospora CBS101635
Halojulella avicenniae GR 00584
Ascocylindrica marina MF416
NeocucurbitariaribicolaCBS142394
Shiraiabambusicola NBRC 30771
Phaeosphaeria thysanolaenicola MFLUCC 10-0563
Neocamarosporium salsolae MFLUCC 17-0826
Cucurbitaria berberidis CBS142401
Leptosphaeria doliolum CBS505.75
Phaeosphaeria musaeMFLUCC 11-0133
Alternaria eureka CBS193.86
Ascochytacoronillae-emeri MFLUCC 13-0820
Bipolaris maydis CBS 134.39
Neophaeosphaeria agaves CPC 21264
Boeremia exigua CBS431.74
Neopyrenochaetatelephoni CBS139022
Alternariaster helianthiCBS327.69
Libertasomyces myopori CPC27354
Stemphyliumvesicarium CBS191.86
Shiraiabambusicola JCM1879
Massarinaeburnea CBS473.64
Coniothyrium telephii CBS856.97
Libertasomyces platani CPC29609
Cucurbitaria oromediterraneaCBS142399
Parapyrenochaeta protearumCBS137997
Dothidotthia negundinicolaMFLUCC 16-1157
CamarosporidiellaaborescentisMFLUCC 17-0660
Pseudopyrenochaeta lycopersiciCBS 306.65
Pyrenochaetopsis botulispora CBS142458
Didymellaglomerata CBS528.66
Halojulella avicenniae BCC20173
Tzeanania taiwanensis NTUCC17-006
Microsphaeropsis spartii-juncei MFLU 16-0100
Halojulella avicenniae BCC28357
Shiraiabambusicola NBRC 30754
NeocucurbitariarhamniCBS142391
Coniothyrium glycines CBS124455
Thyrostromalycii MFLUCC 16-1170
AscocylindricamarinaMD6011
Alternaria alternataCBS916.96
Shiraiabambusicola NBRC 30772
Phaeosphaeria chiangraina MFLUCC 13-0231
Neopyrenochaeta acicolaCBS812.95
Omania hydei SQUCC 3026
Plenodomus sinensis MFLUCC 17-0767
Phaeosphaeriopsisdracaenicol aMFLUCC 11-0157
Neopyrenochaetafragariae CBS101634
Neopyrenochaeta inflorescentiae CBS119222
Parapyrenochaeta acaciae CBS141291
Xenopyrenochaetopsis pratorum CBS445.81
Dothidotthia robiniae MFLUCC 16-1175
Acrocalymmapterocarpi MFLUCC 17-0926
Camarosporiumquaternatum CPC31518
Camarosporidiellaitalica MFLUCC 13-0547
Neophaeosphaeriafilamentosa CBS102203
Halojulella avicenniae BCC18422
Neocamarosporium salsolae MFLUCC 17-0827
caraganicolaMFLUCC 14-0887
98/0.98
62/-
87/1.0
*
*
98/1.0
*
84/0.95
*
*
99/0.99
73/1.0
*
60/-
63/-
85/1.0
*
*
99/1.0
82/-
82/0.91
*
*
84/0.95
*
97/1.0
*
70/0.95
60/-
*
94/0.98
*
65/1.0
70/1.0
*
*
*
99/1.0
*
*
99/1.0
97/0.99
*
97/1.0
99/0.98
*
*
61
*
63/-
81/1.0
*
100
Pleosporaceae
Libertasomycetaceae
Neocamarosporiaceae
Camarosporiaceae
Parapyrenochaetaceae
Camarosporidiellaceae
Leptosphaeriaceae
Coniothyriaceae
Neopyrenochaetaceae
Pyrenochaetopsidaceae
Cucurbitariaceae
Pseudopyrenochaetaceae
Shiraiaceae
Tzeananiaceae
Phaeosphaeriaceae
Neophaeosphaeriaceae
Dothidotthiaceae
Didymellaceae
Halojulellaceae
Acrocalymmaceae
Ascocylindricaceae
*

Fungal Diversity
1 3
Hamathecium with septate, cellular pseudoparaphyses. Asci
2–8-spored, bitunicate, fissitunicate, clavate to cylindrical
clavate. Ascospores uni- to bi-seriate, fusiform, septate.
Asexual morph: Coelomycetous. Conidiomata pycnidial,
immersed or erumpent, uni- to multi-loculate, ostiolate.
Conidiomatal wall comprising cells of textura angularis.
Paraphyses hyaline, flexuous, sparingly septate. Conidio-
phores reduced to conidiogenous cells or with one support-
ing cell. Conidiogenous cell with a supporting cell, hyaline,
percurrent, proliferate. Conidia ellipsoid to subcylindrical,
3(− 4)-euseptate.
Type family: Homortomycetaceae Thambugala, A.J.L.
Phillips & K.D. Hyde
Type species: Homortomyces combreti Crous & M.J.
Wingf. IMA Fungus 3(2): 113 (2012)
Notes—The family Homortomycetaceae was introduced
by Thambugala etal. (2017) as a monotypic family and cur-
rently includes two species, the generic type Homortomyces
combreti Crous & M.J. Wingf. and H. tamaricis Wijayaw.
etal. The LSU sequence analysis of Thambugala etal.
(2017) showed that the closest relative to Homortomyceta-
ceae is Botryosphaeriales. Although Homortomycetaceae
shares feature such as stromatic conidiomata, septate, pig-
mented, thick-walled and diplodia-like conidia, and asci with
a thick endotunicate and a well-developed apical chamber
with Botryosphaeriales, it is clearly distinct from the latter
in having uniloculate, thin-walled ascomata and 3-septate,
pigmented ascospores and distoseptate conidia (Crous etal.
2012; Phillips etal. 2013; Thambugala etal. 2017). Based
on morphology and phylogenetic placement, Thambugala
etal. (2017) tentatively referred to Homortomycetaceae as
an incertae sedis family in Dothideomycetes. Phylogenetic
analyses of combined genes in the present study indicate
that Holmiellales is closely related to the Homortomyceta-
ceae and Catinellales (see notes under Holmiellales). The
order Catinellales was erected by Hongsanan etal. (2020b)
for the genus Catinella Boud. which was previously placed
within Leotiomycetes. The order Catinellales is character-
ised by apothecial, discoid ascomata, excipulum composed
of angularis to globose cells and cylindric, septate, para-
physes clearly distinguish it from Holmiellales and Homor-
tomycetales. Due to phylogenetic distinction and the strong
phenotypic differentiation between Homortomyces Crous &
M.J. Wingf., Catinella, Holmiella and Botryosphaeriales,
the separation into four orders seems straightforward. Future
studies including more taxon sampling will shed additional
light on the classification of Botryosphaeriales, Catinellales,
Homortomycetales and Holmiellales. The order Homorto-
mycetales has a stem age at 97 million of years (Supple-
mentary Fig.1).
Holmiella junipericola Pem, Gafforov, Jeewon & K.D.
Hyde, Cryptog. Mycol. 39(2): 200 (2018) amend.
MycoBank: MB 825002; Fig.10
Other specimens examined: Oman.
Saprobic on dead trunks and branches of Juniperus
seravschanica. Sexual morph: Ascomata 0.4–0.8mm diam.,
apothecial, solitary to gregarious in groups of a few, superfi-
cial, sessile, at first pulvinate and covered with a continuous
outer layer, outer layer splitting irregularly into 3–6 lobes
that fold back to expose a black hymenium, sub-gelatinous
when rehydrated, with smooth outer surface of the recep-
tacle, black. Exciple consisting of a dark-brown, leathery
layer, 20–50μm thick, pseudoparenchymatous, two-layered;
outer layer thick, comprised of dark brown to black, slightly
cracked cells of textura angularis; inner layer, thin, com-
posed of light brown cells of textura epidermoidea. Sub-
hymenium inverted conically, of marginally elongated and
more or less horizontally oriented brown cells. Hypothecium
prosenchymatous consisting of textura intricate, light brown,
10–35μm tall below the subhymenium base to 0.5mm diam.
Hamathecium 1–2μm wide, filiform, branched, interwoven
hyphal filaments, forming a dark brown epithecium above
the asci. Asci 100–145 × 35–45μm, 8-spored, clavate, bitu-
nicate, fissitunicate, with an ocular chamber in the apical
dome. Ascospores 26–38 × 10–18μm, irregularly 2–3 seri-
ate, overlapping, ellipsoidal, 1-septate in the middle and
slightly constricted at the septum, slightly granulated, rang-
ing from hyaline when immature to light brown and dark
brown when mature, with a hyaline papilla at the ends, upper
cell slightly wider than lower cell, thick and smooth-walled.
Asexual morph: corniculariella-like. Conidiomata pycnidial,
0.5–1mm diam., globose, black, superficial or immersed in
PDA. Phialides cylindrical to sub-cylindrical, 12–20μm
long, tapering from 1.5–2μm wide basally to 1–1.5μm
wide at the unflared opening, solitary or in pairs, arising from
short conidiophores. Conidiophores branched, hyaline, sep-
tate. Conidiogenous cells integrated. Conidia stylosporous,
14–22 × 0.5–1.5μm, straight to sharply curved, unicellular,
hyaline.
Culture characteristics—Colonies on PDA slow growing
reaching 2cm diam. In 10days, green to olivaceous green,
flat, with moderate aerial mycelium, smooth, crenate margin.
Material examined—OMAN, Al Jabal al-Akhdar (Green
Mountain), from dead branches of Juniperus seravschanica,
07.2016, SSN Maharachchikumbura OM07 (SQU H-116),
living culture = SQUCC 15186.
Fig. 7 Phylogenetic tree inferred using the combined sequences
of SSU, LSU, RPB2 and TEF of the analysed Halojulellaceae and
related families in Pleosporineae. The tree backbone was constructed
using maximum likelihood (ML) analysis. The MLB and PP above
60% and 0.95, respectively, are given at the nodes (MLB/PP). An
asterisk (*) indicates branches with MLB = 100% and PP values = 1.0.
Species are followed by the strain accession numbers. The new strain
is indicated in blue bold. The scale bar represents the expected num-
ber of changes per site for ML
◂

