Fungal Diversity (2020) 105:319–575
https://doi.org/10.1007/s13225-020-00463-5
Freshwater Dothideomycetes
Wei Dong1,2,3,4,5 · Bin Wang6 · Kevin D. Hyde4,5,7,8 · Eric H. C. McKenzie9 · Huzefa A. Raja10 · Kazuaki Tanaka11 ·
Mohamed A. Abdel‑Wahab12 · Faten A. Abdel‑Aziz12 · Mingkwan Doilom5,13,14,15 · Rungtiwa Phookamsak8,13,14,15 ·
Sinang Hongsanan16 · Dhanushka N. Wanasinghe13,14,15 · Xian‑Dong Yu1,17 · Gen‑Nuo Wang18 · Hao Yang1 ·
Jing Yang4,19 · Kasun M. Thambugala20 · Qing Tian4,7 · Zong‑Long Luo21 · Jian‑Bo Yang13,14,15 · Andrew N. Miller22 ·
Jacques Fournier23 · Saranyaphat Boonmee4,19 · Dian‑Ming Hu24 · Sarunya Nalumpang2,3 · Huang Zhang1,25,26
Received: 12 June 2020 / Accepted: 8 October 2020/ Published online: 28 December 2020
© The Author(s) 2020
Abstract
Freshwater Dothideomycetes are a highly diverse group of fungi, which are mostly saprobic in freshwater habitats worldwide. They are important decomposers of submerged woody debris and leaves in water. In this paper, we outline the genera
of freshwater Dothideomycetes with notes and keys to species. Based on multigene analyses and morphology, we introduce
nine new genera, viz. Aquimassariosphaeria, Aquatospora, Aquihelicascus, Fusiformiseptata, Neohelicascus, Neojahnula,
Pseudojahnula, Purpureofaciens, Submersispora; 33 new species, viz. Acrocalymma bipolare, Aquimassariosphaeria kunmingensis, Aquatospora cylindrica, Aquihelicascus songkhlaensis, A. yunnanensis, Ascagilis submersa, A. thailandensis,
Bambusicola aquatica, Caryospora submersa, Dictyocheirospora thailandica, Fusiformiseptata crocea, Helicosporium thailandense, Hongkongmyces aquaticus, Lentistoma aquaticum, Lentithecium kunmingense, Lindgomyces aquaticus, Longipedicellata aquatica, Neohelicascus submersus, Neohelicomyces dehongensis, N. thailandicus, Neohelicosporium submersum,
Nigrograna aquatica, Occultibambusa kunmingensis, Parabambusicola aquatica, Pseudoasteromassaria aquatica, Pseudoastrosphaeriella aquatica, Pseudoxylomyces aquaticus, Purpureofaciens aquatica, Roussoella aquatica, Shrungabeeja
aquatica, Submersispora variabilis, Tetraploa puzheheiensis, T. yunnanensis; 16 new combinations, viz. Aquimassariosphaeria typhicola, Aquihelicascus thalassioideus, Ascagilis guttulaspora, A. queenslandica, A. seychellensis, A. sunyatsenii,
Ernakulamia xishuangbannaensis, Neohelicascus aquaticus, N. chiangraiensis, N. egyptiacus, N. elaterascus, N. gallicus,
N. unilocularis, N. uniseptatus, Neojahnula australiensis, Pseudojahnula potamophila; 17 new geographical and habitat
records, viz. Aliquandostipite khaoyaiensis, Aquastroma magniostiolata, Caryospora aquatica, C. quercus, Dendryphiella
vinosa, Ernakulamia cochinensis, Fissuroma neoaggregatum, Helicotruncatum palmigenum, Jahnula rostrata, Neoroussoella bambusae, N. leucaenae, Occultibambusa pustula, Paramonodictys solitarius, Pleopunctum pseudoellipsoideum,
Pseudocapulatispora longiappendiculata, Seriascoma didymosporum, Shrungabeeja vadirajensis and ten new collections
from China and Thailand, viz. Amniculicola guttulata, Aquaphila albicans, Berkleasmium latisporum, Clohesyomyces aquaticus, Dictyocheirospora rotunda, Flabellascoma fusiforme, Pseudoastrosphaeriella bambusae, Pseudoxylomyces elegans,
Tubeufia aquatica and T. cylindrothecia. Dendryphiella phitsanulokensis and Tubeufia roseohelicospora are synonymized
with D. vinosa and T. tectonae, respectively. Six orders, 43 families and 145 genera which belong to freshwater Dothideomycetes are reviewed. Of these, 46 genera occur exclusively in freshwater habitats. A world map illustrates the distribution
of freshwater Dothideomycetes.
Keywords 42 new taxa · Freshwater distribution · Phylogeny · Submerged wood · Taxonomy
Introduction
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s13225-020-00463-5) contains
supplementary material, which is available to authorized users.
* Huang Zhang
zhanghuang2002113@163.com
Extended author information available on the last page of the article
Shearer (1993a) reviewed the freshwater ascomycota and
provided a definition and methods to study their taxonomy,
systematics, geographical distribution, ecology, and evolution. This definition and the study techniques have since
commonly been used (e.g., Zhang et al. 2017a; Dong et al.
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2018; Lu et al. 2018b; Luo et al. 2019; Yang et al. 2019b).
Unlike other fungi, which can be often traced to a specific
host, freshwater fungi have mostly been collected from “submerged wood” (Wei et al. 2018; Yu et al. 2018; Wang et al.
2019). Thus, there has been little research on the hosts of
freshwater fungi. The geographical distribution of freshwater fungi is also unclear and little can be concluded as distribution patterns are still largely based on the locations of
mycologists (Shearer 1993a); thus further work are needed
in these areas.
Dothideomycetes and Sordariomycetes, the largest classes
of Ascomycota, have been comprehensively reviewed (e.g.,
Hyde et al. 2013, 2020c; Maharachchikumbura et al. 2016;
Hongsanan et al. 2020a, b). Some websites, e.g. (http://
fungi.life.illinois.edu/) (Shearer and Raja 2010), (https://
www.marinefungi.org/) (Jones et al. 2019), (https://www.
dothideomycetes.org/) (Pem et al. 2019a) and (https://www.
freshwaterfungi.org/) (Calabon et al. 2020), are devoted to
taxonomy and classification of Dothideomycetes, provide
a database of freshwater ascomycetes and online resources
for marine fungi. There has been an increased interest in
the taxonomy and phylogeny of freshwater fungi (Lu et al.
2018b; Yang et al. 2018; Bao et al. 2019b, c; Boonmee et al.
2019), but further research will be carried out. Luo et al.
(2019) gave accounts of freshwater Sordariomycetes and
provided a modified backbone tree and detailed information on distribution, holotypes, sequence data and sexualasexual morph connections of taxa. They also introduced 47
new taxa based on fresh collections from China and Thailand, which indicated there is a high diversity of freshwater
Sordariomycetes.
In this article, a comprehensive study on freshwater Dothideomycetes is carried out based on new collections from
China, Egypt and Thailand, as well as studies of specimens
in fungal herbaria and literature reviews. Multigene phylogenetic analyses and morphological studies have resulted
in nine new genera, 33 new species, 16 new combinations
and 17 new geographical and habitat records being reported.
Considering that many freshwater fungi are scattered in
Dothideomycetes and the study of freshwater fungi have
greatly advanced, there is a need to revise all freshwater
Dothideomycetes with updated knowledge, and to provide
a better understanding of their morphology and phylogeny.
This monograph on freshwater Dothideomycetes provides
the latest generic concepts, descriptions, illustrations and
keys to genera and species. Using data from this study, we
present several common morphological traits that appear
to be adaptations of microfungi that live in flowing freshwater. Additionally, the global distribution of freshwater
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Fungal Diversity (2020) 105:319–575
Dothideomycetes is reviewed in this study to show the
poorly studied areas.
Materials and methods
Sample collection and specimen examination
Submerged wood was mostly collected from lentic or lotic
streams, lakes or rivers in China and Thailand as part of
a north-south survey (Hyde et al. 2016a), and from the
river Nile in Egypt between 2010 to 2019. The principle
of processing samples, including collection, incubation,
isolation and examination, following Shearer (1993a). The
detailed procedure of single spore isolation follows Chomnunti et al. (2014). Pseudoparaphyses are observed at 100×
using a Nikon ECLIPSE Ni compound microscope fitted
with a Canon EOS 600D digital camera. Two types of pseudoparaphyses, cellular and trabeculate, are defined if they
are clearly seen. Herbarium specimens are deposited in the
herbarium of Mae Fah Luang University (MFLU), Chiang
Rai, Thailand and the herbarium of Cryptogams Kunming
Institute of Botany Academia Sinica (KUN-HKAS), Kunming, China. Living cultures are deposited in the Mae Fah
Luang University Culture Collection (MFLUCC), Chiang
Rai, Thailand and Kunming Institute of Botany culture collection (KUMCC), Kunming, China. Facesoffungi and Index
Fungorum numbers are registered as in Jayasiri et al. (2015)
and Index Fungorum (2020).
DNA extraction, PCR amplification and sequencing
Cultures were grown on PDA at room temperature (25–27 °C)
and a Biospin Fungus Genomic DNA Extraction Kit (Bioer
Technology Co., Hangzhou, P.R. China) was used to extract
total genomic DNA from the fresh mycelium according to
the manufacturer’s instructions. Fragments of five loci were
amplified with polymerase chain reaction (PCR). Primer pairs
LR0R/LR5, NS1/NS4, ITS5/ITS4, EF1-983F/EF1-2218R and
fRPB2-5F/fRPB2-7cR are used for LSU, SSU, ITS, TEF and
RPB2, respectively. The amplifications were carried out using
the method described by Zhang et al. (2017a). The PCR products were viewed on 1% agarose electrophoresis gels stained
with ethidium bromide. Sequencing of five loci were carried
out by Shanghai Sangon Biological Engineering Technology
& Services Co. Shanghai, P.R. China.
Phylogenetic analysis
The sequences generated in this study were supplemented
with other Dothideomycetous sequences obtained from
GenBank, based on blast searches and relevant literatures.