Fungal Diversity
1 3
0.03
Sigarispora ononidis MFLUCC15-2667
Crassiclypeus aquaticusKT 970
Sigarispora arundinis KT 651
Neotrematosphaeria biappendiculata KT 975
Lophiopoacea winteri KT 740
Vaginatispor ascabrispora KT 2443
Pseudolophiostoma cornisporum KH 322
Flabellascoma cycadicola KT 2034
Lophiostoma crenatum CBS 629.86
Platystomum actinidiae KT 521
Paucispora versicolor KH 110
Pseudopaucispora brunneospora KH 227
Lophiostoma macrostomum KT 635
Abellascoma minimumKT 2013
Dimorphiopsis brachystegiae CPC22679
Biappendiculispora japonica KT 573
Coelodictyosporium pseudodictyosporium MFLUCC13-0451
Lophiostoma semiliberum KT 828
Sigarispora caudata KT 530
Leptoparies palmarum KT 1653
Platystomum compressum MFLUCC13-0343
Paucispora quadrispora KT 843
Parapaucispora pseudoarmatispora KT 2237
Desertiserpenti hydei SQUCC 15092
Guttulispora crataegi MFLUCC14-0993
Capulatispora sagittiformis KT 1934
Vaginatispora amygdali KT 2248
Abellascoma minimumKT 2040
Teichospora trabicola C 134
Lentistoma bipolare KH 214
Pseudolophiostoma obtusisporum KT 2838
Crassiclypeus aquaticusKH 56
Vaginatispora appendiculata MFLUCC16-0314
Neotrematosphaeria biappendiculata KT 1124
Pseudoplatystomum scabridisporum BCC22835
Pseudolophiostoma vitigenum H26930
Lophiopoacea winteri KT 764
Platystomum salicicola MFLUCC15-0632
Lentistoma bipolare KH 222
Neovaginatispora fuckelii KT 634
Vaginatispora aquatica MFLUCC11-0083
Lophiopoacea paramacrostoma MFLUCC11-0463
Guttulispora crataegi MFLUCC13-0442
Crassiclypeus aquaticusKH 185
Paucispora quadrispora KH 448
Neovaginatispora fuckelii KH 161
Teichospora rubriostiolata TR 7
Lophiohelichrysum helichrysi MFLUCC15-0701
Platystomum crataegi MFLUCC14-0925
Lentistoma bipolare KH 216
Pseudolophiostoma tropicum KT 3134
Pseudoplatystomum scabridisporum BCC22836
Coelodictyosporium muriforme MFLUCC13-0351
Neovaginatispora fuckelii CBS101952
Biappendiculispora japonica KT 686-1
Alpestrisphaeria terricola SC-12
*
98/1.0
*
**
*
61/-
75/0.97
99/1.0
*
67/-
78/1.0
87/1.0
99/1.0
81/1.0
96/1.0
*
96/1.0
*
79/0.99
86/1.0
*
68/0.99
*
71/1.0
89/1.0
89/1.0
79/0.99
70/-
*
*
87/1.0
73/-
*
83/-
-/1.0
-/0.95
*
*
*
*
*
*

Fungal Diversity
1 3
GenBank accession numbers: LSU: MW077151; SSU:
MW077160; ITS: MW077142; TEF: MW075769; RPB2:
MW276074.
Notes—Pem etal. (2018) introduced Holmiella juniperi-
cola, which was collected on dead trunks and branches of
Juniperus seravschanica Kom. (as J. zerawschanica Kom.
in the paper) in Uzbekistan. Hitherto, only the sexual morph
of this species was reported and unidentified asexual struc-
tures of chlamydospores were observed. In this study, isolate
SQUCC 15186 producing both sexual and asexual morphs
was collected on J. seravschanica, which is the dominant
dry, temperate tree species of woodlands in the higher
elevations of the Al Jabal Al Akhdar mountains in Oman
(Fig.11). Pem etal. (2018) clearly showed that H. juniperi-
cola differs from H. sabina (type species of the genus) in
having larger ascomata, different numbers of globules per
ascospore cell and the shape of the ascospore apex. They
also compare the sequence data of H. junipericola with the
GenBank isolate of G.M. 2015-04-29.2 named as H. sabina.
However, it is noted that this isolate is not related to the type
material, nor it is linked to a specimen that can be com-
pared with morphology. In future studies, it is essential to
sequence the type species of the genus Holmiella.
Patellaria Fr., Syst. mycol. (Lundae) 2(1): 158 (1822)
amend.
MycoBank: MB 3765
Saprobic on dead wood, stems or rotten paper in terres-
trial habitats. Sexual morph: Ascomata apothecial, super-
ficial, scattered, sessile, longitudinally wide, exposing
the dark hymenium at the centre, circular, flattened, with
a carbonaceous rim, black. Exciple 2-layered, outer layer
pseudoparenchymatous, black, inner layer composed of
thick-walled cells of textura prismatica, gelatinous, heav-
ily pigmented at the base (hypothecium) where cells are
of textura angularis. Hamathecium composed of hyaline,
septate, branched pseudoparaphyses, slightly swollen and
rounded at the apex, forming a dark and thick epithecium
over the asci. Asci 8-spored, bitunicate, fissitunicate, cylin-
drical to clavate, short and slightly curved pedicel, apically
rounded, with an ocular chamber. Ascospores 2–3-seriate
overlapping, clavate to fusiform, trans-septate, hyaline
(description based on Yacharoen etal. 2015). Asexual
morph: Conidiomata pycnidial, erumpent, globose, ostioles
surrounded by brown, septate, setae with obtuse ends. Con-
idiophores lining the inner cavity, hyaline, smooth, septate,
branched, densely aggregated. Conidiogenous cells hyaline,
smooth, subcylindrical, phialidic, percurrent. Conidia hya-
line, smooth, aseptate, guttulate, subcylindrical, obtuse at
apex, truncate at base.
Notes—We amend the generic description of Patellaria
in order to account for the asexual morphological charac-
teristics of P. atrata (= Patellaria quercus).
Patellaria atrata (Hedw.) Fr., Syst. mycol. (Lundae) 2(1):
158 (1822)
amend. ≡ Patellaria quercus Crous & R.K. Schumach.,
in Hernandez-Restrepo etal., Beih. Sydowia 68: 214 (2016);
Fig.12
Saprobic on dead wood, stems or rotten paper in terrestrial
habitats. Sexual morph: Ascomata 655–900 × 180–320μm
(
̄x
= 722 × 240μm; n = 5), apothecial, superficial, scattered,
sessile, closed at first and open at maturity, longitudinally
wide, exposing the dark hymenium at the centre, circular,
flattened, with a carbonaceous rim, black. Exciple 30–50μm
wide, 2-layered, outer layer pseudoparenchymatous, black,
inner layer composed of thick-walled cells of textura pris-
matica, gelatinous, blackish brown at the base (hypothecium)
with cells of textura angularis. Hamathecium composed of
2.5–4μm wide, hyaline, septate, branched pseudoparaphy-
ses, swollen and rounded at the apex, forming a dark and
thick epithecium over the asci. Asci 75–120 × 12–16μm (
̄x
= 86.8 × 14.6μm; n = 10), 8-spored, bitunicate to fissituni-
cate, cylindric-clavate, with a short and slightly curved pedi-
cel, apically rounded, with an ocular chamber. Ascospores
30–40 × 6–8μm (x = 34.7 × 6.8μm; n = 20), 2-seriate over-
lapping, clavate to fusiform, slightly curved, narrowed at
the lower end, 7–9-septate, hyaline. Asexual morph: Con-
idiomata on pine needle agar pycnidial, solitary, black,
erumpent, globose, to 200µm diam., with 1–3 ostioles sur-
rounded by setae, brown, septate, to 40µm long, with obtuse
ends. Conidiophores lining the inner cavity, hyaline, smooth,
base becoming slightly pigmented, 1–5-septate, branched,
densely aggregated, 10–25 × 3–4µm. Conidiogenous cells
hyaline, smooth, subcylindrical with apical taper, phialidic
with periclinal thickening, but also extending percurrent,
5–10 × 2–3µm. Conidia solitary, hyaline, smooth, asep-
tate, guttulate, subcylindrical, apex obtuse, base truncate,
1.5–2µm diam., (3)4(4.5) × 2µm (asexual morph descrip-
tion based on Hernandez-Restrepo etal., 2016).
Material examined—Oman, Al Jabal al-Akhdar
(Green Mountain), from dead branches of unknown host,
28.02.2017, SSN Maharachchikumbura OM113 (SQU
H-117), living culture = SQUCC 15117.
Fig. 8 Phylogenetic tree inferred using the combined sequences of
LSU, SSU, ITS, RPB2 and TEF of the analysed Lophiostomataceae
genera. The tree backbone was constructed using maximum likeli-
hood (ML) analysis. The MLB and PP above 60% and 0.95, respec-
tively, are given at the nodes (MLB/PP). An asterisk (*) indicates
branches with MLB = 100% and PP values = 1.0. Species are fol-
lowed by the strain accession numbers. The new strain is indicated in
blue bold. The scale bar represents the expected number of changes
per site for ML
◂