Fungal Diversity (2020) 105:319–575
321
Fig. 1 World distribution of freshwater Dothideomycetes. Twelve
morphologically identified freshwater species which occur worldwide are shown with different symbols on the map. The number of
freshwater dothideomycetous species in each country is shown with
different colours, which indicates “richness” from zero to almost 150
species (in China)
The accession numbers used in the multigene analyses were
provided in supplementary material 1 and newly obtained
strains with their accession numbers were shown in supplementary material 2. Multiple sequence alignments were
generated with MAFFT v. 7 (https://mafft.cbrc.jp/alignment/
server/index.html) (Kuraku et al. 2013; Katoh et al. 2019).
The alignments were visually improved with BioEdit (Hall
1999). Phylogenetic analyses of the combined aligned dataset consisted of maximum likelihood (ML) and Bayesian
inference (BI). Maximum likelihood (ML) analysis was
performed at the CIPRES Science Gateway v.3.3 (https://
www.phylo.org/portal2/home.action) (Miller et al. 2010).
The final tree was selected among suboptimal trees from
each run by comparing likelihood scores under the GTRGAMMA substitution model. The best-fit models for the
Bayesian analyses were selected with MrModeltest v. 2.2
(Nylander 2004). The best-fit model GTR+I+G for LSU,
ITS and RPB2, and HKY+I+G for TEF. Six simultaneous
Markov chains were run for one million generations and
trees were sampled every 100th generation and 10000 trees
were obtained. The first 2500 trees representing the burn-in
phase of the analyses were discarded, while the remaining
7500 trees were used for calculating posterior probabilities
in the majority rule consensus tree. Phylogenetic tree was
visualized using FigTree v1.4.0 (http://tree.bio.ed.ac.uk/
software/figtree/, Rambaut 2012). Sequences derived in this
study are deposited in GenBank.
Results
Phylogenetic analysis of combined LSU, ITS, TEF and
RPB2 sequence data
The combined LSU, ITS, TEF and RPB2 sequence dataset
were employed for species of Dothideomycetes. The alignment comprised 714 strains with an alignment length of
5882 total characters including gaps. The RAxML analysis
resulted in a best scoring likelihood tree selected with a final
value for the combined dataset ln L = − 216610.299349. The
matrix has 4462 distinct alignment patterns, with 70.36% of
undetermined characters or gaps. Estimated base frequencies
are as follows: A = 0.238070, C = 0.259088, G = 0.278997,
T = 0.223845; substitution rates AC = 1.423061, AG =
3.547230, AT = 1.643052, CG = 1.083323, CT = 7.578123,
GT = 1.000000; gamma distribution shape parameter a =
0.439170 (Fig. 1).
Taxonomy
In this section, each freshwater genus is treated with
a generic description and notes. A list with freshwater
distribution and a key to freshwater species within each
genus are provided. A key to freshwater genera within
each family is also provided. An illustration of new species, new collections or line-drawings is provided after
each entry, where possible. The freshwater distribution
mainly follows an online resource for freshwater fungi
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(http://fungi.life.illinois.edu/) and published papers. The
species is marked with an asterisk if there are published
sequences deposited in GenBank. Freshwater Dothideomycetes are distributed in six orders, viz. Jahnulales,
Kirschsteiniotheliales, Minutisphaerales, Natipusillales,
Pleosporales and Tubeufiales; a few are scattered in Dothideomycetes families/genera incertae sedis. Classification and list of freshwater Dothideomycetes are shown
in Table 1.
*Denotes species having sequence data in GenBank
Jahnulales K.L. Pang, Abdel-Wahab, El-Shar., E.B.G. Jones
& Sivichai, Mycol. Res. 106(9): 1033 (2002)
Aliquandostipitaceae Inderb., Am. J. Bot. 88(1): 54 (2001)
Key to freshwater sexual genera of Aliquandostipitaceae
1. Ascomata hyaline or slightly pigmented………………2
1. Ascomata dark………………………………………..3
2. Ascospores with longitudinal sulcate striations, without
appendages or sheath……………………Megalohypha
2. Ascospores lacking longitudinal sulcate striations, often
with a large sheath……………………Aliquandostipite
3. Peridium composed of single row of cells…Neojahnula
3. Peridium not as above…………………………………4
4. Ascospores without sheath or appendage………Jahnula
4. Ascospores with sheath or appendage…………………5
5. Ascospores surrounded by a wavy mucilaginous
sheath…………………………………Pseudojahnula
5. Ascospores mostly with a refractive mucilaginous pad at
each end………………………………………Ascagilis
Aliquandostipite Inderbitzin, Am. J. Bot. 88(1): 54 (2001)
Saprobic on submerged wood or fallen, decorticated
branch. Sexual morph: Mycelium visible on the substratum,
of wide hyphae (> 10 µm and up to 50 µm wide), which may
bear ascomata. Ascomata scattered to clustered or gregarious, immersed to erumpent or superficial, uniloculate, globose to broadly ellipsoidal, dimorphic, with or without stalklike hypha attached to the base, hyaline or pale brown when
young, turn to dark brown with age, membranous, papillate.
Peridium thin, one-layered, composed of pale brown, thinwalled cells of textura angularis to globosa. Pseudoparaphyses numerous, cellular, sparsely branched, hyaline, septate.
Asci 8-spored, bitunicate, fissitunicate, clavate, with thickened apical region. Ascospores bi- to multi-seriate, variably
arranged, oval, pale brown, 1-septate, constricted at the
septum, usually asymmetric, mostly with a well-developed
hyaline sheath (Inderbitzin et al. 2001). Asexual morph:
Undetermined.
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Type species: Aliquandostipite khaoyaiensis Inderb.
Notes: Aliquandostipite was introduced to accommodate two species A. khaoyaiensis Inderb. and A. sunyatsenii Inderb., which form dimorphic ascomata (sessile and
stalked) and the widest hyphae known from ascomycetes
(Inderbitzin et al. 2001). However, with Jahnula sequences
added in the phylogenetic tree, A. sunyatsenii clustered in
Jahnula with high bootstrap support, which was therefore
transferred to Jahnula based on limited data (Pang et al.
2002). Among six species recorded in Index Fungorum
(2020), four have been confirmed in Aliquandostipite with
molecular data (Huang et al. 2018; Hyde et al. 2019). Aliquandostipite is similar to Jahnula, but can be distinguished
from the latter by its hyaline or less pigmented ascomata and
larger asci and ascospores.
According to our observations and literature, the
ascospores of A. crystallinus Raja et al., A. khaoyaiensis
and A. minuta Raja & Shearer form acicular crystals when
stored in lactic acid and glycerin (Raja et al. 2005; Raja
and Shearer 2007). These three taxa, however, can be distinguished based on the gelatinous sheath and appendages
on ascospores. Aliquandostipite siamensiae (Sivichai &
E.B.G. Jones) J. Campb. et al. is the only species producing
dimorphic ascospores. Aliquandostipite manochiae Sri-indr.
et al. is distinct in that it has branched and anastomosing
pseudoparaphyses, lacking arcicular crystals within the
spores, and ascospore size (Liu et al. 2015). Aliquandostipite manochiae might be synonymous with A. khaoyaiensis
(Inderbitzin et al. 2001) based on their similar morphological features, overlapping dimension of ascomata, asci, and
ascospores, and their freshwater habitats. Molecular data
are necessary to confirm A. manochii as a unique species
(Fig. 2).
List of freshwater Aliquandostipite species
*Aliquandostipite crystallinus Raja, A. Ferrer & Shearer,
Mycotaxon 91: 208 (2005); Fig. 3e, g–i
Freshwater distribution: Costa Rica (Raja et al. 2005),
Panama (Raja et al. 2005), Peru (Shearer et al. 2015), USA
(Raja et al. 2005, 2009b), Venezuela (Raja et al. 2005)
*Aliquandostipite khaoyaiensis Inderbitzin, Am. J. Bot. 88:
54 (2001)
Facesoffungi number: FoF09158; Figs. 3a–d, f, 4
Freshwater distribution: Costa Rica (Raja et al. 2005),
USA (Raja et al. 2009b), Thailand (this study)
Saprobic on submerged wood. Sexual morph: Ascomata
220–240 μm high, 190–210 μm diam., hyaline to pale brown,
become dark brown with age, scattered, superficial, sometimes seated in a pseudostroma, globose or subglobose, sessile, membranous, with ostiolate papilla. Peridium 25–40 μm
thick, membranous, composed of thin-walled, pale brown,
compressed cells of textura angularis. Pseudoparaphyses 2
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Table 1 Classification and list of freshwater Dothideomycetes
Order
Family
Genus (number of freshwa- Species treated in this
ter species)
study
Jahnulales
Aliquandostipitaceae
Aliquandostipite (5)
Ascagilis^ (7)
Brachiosphaera^ (2)
Jahnula^ (11)
Megalohypha^ (1)
Neojahnula^ (1)
Al. khaoyaiensis
As. guttulaspora
As. queenslandica
As. seychellensis
As. submersa
As. sunyatsenii
As. thailandensis
New geographical record
New combination
New combination
New combination
New species
New combination
New species
J. rostrata
New geographical record
N. australiensis
Pseudojahnula^ (1)
Kirschsteiniotheliales
Minutisphaerales
Natipusillales