Fungal Diversity
1 3
GenBank accession numbers: LSU: MW077152; ITS:
MW077143; TEF: MW075770; RPB2: MW276075.
Notes—Patellaria atrata was first described by Hedwig
(1787–1789) and Fries (1822) placed it as the type species
of the newly erected genus Patellaria (lat. patella: cup, bowl;
atra: black) (Unterseher etal. 2003). The genus Patellaria
is characterised by superficial, black, apothecioid ascomata,
with a greenish-black epithecium formed from the branched
and swollen paraphyses, bitunicate, fissitunicate asci, and
hyaline, clavate to cylindrical, up to more than 5-phragmo-
septate ascospores (Kutorga and Hawksworth 1997; Yacha-
roen etal. 2015). Patellaria atrata differs from other species
in having claviform spores (Méndez-Mayboca etal. 2010).
Patellaria atrata is seldom recorded and not mentioned often
in the literature. It decays woods and, interestingly, uses a
type of decaying mode more common in white rot fungi;
0.02
Cryptocoryneum longicondensatum KT2913
Lophiotrema fallopiae KT2748
Cryptoclypeus oxysporus KT2772
Lophiostoma crenatum CBS 629.86
Atrocalyx bambusae RP0030
Pseudotetraploa curviappendiculata HC4930
Anteaglonium abbreviatum ANM925a
Quadricrura septentrionalis HC4983
Cryptocoryneum brevicondensatum yone152
Halotestudina muriformis MFLUCC 17-0395
Angustospora nilensis MFLU 15-1511
Verruculina enalia MFLUCC 17-0406
Verruculina enalia BCC18402
Atrocalyx lignicola CBS 122364
Lophiotrema eburneoides KT1424 1
Aquasubmersa japonica KT2863
Hermatomyces tectonae MFLUCC 14-1141
Aquasubmersa mircensis MFLUCC 11-0401
Montanitestudina hydei SQUCC 15173
Trematosphaeria wegeliniana CBS 123124
Massarina albocarnis CBS 119345
Atrocalyx acutisporus KT2436
Anteaglonium parvulum MFLUCC 14-0821
Muritestudina chiangraiensis MFLUCC 17-2551
Hermatomyces tectonae MFLUCC 14-1142
Xenolophium applanatum CBS 123123
Pseudolophiotrema elymicola KT1450
Ulospora bilgramii CBS 101364
Xenolophium applanatum CBS 123127
Anteaglonium thailandicum MFLUCC 14-0816
Tetraploa sasicola KT563
Triplosphaeria maxima KT870
Anteaglonium parvulum MFLUCC 14-0823
Lepidosphaeria nicotiae CBS 101341
Cryptocoryneum paracondensatum KT3241
Cryptocoryneum japonicum KT3300
Polyplosphaeria fusca KT1616
Pseudocryptoclypeus yakushimensis KT2186
Aquasubmersa japonica KT2862
Hermatomyces thailandicus MFLUCC 14-1144
Anteaglonium parvulum MFLUCC 14-0817
Aquasubmersa japonica KT2813
Hermatomyces tectonae MFLUCC 14-1140
Cryptocoryneum pseudorilstonei CBS 113641
Lophiostoma arundinis CBS 621.86
Anteaglonium parvulum MFLUCC 14-0815
Hermatomyces thailandicus MFLUCC 14-1145
Neotestudina rosatii CBS 690.82
Amniculicola parva CBS 123092
Amniculicola immersa CBS 123083
Anteaglonium globosum ANM925 2
Hermatomyces thailandicus MFLUCC 14-1143
Halotestudina muriformis MFLUCC 18-0392
63/0.97
60/0.98
*
*
79/-
*
86/-
*
*
99/1.00
60/-
*
77/1.0
63/0.97
*
60/-
98/1.0
*
77/-
*
80/0.99
88/1.0
78/0.93
6/0.99
*
*
99/1.00
*
67/-
*
-/0.96
Cryptocoryneaceae
Lophiotremataceae
Hermatomycetaceae
Aquasubmersaceae
Pseudolophiotremataceae
Anteagloniaceae
Tetraplosphaeriaceae
Testudinaceae
Amniculicolaceae
Pseudoastrosphaeriella thailandensis MFLUCC 10-0553
Pseudoastrosphaeriella longicolla MFLUCC 11-0171
Pseudoastrosphaeriella bambusae MFLUCC 11-0205
86/0.99
Pseudoastrosphaeriellaceae
2x
Fig. 9 Phylogenetic tree inferred using the combined sequences of
SSU, LSU, 5.8S, RPB2 and TEF of the analysed Testudinaceae gen-
era. The tree backbone was constructed using maximum likelihood
(ML) analysis. Families are indicated in coloured blocks. The MLB
and PP above 90% and 0.90, respectively, are given at the nodes
(MLB PP). An asterisk (*) indicates branches with MLB = 100% and
PP values = 1.0. Species are followed by the strain accession num-
bers. The new strain is indicated in blue bold. The scale bar repre-
sents the expected number of changes per site for ML.

Fungal Diversity
1 3
generally, ascomycetes are soft rotting fungi (Unterseher
etal. 2003). The asexual morph of the genus Patellaria was
unknown until phoma-like, P. quercus Crous & R.K. Schu-
mach. was described by Hernandez-Restrepo etal. (2016).
Phylogenetic results from the present study showed that P.
quercus forms a sister relationship to our isolate and other
GenBank isolates of P. atrata (Fig.3). This confirms that P.
quercus from twigs of Quercus sp. in Germany is conspecific
with P. atrata. Giving priority to the oldest epithet, here we
synonymised P. quercus under P. atrata. Over 1,200 species
epithets are listed in the Index Fungorum (2020), however,
ten species were accepted in Patellaria by Hawksworth etal.
(1995) and presently P. atrata and ex-type-based sequences
of P. apiculatae Dayarathne & K.D. Hyde, P. chromolaenae
Mapook & K.D. Hyde and P. microspora Ekanayaka & K.D.
are available (Hernandez-Restrepo etal. 2016; Dayarathne
etal. 2020; Hongsanan etal. 2020b; Mapook etal. 2020).
Pleosporales Luttr. ex M.E. Barr, Prodromus to class Locu-
loascomycetes: 67 (1987)
Notes—The taxonomy of Pleosporales has changed rap-
idly in recent years due to additions of numerous families,
genera and species. With 91 families and 614 genera (includ-
ing 48 genera incertae sedis) Pleosporales is recognized as
the largest order in Dothideomycetes (Wijayawardene etal.
2020). These taxa are reported from various habitats in dif-
ferent ecological niches and in this study, we introduce three
new genera and six new species from Oman. The divergence
time for Pleosporales is estimated as 205 Mya (Hongsanan
etal. 2020a).
0.04
Omphalina foliacea Palice 4369
Laetisaria roseipellis CBS 299.82
Leptocorticium tenellum MG143
Galzinia incrustans HHB-12952
Omphalina foliacea Palice 2509
Punctulariopsis subglobispora FCUG 2535
Marchandiomyces corallinus JL128-98
Tretopileus sphaerophorus JCM10092
Vuilleminia pseudocystidiata CBS 347.95
Dendrominia ericae MG162
Erythricium aurantiacum CBS 718.97
Corticium roseum AFTOL-ID 1943
Dendrocorticium polygonioides MG48
Gloeophyllum sepiarium Wilcox 3BB
Dendrominia dryina MG159
Waitea circinata AFTOL-ID 1129
Corticium sp. S. Feusi 05.06.2017
Vuilleminia comedens CBS 428.72
Australovuilleminia coccinea MG74
Vuilleminia comedens T-583
Basidiodesertica hydei SQUCC 15289
Laetisaria nothofagicola JL 261-04
Erythricium aurantiacum JL391-10
Laetisaria marsonii ATCC MYA 4210
Marchandiomyces corallinus ATCC 200796
Cytidia salicina MG49
Corticium roseum EL14 98
Vuilleminia coryli MG136
Giulia tenuis BCC 13066
Erythricium laetum MG72
Punctularia strigosozonata CBS 345.34
98/1.0
60/0.95
60/-
95/1.0
85/1.0
100/-
96/1.0
96/1.0
96/1.0
92/1.0
93/0.99
*
65/0.98
98/1.0
87/0.99
85/0.99
-/0.97
73/1.0
-/0.98
82/1.0
Corticiaceae
Vuilleminiaceae
Punctulariaceae
Dendrominiaceae
*
Fig. 10 Phylogenetic tree inferred using the combined sequences
of SSU, LSU and ITS of the analysed Corticiales families. The tree
backbone was constructed using maximum likelihood (ML) analysis.
Families are indicated in coloured blocks. The order name is indi-
cated to the right of the clade. The MLB and PP above 90% and 0.90,
respectively, are given at the nodes (MLB/PP). An asterisk (*) indi-
cates branches with MLB = 100% and PP values = 1.0. Species are
followed by the strain accession numbers. The new strain is indicated
in blue bold. The scale bar represents the expected number of changes
per site for ML