Pleosporales
Kirschsteiniotheliaceae
Acrogenosporaceae
Minutisphaeraceae
Natipusillaceae
Acrocalymmaceae
Aigialaceae
Amniculicolaceae
Anteagloniaceae
Aquasubmersaceae
Astrosphaeriellaceae
Bambusicolaceae
Corynesporascaceae
Delitschiaceae
Dictyosporiaceae
P. potamophila
Xylomyces (5)
Kirschsteiniothelia (6)
Acrogenospora (13)
Minutisphaera^ (5)
Natipusilla^ (4)
Acrocalymma (2)
A. bipolare
Fissuroma (1)
F. neoaggregatum
Neoastrosphaeriella (1)
Amniculicola (6)
A. guttulata
Murispora (4)
Neomassariosphaeria^ (1)
Vargamyces^ (1)
Flammeascoma (1)
Purpureofaciens^ (1)
P. aquatica
Aquasubmersa (2)
Aquatospora^ (1)
A. cylindrica
Astrosphaeriella (7)
Caryospora (4)
C. aquatica
C. quercus
C. submersa
Pithomyces (1)
Xenoastrosphaeriella (2)
Bambusicola (1)
B. aquatica
Corynespora (3)
Delitschia (3)
Aquadictyospora^ (1)
Aquaticheirospora^ (1)
Cheirosporium^ (2)
Dendryphiella (1)
De. vinosa
Dictyocheirospora (9)
Notes
Di. rotunda
Di. thailandica
New genus
New combination
new genus
new combination
New species
New habitat record
New collection
New genus
New species
New genus
New species
New geographical record
New habitat record
New species
New species
New habitat record
New synonym
New collection
New species
Dictyosporium (11)
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Table 1 (continued)
Order
Family
Didymosphaeriaceae
Latoruaceae
Lentitheciaceae
Lindgomycetaceae
Longipedicellataceae
Genus (number of freshwa- Species treated in this
ter species)
study
Digitodesmium (3)
Jalapriya (1)
Pseudocoleophoma (1)
Pseudodictyosporium (1)
Vikalpa (1)
Didymosphaeria (1)
Paraphaeosphaeria (2)
Paraconiothyrium (1)
Pseudotrichia (1)
Pseudoasteromassaria (2)
Keissleriella (1)
Lentithecium (6)
Poaceascoma (1)
Setoseptoria (2)
Tingoldiago^ (3)
Arundellina (1)
Clohesyomyces^ (1)
Hongkongmyces (2)
Aquimassariosphaeria (2)
Lindgomyces^ (14)
Lolia^ (2)
Neolindgomyces (1)
Longipedicellata (2)
Pseudoxylomyces^ (2)
P. aquatica
New species
L. kunmingense
New species
C. aquaticus
H. aquaticus
New collection
New species
New genus
New species
New combination
New species
A. kunmingensis
A. typhicola
L. aquaticus
L. aquatica
P. aquaticus
P. elegans
Submersispora^ (1)
S. variabilis
Lophiostomataceae
Lophiotremataceae
Massarinaceae
Melanommataceae
Morosphaeriaceae
13
Biappendiculispora (1)
Crassiclypeus^ (1)
Pseudocapulatispora (1)
Flabellascoma (1)
Lentistoma (2)
Lophiostoma (6)
Neotrematosphaeria^ (1)
Neovaginatispora (1)
Sigarispora (1)
Vaginatispora (2)
Lophiotrema (1)
Byssothecium (2)
Helminthosporium (2)
Byssosphaeria (1)
Camposporium (6)
Herpotrichia (1)
Mamillisphaeria^ (1)
Melanomma (1)
Aquihelicascus^ (3)
Notes
P. longiappendiculata
F. fusiforme
L. aquaticum
New species
New species
New collection
New genus
New species
New habitat and geographical record
New collection
New species
New genus
Fungal Diversity (2020) 105:319–575
325
Table 1 (continued)
Order
Family
Genus (number of freshwa- Species treated in this
ter species)
study
A. songkhlaensis
A. thalassioideus
A. yunnanensis
Neohelicascus^ (8)
Nigrogranaceae
Occultibambusaceae
Nigrograna (2)
Occultibambusa (3)
Ne. aquaticus
Ne. chiangraiensis
Ne. egyptiacus
Ne. elaterascus
Ne. gallicus
Ne. submersus
Ne. unilocularis
Ne. uniseptatus
Ni. aquatica
O. kunmingensis
O. pustula
Seriascoma (1)
S. didymosporum
Parabambusicolaceae
Aquastroma (1)
Parabambusicola (1)
Paramonodictys (1)
A. magniostiolata
Parab. aquatica
Param. solitarius
Periconiaceae
Phaeoseptaceae
Periconia (6)
Phaeoseptum (1)
Pleopunctum (1)
Pl. pseudoellipsoideum
Splanchnonema (1)
Pleomonodictys (1)
Pseudoastrosphaeriella (5) Ps. aquatica
Pleomassariaceae
Pleomonodictydaceae
Pseudoastrosphaeriellaceae
Roussoellaceae
Roussoella (3)
Neoroussoella (2)
Ps. bambusae
R. aquatica
N. bambusae
N. leucaenae
Testudinaceae
Tetraplosphaeriaceae