Fungal Diversity
1 3
Fig. 11 Holmiella junipericola (SQU H-116). a, b, Appearance of
ascomata on host surface. c, d Section through an ascoma. e Perid-
ium. f Pseudoparaphyses. g–i Asci. j–k Ascospores. l Colony on
PDA, m, n Phialides. o Conidia. Scale bars: c, d = 200µm, f = µm 5,
g–i = 50µm, k, o = 25µm, m, n = 10µm

Fungal Diversity
1 3
Didymellaceae Gruyter, Aveskamp & Verkley, Mycological
Research 113 (4): 516 (2009)
Notes—The majority of members in Didymellaceae are
plant associated fungi that can be pathogens across a wide
range of hosts in various habitats, largely causing leaf and
stem lesions, with some of particular relevance for quaran-
tine measures (Wanasinghe etal. 2018). These species are
cosmopolitan and able to adapt to extreme environmental
conditions (Chen etal. 2017). Didymellaceae is one of the
most species-rich families in Pleosporales and in the recent
Outline of Fungi and fungus-like taxa (Wijayawardene etal.
2020) 33 genera are listed in this family. In a recent study,
Huo etal. (2020a) introduced seven genera in Didymellaceae
viz. Dimorphoma L.W. Hou etal.; Ectodidymella L.W. Hou
Fig. 12 Patellaria atrata (SQU H-117). a Apothecia on host substrate. b Section of apothecium. c, d Peridium. e, f Pseudoparaphyses. g–j Asci.
k–r Ascospores. Scale bars: b = 200µm, c, d, g–j = 20µm, e, f, k–r = 10µm

Fungal Diversity
1 3
etal.; Longididymella L.W. Hou etal.; Macroascochyta L.W.
Hou etal.; Paramicrosphaeropsis L.W. Hou etal.; Pseu-
dopeyronellaea L.W. Hou etal.; and Sclerotiophoma L.W.
Hou etal. See Valenzuela-Lopez etal. (2018), Wanasinghe
etal. (2018). Marin-Felix etal. (2019), Tennakoon (2019),
and Hou etal. (2020a, b) for recent updates on this family.
Calophoma Qian Chen & L. Cai, Studies in Mycology 82:
162 (2015)
Notes—Woudenberg etal. (2009) designated an epitype
for Phoma clematidina (Thüm.) Boerema for the isolate col-
lected from the stem of Clematis sp. in the Netherlands and
later Chen etal. (2015) introduced the genus Calophoma
and synonymised P. clematidina as Calophoma clematidina.
Thirteen species epithets are recognised in Calophoma (Huo
etal. 2020) and we introduce the fourteenth species from
this study. This is the first report of Calophoma from Oman.
Calophoma hydei Maharachch., Wanas. &
Al‑Sadi, sp. nov.
MycoBank: MB 837550; Fig.13
Etymology. Named in honour of British mycologist K.D.
Hyde, for his immense contributions to ascomycete
taxonomy.
Saprobic on dead wood of unknown host. Sexual morph:
Not observed. Asexual morph: Conidiomata on PDA pyc-
nidial, solitary, brown, globose, semi immersed or superfi-
cial, ostiolate, 220–550 × 150–330μm, with wall comprising
3–5 layers of brown textura angularis. Micropycnidia pre-
sent. Conidiophores reduced to conidiogenous cells lining
the inner cavity, ampulliform to doliiform, hyaline, smooth,
phialidic, 4–8 × 2–5μm. Conidia ellipsoidal to oblong, hya-
line, smooth, aseptate, guttulate, 4.5–7 × 2–5μm, Conidial
matrix cream.
Culture characteristics—Colonies on PDA, white, flat,
spreading, with moderate aerial mycelium, smooth, undulate
margin, fast growing, reaching 5cm within 7days.
Material examined—OMAN, Al Jabal al-Akhdar (Green
Mountain), from dead wood of unknown host, 22.02.2017,
SSN Maharachchikumbura (SQU H-118, holotype), ex-type
living culture = SQUCC 13750.
GenBank accession numbers: LSU: MW077153; ITS:
MW077144; TUB: MW075776; RPB2: MW276076.
Notes—The phylogenetic analysis based on the combined
sequences of LSU, ITS, TUB and RPB2 shows that C. hydei
is most closely related to C. clematidina, the type species of
the genus Calophoma. Calophoma hydei is distinct from C.
clematidina (Thüm.) Qian Chen & L. Cai in having broader
conidia (4.5–7 × 2–5μm vs 3.5–9 × 2–3.5μm). Calophoma
clematidina produces chlamydospores. However, we did
not observe chlamydospores in C. hydei in cultures. Fur-
ther, there is no report that C. clematidina produces micro-
pycnidia as in C. hydei. Morphological, geographical, and
phylogenetic evidence confirm that C. hydei is a unique
member of the genus Calophoma and deserves species level
recognition.
Didymosphaeriaceae Munk, Dansk botanisk Arkiv 15 (2):
128 (1953)
Notes—The family Didymosphaeriaceae comprises a
miscellaneous group of fungi and currently includes 32 gen-
era (Wijayawardene etal. 2020). Species in this family are
widely distributed among various habitats in different geo-
graphical regions. In this study, we provide the first report
of a Didymosphaeriaceae taxon from Oman.
Dictyoarthrinium S. Hughes, Mycological Papers 48: 29
(1952)
Notes—Dictyoarthrinium is characterized by monon-
ematous or synnematous conidiophores with integrated
conidiogenous cells and septate conidia (Leão-Ferreira and
Gusmão 2010). Members of the genus consist of cruciately
Fig. 13 Calophoma hydei (SQU H-118, holotype). a, b, Appearance of pycnidia on PDA. c, Pycnidium. d Micropycnidia. e Conidia. Scale
bars: a–c = 200μm, d = 10μm, e = 20μm

Fungal Diversity
1 3
septate 4-celled conidia, but the species D. africanum S.
Hughes (Hughes 1952) has 16-celled conidia. The genus
has long been placed in the family Apiosporaceae in the
class Sordariomycetes by various authors (Samarakoon
etal. 2020b). Hughes (1952) compared the morphologi-
cal description of Tetracoccosporium sacchari J.A. Stev.
and type D. quadratum S. Hughes and noted that the two
taxa are similar. This was confirmed by Damon (1953) by
observing the type material of T. sacchari and he trans-
ferred T. sacchari to Dictyoarthrinium as D. sacchari (J.A.
Stev.) Damon. The sequence data derived by Vu etal.
(2019) placed D. sacchari as the heterotypic synonym of
D. quadratum S. Hughes in Dothideomycetes. Samarakoon
etal. (2020b) provided the first detailed molecular investi-
gation of Dictyoarthrinium and confirmed its placement in
Didymosphaeriaceae and our results support their findings.
Hughes (1952) introduced the genus Dictyoarthrinium with
D. quadratum as the type species, which was collected from
Ghana. The genus is distributed worldwide and comprises
ten species, including the newly described species in the
present study.
Dictyoarthrinium hydei Maharachch., Wanas. & Al-Sadi,
sp. nov.
MycoBank: MB 837552; Fig.14
Etymology: Named in honour of British mycologist K.D.
Hyde, for his huge contributions to ascomycete taxonomy.
Saprobic on decaying wood submerged in freshwater
habitats. Sexual morph: Not observed. Asexual morph:
Colonies black, irregular, powdery, up to 3mm broad,
often coalescing. Mycelium composed of septate, branched,
anastomosing, 3–4μm in width pale brown hyphae. Con-
idiophores macronematous, basauxic, cylindrical, flexuous,
subhyaline or pale brown, smooth- and thin-walled, up to
Fig. 14 Dictyoarthrinium hydei (SQU H-119, holotype), a, Appearance of colonies on host surface. b–f, h, i Conidia with conidiophores, g
Colonies on PDA. j–k Conidia. Scale bars: a = 500μm, e, f, h, b–p = 5μm. Scale bar of bapplies to c. Scale bar of d applies to e–f, i

Fungal Diversity
1 3
400μm long, 3–5.5μm wide, terminating in Conidiophore
mother cells. Conidiogenous mother cells cup-shaped, dark
brown, verrucose, give rise to conidiogenous cells. Conid-
iogenous cells integrated, separated by dark-banded septa
(distance between septa 2.5–5.5μm), intercalary and termi-
nal, cylindrical, 2–8μm long, 2–5μm wide. Conidia holo-
blastic, hyaline and spherical when young and brown to dark
brown when mature, thick-walled, verrucose, cruciately sep-
tate with 4-cells, constricted at the septa, 9–17 × 8–13μm,
echinulate, attached to the conidiogenous cells apically or
laterally; apical conidia sessile, borne singly at the apex of
the conidiophore; many lateral conidia, arising from cells of
the conidiophore in single whorls and borne on short stalks.
Culture characteristics—Colonies on PDA slow growing
reaching 6cm diam. in ten days, brownish yellow periph-
ery yellowish, flat, with moderate aerial mycelium, smooth,
undulate margin.
Materials examined—OMAN, Dhofar province, Salalah,
from floating wood in a freshwater stream, 13.10.2016, SSN
Maharachchikumbura OM33b (SQU H-119, holotype), ex-
type living culture SQUCC 13296).
GenBank accession numbers: LSU: MW077154; SSU:
MW077161; ITS: MW077145; TEF: MW075771.
Notes—A new species ofDictyoarthrinium is described
and was determined to be genetically distinct from all other
currently sequenced Dictyoarthrinium in a phylogenetic
analysis based on SSU, LSU, TEF and ITS sequence data.
Phylogenetically, D. hydei closely resembles D. sacchari.
However, it is clearly distinct from D. sacchari (11–13μm)
in having larger conidia (9–17 × 8–13μm) Damon (1953).
Furthermore, D. hydei is geographically distinct from D.
sacchari, which was isolated from decaying terrestrial plant
materials from Brazil, Cuba, Ghana, India, Malaysia, Paki-
stan, Puerto Rico, Spain, States of Micronesia, Thailand,
Venezuela and Zambia (Hughes 1952; Subramanian 1952;
Photita etal. 2003; Saravanan and Vittal 2007; Tarda etal.
2019; Samarakoon etal. 2020b).
Halojulellaceae Suetrong, K.D. Hyde & E.B.G. Jones, Phy-
totaxa 130 (1): 18 (2013)
Notes—Ariyawansa etal. (2013) introduced the family
Halojulellaceae to accommodate the Halojulella avicenniae
(Borse) Suetrong etal. found growing on woody substrata,
especially on the branches of Avicennia marina (Forssk.)
Vierh. The type species H. avicenniae was initially described
as Pleospora avicenniae Borse by Borse (1987). Phoma-
like thin-walled, ostiolate brown pycnidia; filiform, septate,
branched conidiophores and ellipsoidal conidia that are hya-
line, aseptate, thin-walled and guttulate asexual morph is
produced by Halojulella in culture (Hyde 1992; Ariyawansa
etal. 2013). Marine fungi are distributed over 21 families
of Pleosporales (Jones etal. 2015, 2019) and in this study,
we introduce Omania, the second genus in Halojulellaceae.
Omania Maharachch., Wanas. & Al‑Sadi, gen.
nov.
MycoBank: MB 837553
Etymology: The genus was named after the country where
the fungus was collected, Oman.
Saprobic on decaying mangrove roots submerged in
marine habitats. Sexual morph: Not observed. Asexual
morph: Colonies on the PDA superficial to erumpent, dark
brown to grey. Mycelium immersed and superficial. Conidi-
ophores arising from basal cells, septate, branched, cylin-
drical, hyaline, forming conidia laterally and terminally;
conidiophores frequently aggregated into brown stroma.
Conidiogenous cells holoblastic, integrated, determinate,
ampulliform to subcylindrical, smooth to finely verrucu-
lose. Conidia hyaline at immaturity, dark olivaceous-brown
at maturity, smooth to verruculose, guttulate, globose to sub-
globose, thick-walled, mostly solitary, rarely in short chains.
Type: Omania hydei Maharachch., Wanas. & Al-Sadi.
Omania hydei Maharachch., Wanas. &
Al‑Sadi, sp. nov.
MycoBank: MB 837554; Fig.15
Etymology: Named in honour of British mycologist K.D.
Hyde, for his huge contributions to ascomycete taxonomy.
Saprobic on decaying, submerged Avicenia marina in
marine habitats. Sexual morph: Not observed. Asexual
morph: Conidiophores arising from basal cells, cylindri-
cal, hyaline, forming conidia laterally and terminally, fre-
quently aggregated into a brown stroma, septate, branched,
up to 250μm long, 2.5–5.0 μm wide. Conidiogenous
cells 2.0–5.0 × 1.5–3.0 μm, integrated, ampulliform to
subcylindrical, smooth to finely verruculose. Conidia
7.0–11.0 × 7.0–10.5µm, (
̄x
= 9.5 × 9.2µm, n = 30), hyaline
at immaturity, dark olivaceous-brown at maturity, smooth to
verruculose, guttulate, globose to subglobose, thick-walled
(0.3 − 1.0µm wide), mostly solitary, rarely in short chains.
Culture characteristics—Colonies on the PDA were
slow growing reaching 5cm diam. in ten days, superfi-
cial to erumpent, dark brown to grey, with moderate aerial
mycelium, undulate margin.
Material examined—OMAN, Muscat, Al Qurum, from
dead root of Avicenia marina, 11.10.2017, SSN Maharach-
chikumbura & AM Al-Sadi H26 (SQU H-120, holotype),
ex-type living culture = SQUCC 3026.

Fungal Diversity
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GenBank accession numbers: LSU: MW077155; SSU:
MW077162; ITS: MW077146; TEF: MW075772; RPB2:
MW276077.
Notes—Omania hydei, isolated from dead roots of Avi-
cenia marina collected from the Al Qurum beach in Mus-
cut, Oman, is clearly distinct from Halojullela in having
an asexual morph that is not arthrinium-like. Halojulel-
laceae is sister to the monotypic marine family Ascocylin-
dricaceae, which includes the type species Ascocylindrica
marina Abdel-Wahab etal., collected from Saudi Arabian
mangroves (Ariyawansa etal. 2015). Ascocylindrica marina
is characterised by small ascomata, cylindrical asci and bi-
celled dark brown to black ascospores (Ariyawansa etal.
2015). We were unable to compare the morph of Omania
hydei with A. marina as O. hydei is only reported as the
asexual form. Further fresh collections and DNA based
sequence data are warranted to understand the exact phylo-
genetic relationship between these two families.
Lophiostomataceae Sacc., Sylloge Fungorum 2: 672 (1883)
Notes—Members of Lophiostomataceae are mainly sap-
rotrophs found in woody and herbaceous plants in terrestrial,
freshwater and marine habitats (Hashimoto etal. 2018). The
family was erected by Nitschke (1869), with Lophiostoma
macrostomum (Tode) Ces. & De Not. as the type species.
Lophiostomataceae are highly diverse and characterized by
immersed to erumpent, carbonaceous to coriaceous asco-
mata with rounded or slit-like ostioles, mostly clavate asci
and uni- to multi-septate, hyaline to dark brown ascospores
with terminal appendages or mucilaginous sheaths (Tham-
bugala etal. 2015; Bao etal. 2019). Here we introduce a
new hyphomycetous genus Desertiserpentica to the family
Lophiostomataceae.
Desertiserpentica Maharachch., Wanas. &
Al‑Sadi, gen. nov.
MycoBank: MB 837555
Etymology: The generic epithet is from the combination
of two words “desertum” (desert) for the habitat where the
fungus was isolated and “serpens” (Latin: crawling, snake),
which refers to the somewhat snake like conidia.
Colonies on substratum superficial, effuse, hairy or vel-
vety, dark brown, grey or black. Mycelium partly superficial,
partly immersed, composed of branched, septate, smooth,
pale brown hyphae. Sexual morph: Not observed. Asexual
morph: Hyphomycetous. Conidiophores macronematous,
mononematous, simple, cylindrical, unbranched, smooth,
olive-green to brown, septate, without percurrent extensions.
Conidiogenous cells monoblastic, integrated, terminal,
cylindrical or doliiform, determinate, olive-green to brown.
Conidia acrogenous, solitary, flexuous, dry, pale brown to
brown, distoseptate, long cylindrical with rounded apex.
Type: Desertiserpentica hydei Maharachch., Wanas. &
Al-Sadi
Fig. 15 Omania hydei (SQU H-120, holotype) a, b Colonies on PDA. c–h Conidiophores, conideogenous cells and conidia. i Conidia. Scale
bars: c = 50μm, d, f, h = 20μm. Scale bar of d applies to e. Scale bar of f applies to g. Scale bar of h applies to i