Angustospora^ (1)
Ernakulamia (2)
E. cochinensis
Shrungabeeja (2)
E. xishuangbannaensis
S. vadirajensis
Tetraploa (6)
Torulaceae
Trematosphaeriaceae
S. aquatica
T. puzheheiensis
T. yunnanensis
Notes
New species
New combination
New species
New genus
New combination
New combination
New combination
New combination
New combination
New species
New combination
New combination
New species
New species
New habitat and geographical record
New habitat and geographical record
New geographical record
New species
New habitat and geographical record
New habitat record
New species
New collection
New species
New habitat and geographical record
New habitat and geographical record
New habitat and geographical record
New combination
New habitat and geographical record
New species
New species
New species
Triplosphaeria (1)
Dendryphion (4)
Neotorula^ (2)
Rostriconidium^ (1)
Torula (4)
Falciformispora (3)
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Table 1 (continued)
Order
Family
Wicklowiaceae
Pleosporales genera
incertae sedis
Genus (number of freshwa- Species treated in this
ter species)
study
Hadrospora (1)
Trematosphaeria (4)
Wicklowia^ (2)
Ascorhombispora^ (1)
Fusiformiseptata^ (1)
Tubeufiales
Tubeufiaceae
Isthmosporella^ (1)
Rebentischia (1)
Aquaphila (1)
Berkleasmium (5)
Boerlagiomyces (3)
Chlamydotubeufia (4)
Dematiohelicomyces^ (1)
Dictyospora^ (1)
Helicoarctatus^ (1)
Helicodochium^ (2)
Helicohyalinum^ (2)
Helicoma (8)
Helicomyces (5)
Helicosporium (6)
Helicotruncatum (1)
Helicotubeufia (2)
Muripulchra^ (1)
Neochlamydotubeufia (2)
Neohelicomyces (6)
Neohelicosporium (11)
Neotubeufia^ (1)
Pseudohelicomyces (3)
Pseudohelicoon^ (2)
Thaxteriellopsis (1)
Tubeufia (29)
Wiesneriomycetaceae
Dothideomycetes families Argynnaceae
incertae sedis
Pseudorobillardaceae
Dothideomycetes
genera incertae sedis
Wiesneriomyces (1)
Lepidopterella^ (2)
Pseudorobillarda (1)
Alascospora^ (1)
Ascominuta^ (2)
Lucidascocarpa^ (1)
Monodictys (10)
Ocala^ (1)
Quintaria (2)
Wettsteinina (2)
^Genera known only from freshwater habitats (before this study)
13
Notes
F. crocea
New genus
New species
A. albicans
B. latisporum
New collection
New collection
Helicos. thailandense
Helicot. palmigenum
New species
New habitat and geographical record
Neohelicom. dehongensis
Neohelicom. thailandicus
Neohelicos. submersum
New species
New species
New species
T. aquatica
T. cylindrothecia
T. tectonae
New collection
New collection
New synonym
Fungal Diversity (2020) 105:319–575
Fig. 2 Phylogram generated
from maximum likelihood
analysis of combined LSU,
ITS, TEF and RPB2 sequence
data for species of Dothideomycetes. Bootstrap values for
maximum likelihood equal to or
greater than 75% and Bayesian
posterior probabilities equal to
or greater than 0.95 are placed
near the branches as ML/BYPP.
Newly generated sequences are
in red and ex-type strains are
in bold. The new species introduced in this study are indicated
with underline. Freshwater
strains are indicated with a red
letter “F”. Orders not treated in
this study are compressed. The
tree is rooted to Capronia pilosella AFTOL-ID 657 and Endocarpon pallidulum AFTOL-ID
661 (Eurotiomycetes)
327
100/1.00
Periconia pseudodigitata KT 644
100/1.00
Periconia pseudodigitata KT 1195A F
100/1.00
Periconia pseudodigitata KT 1395 F
Periconia digitata CBS 510.77
Noosia banksiae CPC 17282
84/1.00
Periconia igniaria CBS 845.96
100/1.00
Periconia igniaria CBS 379.86
87/0.99
Periconia minutissima MFLUCC 15-0245 F
100/1.00
Periconiaceae
Flavomyces fulophazii CBS 135761
Periconia byssoides H 4600
100/1.00
Periconia byssoides H 4432
83/1.00
Periconia byssoides H 4853
Periconia aquatica MFLUCC 16-0912 F
92/-98/1.00
Periconia submersa MFLUCC 16-1098 F
97/1.00
Stagonospora duoseptata CBS 135093
--/0.96
Stagonospora paludosa CBS 135088
86/--