Fungal Diversity
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Desertiserpentica hydei Maharachch., Wanas.
& Al‑Sadi, sp. nov.
MycoBank: MB 837556; Fig.16
Etymology: Named in honour of British mycologist K.D.
Hyde, for his huge contributions to ascomycete taxonomy.
Colonies growing saprotrophically on natural substrate,
superficial, effuse, hairy or velvety, dark brown, grey or
black. Mycelium partly superficial, partly immersed, com-
posed of branched, septate, smooth, pale brown hyphae.
Sexual morph: Not observed. Asexual morph: Hypho-
mycetous. Conidiophores macronematous, mononematous,
simple, cylindrical, septate, branched, smooth, brown, sep-
tate, without percurrent extensions. Conidiogenous cells
monoblastic, integrated, terminal, cylindrical or dolii-
form, determinate, brown. Conidia up to 1,100μm long,
9.5–15.0μm wide, acrogenous, solitary, flexuous, dry, pale
brown to brown, distoseptate 8–14μm, long cylindrical
with rounded apex.
Colonies on PDA moderately growing, reaching 4cm
diam. after seven days at 25°C, circular, white to olive-
green and margin orange or dark brown, with moderate
aerial mycelium, margin undulate. Mycelium partly super-
ficial, partly immersed, composed of branched, septate,
smooth, pale brown hyphae. Sexual morph: Not observed.
Asexual morph: Conidiophores and Conidiogenous cells
similar to those on natural substrate. Conidia up to 400μm
long, 8.0–10.0μm wide, acrogenous, rostrate, sometimes
furcate with several arms, olive-green to brown, distoseptate
8–14μm, long and cylindrical with a rounded apex.
Material examined—OMAN, Nizwa, from dead wood,
25.09.2016, SSN Maharachchikumbura OM21 (SQU H-121,
holotype), ex-type living culture = SQUCC 15092.
GenBank accession numbers: LSU: MW077156; SSU:
MW077163; ITS: MW077147; TEF: MW075773.
Fig. 16 Desertiserpentica hydei (SQU H-121, holotype) a–c Appear-
ance of colonies on host surface. d–f Acrogenous, solitary, flexuous,
distoseptate conidia on natural substrate. g Colonies on PDA. h–m
Conidiophores, conidiogenous cells and conidia on PDA. Scale bars:
c = 500μm, d, h, k, m = 100μm, e = 20 μm, i, j = 50μm. Scale bar of
e applies to f. Scale bar of k applies to l

Fungal Diversity
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Notes—Desertiserpentica is allied to Dimorphiopsis
(Fig.4), an asexual, unusual monotypic genus associated
with leaves of Brachystegia spiciformis (Fabaceae) col-
lected from Zambia (Crous etal. 2013). Crous etal. (2013)
were not sure if Brachystegia was a coelomycete or hypho-
mycete because in pine needle agar it was a hyphomycete,
whereas in water agar it behaved like a coelomycete with
immersed conidiomata. Dimorphiopsis is characterised by
pale to brown, septate, branched hyphae, varying conidio-
mata from immersed pycnidia to superficial sporodochia and
solitary, pale brown to dark brown, 1-distoseptate conidia
(Crous etal. 2013). Thambugala etal. (2015) introduced
Amorosiaceae, as sister to the family Lophiostomataceae.
Two genera, viz. Amorosia Mantle & D. Hawksw. and
Angustimassarina Thambugala etal. were accommodated in
Amorosiaceae and both genera have hyphomycetous asexual
morphs but are clearly distinct from Desertiserpentica.
Phaeosphaeriaceae M.E. Barr, Mycologia 71(5): 948
(1979)
Notes —Phaeosphaeriaceae was erected by Barr (1979)
to include 15 genera. It is one of the largest heterogeneous
families currently recognised in the order Pleosporales and
consists of 82 genera (Wijayawardene etal. 2020). The fam-
ily is based on Phaeosphaeria I. Miyake which was typified
by P. oryzae I. Miyake. Most of the species and genera of
Phaeosphaeriaceae are saprobes or endophytes. However,
some members are well known phytopathogens that infect
economically important cereal crops such as barley, wheat
and rye (Cunfer 2000; Quaedvlieg etal. 2013; Karunar-
athna etal. 2017; Phookamsak etal. 2017; Wanasinghe
etal. 2018; Maharachchikumbura etal. 2019). Members of
the Phaeosphaeriaceae are ubiquitous in soils, freshwater
and marine ecosystems and even cause a variety of human
diseases (Ahmed etal. 2017). Species of Phaeosphaeriaceae
are characterised by immersed to superficial ascomata with
globose to subglobose, short-papillate ostiole, bitunicate
asci, and hyaline, yellowish or brown, uni- or multi-septate
or muriform ascospores (Shoemaker 1984; Shoemaker and
Babcock 1992; Phookamsak etal. 2017). In the present con-
tribution, we introduce a third species to the genus Hydeo-
myces Maharachch. etal. from Oman.
Hydeomyces Maharachch., H.A. Ariyaw., Wanas. & Al-
Sadi, Phytotaxa 391(1): 33 (2019)
Notes—The genus Hydeomyces was introduced by Maha-
rachchikumbura etal. (2019), for the type species H. deserti-
pleosporoides Maharachch. etal., which was collected from
Juniperus excels M. bieb. in the Al Jabal al-Akhdar moun-
tains, Oman. The second species H. pinicola J.F. Zhang
etal., a fungus collected from the dead wood of Pinus sp.
in Guizhou Province of China, was added to the genus by
Zhang etal. (2019). The genus is characterised by partly
immersed to erumpent ascomata, heavily pigmented, 2–5
layered peridium, 8-spored, bitunicate, fissitunicate, cylin-
drical asci, uniseriate, ellipsoidal to subfusiform, muriform
ascospores and phoma-like asexual morph (Maharachchi-
kumbura etal. 2019).
Hydeomyces hydei Maharachch., Wanas. &
Al‑Sadi, sp. nov.
MycoBank: MB 837557; Fig.17
Etymology: Named in honour of British mycologist K.D.
Hyde, for his huge contributions to ascomycete taxonomy.
Saprobic on dead woods. Sexual morph: Ascomata
220–280µm high, 250–320µm diam. (
̄x
= 248.8 × 276.2µm,
n = 10), black, immersed, scattered, slightly erumpent,
globose, ostiolate. Ostiole central, short, slightly raised,
smooth, ostiolar canal filled with brown cells. Peridium
12–20µm wide at the base, 30–40µm wide at the apex,
comprising 4–5 layers, with heavily pigmented, thick-
walled, comprising dark brown cells of textura angularis.
Hamathecium comprising numerous, 2–3µm (n = 30) wide,
filamentous, branched, septate, guttulate, pseudoparaphyses.
Asci 110–150 × 13–17 µm (
̄x
= 127.3 × 14.5µm, n = 40),
8-spored, bitunicate, fissitunicate, cylindrical, pedicellate
(15–30µm long), rounded at apex with an ocular chamber.
Ascospores 18–22 × 9.5–12µm (
̄x
= 20 × 10.3µm, n = 50),
overlapping uniseriate, muriform, ellipsoidal, 4 − 5-trans-
versely septate, with one vertical septum, deeply constricted
at middle septum, initially hyaline, becoming brown at matu-
rity, rounded at both ends, not surrounded by a mucilaginous
sheath. Asexual morph: Not observed.
Culture characteristics—Colonies on the PDA slow grow-
ing reaching 2cm diam. in four weeks, irregular, superficial
to erumpent, blackish-green to black, lacking aerial myce-
lium, top surface raised centrally and wrinkled, undulate
margin, reverse black.
Material examined—OMAN, Al Jabal al-Akhdar (Green
Mountain), from dead wood of unknown host, 06.2017, SSN
Maharachchikumbura (SQU H-122, holotype), ex-type liv-
ing culture = SQUCC 15290.
GenBank accession numbers: LSU: MW077157; SSU:
MW077164; ITS: MW077148; TEF: MW075774; RPB2:
MW276078.
Notes—Hydeomyces hydei shares similar morphologi-
cal features with H. desertipleosporoides (type species),
such as globose ascomata, cylindrical asci and muriform
ascospores. Phylogenetically these two taxa constitute a
monophyletic clade with absolute support (100% MLB, 1.00
PP, Fig.6). However, there are noteworthy morphological
differences between them. Hydeomyces desertipleosporoides

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Fig. 17 Hydeomyces hydei (SQU H-122, holotype). a, b Ascomata on host substrate. c Section of ascoma. d Closeup of ostiole. e Peridium. f
Pseudoparaphyses. g–j Asci. k–p Ascospores. Scale bars: c = 100µm, d, e, g–j = 20µm, f k–p = 10µm

Fungal Diversity
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has semi-immersed to erumpent ascomata, short pedicellate
asci, 2−4-transversely septate ascospores with conical lower
ends, whereas H. hydei has immersed ascomata, 15–30µm
long pedicellate asci, and 4 − 5-transversely septate
ascospores that have rounded end cells. The size of the asci
and ascospores of H. hydei (127.3 × 14.5µm, 20 × 10.3µm
correspondingly) are comparatively larger than those are of
H. desertipleosporoides (75 × 12.5µm, 13.5 × 5.8µm). Due
to these morphological variations, we introduce Hydeomyces
hydei sp. nov. for SQUCC 15290.
Testudinaceae Arx, Persoonia 6 (3): 366 (1971)
Notes—Testudinaceae was introduced by Von Arx (1971)
to accommodate Argynna Morgan, Lepidosphaeria, Neotes-
tudina, Pseudophaeotrichum Arx, E. Müll. & C. Stoll and
Testudina Bizz. based on the presence of ascomata with a
dark peridium, bitunicate asci and dark, 2-celled ascospores
as significant delimiting characters at the familial level
(Wanasinghe etal. 2017). From a morphological perspec-
tive, the genera of Testudinaceae appear to have different
morphologies and its intergeneric classification is still poorly
understood. Currently, there are nine genera accepted in Tes-
tudinaceae and in this study, we introduce Montanitestudina,
a novel genus for a fungus collected from Al Jabal al-Akhdar
mountains in Oman.
Montanitestudina Maharachch., Wanas. &
Al‑Sadi, gen. nov.
MycoBank: MB 837558
Etymology: The generic epithet is from the combination
of two words “montānus” (mountain) habitat where the
fungus was isolated and “testudina” similar to Testudina.
Saprobic on dead wood of unknown plants. Sexual morph:
Ascomata scattered or sometimes gregarious beneath the
host periderm or on decorticated wood, coriaceous, black,
globose to subglobose, ostiolate. Ostiole central, papillate,
with an irregular, pore-like opening. Peridium composed
of 4–5-layers with brown to dark-brown, cells of textura
angularis fusing the host tissues. Hamathecium comprising
septate, cellular pseudoparaphyses. Asci 8-spored, bituni-
cate, fissitunicate, cylindrical to cylindric-clavate, with a
distinct pedicel, apically rounded, with an ocular chamber.
Ascospores overlapping uniseriate, brown, ellipsoid, oblong
to fusoid, 3–5-transversely septate, with 1–2-longitudinal
septa, muriform, smooth-walled, with or without surrounded
by a mucilaginous sheath. Asexual morph: Not observed.
Type: Montanitestudina hydei Maharachch., Wanas. &
Al-Sadi
Notes—Montanitestudina morphologically resembles
some of the species in Camarosporidiella Wanas. etal.,
Cucurbitaria Gray, Fenestella Tul. & C. Tul., Hawkswor-
thiana U. Braun, Marjia Wanas. etal., Neocucurbitaria
Wanas. etal., Pseudostrickeria Q. Tian etal., Sporormuris-
pora Wanas. etal., and Uzbekistanica Wanas. etal. by its
cylindrical asci and uniseriate, brown, muriform ascospores
(Wanasinghe etal. 2017, 2018; Jaklitsch etal. 2018).
Although there is some morphological overlap between
Montanitestudina and the above-mentioned genera, they are
phylogenetically apart from Montanitestudina in multi-gene
phylogenetic analyses. Montanitestudina is phylogenetically
closely related to Lepidosphaeria (Fig.8). However, this is
strongly supported only by Bayesian analysis. There are sub-
stantial morphological differences between these two taxa to
warrant generic ranks.
Montanitestudina hydei Maharachch., Wanas.
& Al‑Sadi, sp. nov.
MycoBank: MB 837560; Fig.18
Etymology: Named in honour of British mycologist K.D.
Hyde, for his huge contributions to ascomycete taxonomy.
Saprobic on dead wood of unknown plants. Sexual
morph: Ascomata 480–550µm high, 440–530µm diam.
(
̄x
= 529.4 × 500.1 µm, n = 10), black, immersed, scat-
tered, sometimes gregarious beneath the host periderm
or on decorticated wood, partly erumpent, globose, ostio-
late. Ostiole central, short, slightly raised, smooth, ostiolar
canal filled with hyaline cells. Peridium 35–50µm wide,
comprising 4–5 layers, with outer layer heavily pigmented,
thick-walled, comprising dark brown cells of textura angu-
laris, cells towards inside lighter, with inner layer com-
posed of pale brown to hyaline, flattened, thin-walled cells
of textura angularis. Hamathecium comprising numerous,
1.5–2.5µm (n = 30) wide, filamentous, branched, septate,
guttulate, pseudoparaphyses. Asci 115–130 × 11–15µm
(
̄x
= 120.6 × 12.6µm, n = 40), 8-spored, bitunicate, fissituni-
cate, cylindrical, pedicellate, rounded at apex with an ocular
chamber. Ascospores 14–18 × 7–10µm (
̄x
= 16.1 × 7.8µm,
n = 50), overlapping uniseriate, muriform, mostly ellipsoidal,
3 − 4-transversely septate, with 1 − 2 vertical septa, slightly
constricted at middle septum, initially hyaline, becoming
brown at maturity, upper end rounded, lower end conical,
not surrounded by a mucilaginous sheath. Asexual morph:
Not observed.
Culture characteristics—Colonies on PDA slow grow-
ing reaching 2cm diam. in four weeks, irregular, superfi-
cial to erumpent, blackish-green to black, lacking aerial

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Fig. 18 Montanitestudina hydei (SQU H-123, holotype). a, b Ascomata on host substrate. c Section of ascoma. d Peridium. e Pseudoparaphy-
ses. f–k Asci. l–q Ascospores. r, s Culture on PDA (note s reverse). Scale bars: c = 200µm, d, f–k = 20µm, e, l–q = 10µm