Helminthosporium velutinum H 4626
Massarinaceae
Byssothecium circinans CBS 675.92
Massarina eburnea CBS 473.64
Fuscostagonospora cytisi MFLUCC 16-0622
99/0.99
Fuscostagonospora sasae KT 1467
Fuscostagonosporaceae
Bimuria novae-zelandiae
100/1.00
Deniquelata barringtoniae MFLUCC 11-0422
Spegazzinia deightonii yone 66
Didymosphaeriaceae
Montagnula opulenta AFTOL-ID 1734
85/--
Didymosphaeria rubi ulmifolii MFLUCC 14-0023
--/0.98
Lentithecium unicellulare isolate MD 6004 F
Lentithecium carbonneanum CBS 144076 F
Lentithecium voraginesporum CBS H-22560
--/0.96
Lentithecium kunmingense KUMCC 19-0101 F
Lentithecium cangshanense DLUCC 0143 F
99/0.95
100/--
84/--
Lentitheciaceae
Poaceascoma aquaticum MFLUCC 18-0981 F
Poaceascoma aquaticum MFLUCC 14-0048 F
Poaceascoma helicoides MFLUCC 11-0136
Poaceascoma halophilum MFLUCC 15-0949
Dictyocheirospora thailandica MFLUCC 18-0987 F
Dictyocheirospora rotunda KUMCC 19-0103 F
Dictyocheirospora rotunda MFLUCC 17-1313 F
100/1.00
Dictyocheirospora rotunda KUMCC 19-0105 F
Dictyosporiaceae
Dictyocheirospora rotunda MFLUCC 17-0222 F
96/--
Dendryphiella vinosa
100/0.99
Dendryphiella vinosa MFLUCC 17-2321 F
Dendryphiella eucalyptorum CBS 137987
99/-99/--
Neobambusicola strelitziae CBS 138869
Sulcatispora acerina KT 2982
Sulcatisporaceae
Magnicamarosporium iriomotense KT 2822
Ascorhombispora aquatica HKUM 10859 F
μm diam., numerous, cellular, sparsely branched, hyaline,
septate. Asci 115–175 × 35–65 μm ( x̄ = 145 × 45 μm, n =
20), 8-spored, bitunicate, fissitunicate, clavate, thickened at
the apex, sessile or short pedicellate, with a well-developed
ocular chamber. Ascospores 50–70 × 14.5–19.5 μm (x̄ = 63
× 17 μm, n = 20), variably arranged in asci, oval, hyaline
to pale brown, 1-septate, constricted at the septum, asymmetric, upper cell slightly shorter and wider than lower cell,
13
328
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
Bambusicola didymospora MFLUCC 15-0189
100/1.00
Bambusicola didymospora MFLUCC 10-0557
Bambusicola splendida MFLUCC 11-0439
100/1.00
Bambusicola triseptatispora MFLUCC 11-0166
76/1.00
Bambusicola loculata MFLU 15-0056
Bambusicola massarinia MFLUCC 11-0389
87
100/--
Bambusicola dimorpha MFLUCC 13-0282
100/0.99
98/0.99
Bambusicola bambusae MFLUCC 11-0614
100/1.00
Bambusicolaceae
Bambusicola pustulata MFLUCC 15-0190
Bambusicola aquatica MFLUCC 18-1031 F
100/1.00
Bambusicola irregulispora MFLUCC 11-0437
99/0.99
Bambusicola sichuanensis SICAU 16-0004
89/--
Bambusicola thailandica MFLUCC 11-0147
99/0.99
Bambusicola subthailandica SICAU 16-0005
Palmiascoma gregariascomum MFLUCC 11-0175
Leucaenicola aseptata MFLUCC 17-2423
100/1.00
Leucaenicola phraeana MFLUCC 18-0472
Polyschema larviformis CBS 463.88
77/--
Polyschema sclerotigenum UTHSC:DI14-305
100/1.00
Polyschema congolensis CBS 542.73
98/1.00
Polyschema terricola CBS 301.65
96/1.00
Latorua caligans CBS 576.65
--/1.00
Latorua grootfonteinensis CBS 369.72
Matsushimamyces bohaniensis CBEC001
Matsushimamyces venustum CBS 140212
85/--
Latoruaceae
Pseudoasteromassaria spadicea MFLUCC 15-0973 F
100/1.00
Pseudoasteromassaria fagi KT3432
Pseudoasteromassaria fagi KT2966
98/1.00
Pseudoasteromassaria aquatica MFLUCC 18-1397 F
84/--
Longipedicellata aptrootii MFLUCC 16-0244
99/--
Longipedicellata aptrootii MFLUCC 18-0988 F
89/-100/0.98
100/0.99
Longipedicellata aptrootii MFLUCC 10-0297 F
Longipedicellata aptrootii MFLUCC 16-0384
100/1.00
Longipedicellata aquatica MFLUCC 19-0340 F
100/0.99
Longipedicellata aquatica MFLUCC 17-2334 F
77/--
Longipedicellataceae
Longipedicellata aquatica MFLUCC 19-0324 F
100/1.00
Submersispora variabilis MFLUCC 17-2360 F
99/--
Pseudoxylomyces elegans MFLUCC 17-2350 F
Pseudoxylomyces elegans KT 2887 F
Pseudoxylomyces aquaticus KUMCC 17-0312 F
99/0.95
guttulate, straight or curved, thin-walled, smooth, sheathed.