Fungal Diversity
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mycelium, top surface raised centrally and wrinkled, undu-
late margin, reverse black.
Material examined—OMAN, Al Jabal al-Akhdar
(Green Mountain), from dead wood of unknown host,
25.09.2017, SSN Maharachchikumbura (SQU H-123,
holotype), ex-type living culture = SQUCC 15173.
GenBank accession numbers: LSU: MW077158; SSU:
MW077165; ITS: MW077149; TEF: MW075775.
Notes—Montanitestudina hydei is similar to Murites-
tudina chiangraiensis Wanas etal. in Testudinaceae due
to its globose ascomata with a thin peridium and muri-
form ascospores but differs by having biseriate, hyaline
ascospores with large guttules whereas M. hydei has uni-
seriate, brown ascospores without guttules. Also, these
genera are phylogenetically separated in multi-gene phy-
logenetic analyses (Fig.8). Montanitestudina hydei and
Lepidosphaeria nicotiae Parg.-Leduc are monophyletic in
the multigene phylogenetic analyses. However, they are
separated by morphological features. Lepidosphaeria nico-
tiae has globose to subglobose ascomata, clavate asci and
1-septate ascospores with a granulated surface whereas M.
hydei has globose ascomata, cylindrical asci and muriform
ascospores with a smooth surface.
Basidiomycota Whittaker ex R.T. Moore, Botanica Marina
23 (6): 371 (1980)
Notes—The Basidiomycota is the second largest phy-
lum of the kingdom Fungi after Ascomycota and includes
approximately one third of all fungi (Taylor etal. 2014).
Four subphyla, 18 classes and 68 orders are presently rec-
ognized in Basidiomycota (He etal. 2019). Members of
the Basidiomycota can be mushrooms, smuts or rusts and
are characterized by a club-shaped fertile structure in their
sexual stage, the basidium, on which meiospores (basidi-
ospores) are produced. Members of the Basidiomycota
are involved in ecosystem functioning and are the major
degraders of wood and some are major crop and cereal
plant pathogens (Taylor etal. 2014).
Agaricomycetes Doweld, Prosyllabus Tracheophytorum,
Tentamen Systematis Plantarum Vascularium (Tracheo-
phyta) (Moscow): LXXVIII (2001)
Notes—Members of the Agaricomycetes are generally
mushroom-forming fungi and are found commonly in ter-
restrial and aquatic habitats (Hibbett etal. 2014). They are
responsible for decaying wood, some are well known plant
pathogens, and some function as mutualists (Hibbett etal.
2014). The class includes most of the edible mushrooms
and the taxa are included in Agaricomycetes, which were
previously included in Homobasidiomycetes (Hibbett and
Thorn 2001).
Corticiales K.H. Larss., Mycological Research 111 (5): 540
(2007)
Notes—The order Corticiales was introduced by Hibbett
etal. (2007) based on phylogeny for the taxa previously in
the Vuilleminiales, the Dendrocorticium group, and the cor-
ticioid group (Hibbett etal. 2014). The four families viz.
Corticiaceae, Dendrominiaceae, Punctulariaceae and Vuil-
leminiaceae are presently recognised in Corticiales (Wijaya-
wardene etal. 2020). Most of the members are resupinate
species that produce smooth hymenophores, with or with-
out clamps, and smooth basidiospores (Hibbett etal. 2014)
and some species are only known from their asexual states.
Members of the Corticiales are saprotrophs, plant patho-
gens, lichen pathogens, and lichenized species (Lawrey etal.
2008; Hibbett etal. 2014; Diederich etal. 2018).
Corticiaceae Herter, Kryptogamen-Flora der Mark
Brandenburg 6 (1): 70 (1910)
Notes—Ecologically and phenotypically Corticiaceae is
the most diverse family in Corticiales that includes a sexual
morph with corticioid type fruiting bodies and coelomycetes
and hyphomycetes asexual genera with diverse nutritional
modes. The genera Capillosclerotium Prameela & Deeba,
Corticirama Pilát, Corticium Pers., Erythricium J. Erikss.
& Hjortstam, Galzinia Bourdot, Giulia Tassi, Laetisaria
Burds., Lawreymyces Lücking & Moncada, Marchandio-
myces Diederich & D. Hawksw., Necator Massee, Tretopi-
leus B.O. Dodge and Waitea Warcup & P.H.B. Talbot are
presently recognized in Corticiaceae (Hibbett etal. 2014).
Species of the family are saprobes, plant pathogens, myco-
parasites, lichenized or lichenicolous taxa (Jayawardena
etal. 2019).
Basidiodesertica Maharachch., Wanas. & Al-Sadi, gen. nov.
MycoBank: MB 837561
Etymology. The species epithet is from the combination
of two words “Basidio” (belongs to Basidiomycetes) and
“desertum” (desert). Saprobic on dead leaves of unknown
plants. Asexual morph: Synnemata arising from stromatic
base, erect, cylindrical to subulate, solitary to scattered,
brownish, wide and foot-like at the very base. Conidi-
ophores of synnema parallelly compacted, flexuous, hya-
line to brown, septate with or without clamp connections,
branched. Conidiogenous cells holoblastic, integrated, ter-
minal, cylindrical. Conidia dictyosporous bulbil-like, mul-
ticellular, oblong to fusiform, hyaline when young, brown
grey to dark brown violet at maturity, with several (4–6)
holoblastic conidiogenous cells contributing to development
of the conidium.
Notes—The majority of hyphomycete and coelomycet-
ous fungi have sexual states in the Ascomycota (Kirk etal.

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Fungal Diversity
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2008). Presently 13 basidiomycete genera are identified with
coelomycetous asexual morphs (Rungjindamai etal. 2008).
Some of these genera are phylogenetically linked to their
respective basidiomycete groups. For example, based on
the sequence data, Mycotribulus Nag Raj & W.B. Kendr.
is phylogenetically affiliated with Physalacriaceae (Agari-
cales), Chaetospermum Sacc. to Sebacinaceae (Sebacina-
les) whereas monotypic Giulia Tassi is well-placed in the
Corticiaceae (Corticiales) and Basidiopycnis Oberw. etal.
and Proceropycnis M. Villarreal etal. are related to Hoeh-
nelomycetaceae (Atractiellales) (Oberwinkler etal. 2006;
Rungjindamai etal. 2008; Tangthirasunun etal. 2014).
The order Corticiales is characterized by effused or discoid
basidiomata, a smooth hymenophore, and a monomitic
hyphal system with clamped, rarely single septate hyphae
(Hibbett etal. 2007). However, the family Corticiaceae
is linked to coelomycetous and huphomyceteous asexual
morphs, which are not typical members for the Basidi-
omycetes. Here we introduce a new hyphomycetes genus,
Basidiodesertica, based on molecular phylogenetic results
and distinct morphological characters.
Type: Basidiodesertica hydei Maharachch., Wanas. &
Al-Sadi
Basidiodesertica hydei Maharachch., Wanas. & Al-Sadi,
sp. nov.
MycoBank: MB 837562; Fig.19
Etymology. Named in honour of British mycologist K.D.
Hyde, for his huge contributions to fungal taxonomy.
Saprobic on dead leaves of unknown plants. Asexual
morph: Synnemata arising from stromatic base, erect,
cylindrical to subulate, solitary to scattered, brownish
800–1600 × 40–80µm, widened, foot-like at the base. Conid-
iophores macronemeatous, compact and parallel in synnema,
2.5–5µm diam., flexuous, hyaline to brown, septate, with/
without clamp connections. Conidiogenous cells holoblastic,
integrated and terminal, cylindrical, 6–12 × 4–8µm. Several
of a moderate to dark brown, Conidia dictyosporous bulbil-
like 50–80 × 20–25µm (
̄x
= 63.4 × 22.9µm, n = 30), mul-
ticellular, oblong to fusiform, hyaline when young, brown
grey to brown violet when mature, with several (4–6) holo-
blastic conidiogenous cells contributing to development of
conidium.
Culture characteristics—Colonies on the PDA slow
growing reaching 2cm diam. in four weeks, irregular,
superficial to erumpent, blackish-green to black, lacking
aerial mycelium, top surface raised centrally and wrinkled,
undulate margin, reverse black.
Material examined—OMAN, Sur, Wadi Shab, from dead
leaves of unknown plants, 07.2016, SSN Maharachchikum-
bura OM 01 (SQU H-124, holotype), ex-type living cul-
ture = SQUCC 15289.
GenBank accession numbers: LSU: MW077159; SSU:
MW077166; ITS: MW077150.
Notes—Phylogenetically Basidiodesertica hydei shows
affinities to the coelomycetous monotypic and saprobic
genus Giulia which was typified by G. tenuis (Tassi 1904),
and hyphomycetes Tretopileus sphaerophorus (Okada 1998).
Giulia tenuis is characterised by dark brown to black conidi-
omata, hyaline, phialidic, ampulliform to conical, smooth-
walled conidiogenous cells and hyaline, cylindrical, asep-
tate conidia bearing flexuous, unbranched, tubular, apical
appendages (Li etal. 2020). Tretopileus sphaerophorus has
been reported from Cuba, USA, Sierra Leone, Indonesia and
Thailand (Berkeley 1868; Deighton 1960; Weilbacher 1970;
Okada 1998). This synnematous hyphomycete produced first
a bulbil as a propagule on the top of a synnema and when
the bulbil has fallen, the synnema proliferated about seven
times to produce new bulbils, each time making conspicuous
nodes at the upper part (Okada 1998). Based on phenotype
and sequence data, Basidiodesertica is clearly distinct from
the members of the Giulia and Tretopileus.
Discussion
‘Discomycetes’ is an artificial term for apothecia-produc-
ing fungi. The discoid fruiting bodies are found among ten
classes of Ascomycota (Johnston etal. 2014; Ekanayaka
etal. 2017). The orders Catinellales, Eremithallales, Hys-
teriales, Mytilinidiales, Patellariales and Pleosporales of
the class Dothideomycetes produce apothecial ascomata.
Among them, Patellariales was erected by Hawksworth and
Eriksson (1986), comprising the family Patellariaceae. The
taxonomic circumscriptions of Patellariaceae have rapidly
changed. Subsequently, several genera were included in
Patellariaceae while some were transferred to different tax-
onomic groups (Zhang and Hyde 2009). For instance, nine
(Saccardo 1884), 21 (Lindau 1897), 33 (Saccardo 1899),
30 (Boudier 1907), 20 (Ramsbottom 1912), 35 (Clements
and Shear 1931), two (Butler 1940), nine (Luttrell 1973),
13 (von Arx 1975), 16 (Barr 1979), 12 (Kutorga and Hawk-
sworth 1997), 15 (Lumbsch and Huhndorf 2007), 13 (Hyde
etal. 2013), 13 (Yacharoen etal. 2015), 18 (Ekanayaka etal.
2017), 21 (Wijayawardene etal. 2020) and 21 (Hongsanan
etal. 2020b) genera were accepted in Patellariaceae. With
Fig. 19 Basidiodesertica hydei (SQU H-124, holotype). a Synne-
mata on natural host surface. b, c Three-day-old culture on PDA
(note c reverse). d, e Two-week-old culture on PDA (note e reverse). f
Synnemata and conidia on PDA. g–i Synnemata. j Hyphae of synne-
mata k–o Conidiophore and conidiogenous cells. p–v Bulbils at dif-
ferent maturities. Scale bars: f, g = 200 µm, h–k, n–v = 20 µm, l,
m = 50µm
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