Sheath first appressed to the ascospore wall, expanding and
detaching from the polar regions when mounted in water,
becoming balloon-like at the two poles, finally surrounding
the entire ascospore. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 30 mm in 45 days at 25 °C, grey from above, dark brown
13
Macrodiplodiopsidaceae
from below, surface rough, with dense mycelium, mostly
immersed in culture, dry, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat827-1 (MFLU 18-1573), living culture MFLUCC
18-1249; ibid., hat827-2 (HKAS 105024), living culture
KUMCC 19-0039; Chestnut Hill, on submerged wood in a
stream, 10 May 2018, W. Dong, hat460-1 (MFLU 18-1554);
Fungal Diversity (2020) 105:319–575
329
Fig. 2 (continued)
Paramonodictys solitarius GZCC 20-0007
Paramonodictys solitarius MFLUCC 17-2353 F
89/-100/1.00
Monodictys sp. KH 331
Monodictys sp. JO 10
100/1.00
Aquastroma magniostiolata MFLUCC 18-0976 F
Aquastroma magniostiolata KT 2485 F
100/1.00
Paratrimmatostroma kunmingensis HKAS 102224A
Paratrimmatostroma kunmingensis HKAS 102224B
Pseudomonodictys tectonae MFLUCC 12-0552
100/--
Lonicericola hyaloseptispora KUMCC 18-0149
100/1.00
Lonicericola hyaloseptispora KUMCC 18-0150
Multilocularia bambusae MFLUCC 11-0180
Parabambusicola bambusina KH 139
86/-100/1.00
Parabambusicolaceae
Parabambusicola bambusina KT 2637
98/--
Parabambusicola bambusina H 4321
Parabambusicola aquatica MFLUCC 18-1140 F
100/0.97
Parabambusicola thysanolaenae KUMCC 18-0148
100/1.00
Parabambusicola thysanolaenae KUMCC 18-0147
Neoaquastroma bauhiniae MFLUCC 16-0398
100/1.00
100/1.00
Neoaquastroma bauhiniae MFLUCC 17-2205
Neoaquastroma guttulatum MFLUCC 14-0917
100/1.00
Neoaquastroma krabiense MFLUCC 16-0419
100/1.00
Multiseptospora thailandica MFLUCC 12-0006
100/1.00
Multiseptospora thailandica MFLUCC 11-0204
Multiseptospora thailandica MFLUCC 11-0183
79/--
Aquihelicascus songkhlaensis MFLUCC 18-1154 F
99/1.00
Aquihelicascus songkhlaensis MFLUCC 18-1278 F
Aquihelicascus songkhlaensis MFLUCC 18-1273 F
100/1.00
Aquihelicascus yunnanensis MFLUCC 18-1025 F
100/1.00
Aquihelicascus thalassioideus MFLUCC 10-0911 F
100/1.00
Aquihelicascus thalassioideus KUMCC 19-0094 F
Neohelicascus aquaticus MFLUCC 10-0918 F
100/--
Neohelicascus aquaticus KUMCC 19-0107 F
93/0.99
Morosphaeriaceae
Neohelicascus chiangraiensis MFLUCC 13-0883 F
100/--
100/--
Neohelicascus elaterascus MFLUCC 18-0985 F
100/1.00
Neohelicascus elaterascus MFLUCC 18-0993 F
100/1.00
100/--
Helicascus nypae BCC 36752
Helicascus nypae BCC 36751
Helicascus kanaloanus ATCC 18591
Multiseptospora thysanolaenae MFLUCC 11-0202
87/0.99
98/0.96
100/1.00
100/--
Trematosphaeria pertusa KT 3314
Trematosphaeria pertusa KT 3315
Trematosphaeria pertusa KT 1496 F
Trematosphaeria pertusa CBS 122368
Trematosphaeria grisea CBS 332.50
97/-- 100/1.00
Trematosphaeriaceae
Trematosphaeria grisea CBS 135984
Falciformispora uttaraditense MFLUCC 19-0485 F
96/--
Falciformispora lignatilis BCC 21117
Falciformispora aquatica MFLUCC 18-0212 F
13
330
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
100/1.00
86/--
Pleomonodictys capensis CBS 968.97
Pleomonodictys descalsii CBS 142298
Pleomonodictydaceae
Asteromassaria pulchra CBS 124082
100/0.97
Dacampiaceae
85/1.00
Coniothyriaceae
100/1.00
Coniothyriaceae
83/0.95
Camarosporidiellaceae
100/1.00
Neophaeosphaeriaceae
100/1.00
87/-97/1.00
Libertasomycetaceae
Cucurbitariaceae
Neocamarosporiaceae
100/1.00
Leptosphaeria dryadis CBS 643.86
Pleosporaceae
Plenodomus biglobosus CBS 303.51
95/--
99/--
Phaeosphaeriaceae
100/1.00
77/--
Didymellaceae
Dothidotthia aspera CPC 12933
88/--
Dothidotthia symphoricarpi CPC 12929
Dothidotthiaceae
Acrocalymma vagum CPC 24226
77/-100/--
Acrocalymma walkeri UTHSCDI 16-195
Acrocalymma vagum CPC 24225
Acrocalymma ficus CBS 317.76
Acrocalymma aquatica MFLUCC 11-0208 F
98/1.00
Acrocalymmaceae
Acrocalymma medicaginis CPC 24340
Acrocalymma pterocarpi MFLUCC 17-0926
76/--
100/-85/--
Acrocalymma bipolare MD1321 F
Acrocalymma cycadis CBS 137972
100/0.99
Halojulellaceae
Ascocylindricaceae
100/1.00
Corynespora cassiicola CBS 100822
100/1.00
Corynespora smithii CABI 5649b
Corynesporascaceae
Cyclothyriella rubronotata TR9
Cyclothyriella rubronotata TR1
100/1.00
Cyclothyriella rubronotata CBS 419.85
Cyclothyriellaceae
Cyclothyriella rubronotata TR3
Massariosphaeria phaeospora CBS 611.86
Prosthemium orientale KT1669
100/1.00
Prosthemium betulinum VM20040721
96/0.99
100/0.99
Pseudotrichia mutabilis SMH 1541
77/-87/0.99
Aposphaeria populina CBS 350.82
Pleomassariaceae
Pseudotrichia mutabilis SMH 5288
Aposphaeria populina CBS 543.70
100/1.00
100/--
Melanomma pulvis pyrius CBS 124080
Melanomma pulvis pyrius CBS 109.77
Byssosphaeria schiedermayeriana SMH3157
100/--
Melanommataceae
Byssosphaeria schiedermayeriana GKM 152 N
Byssosphaeria schiedermayeriana MFLUCC 10-0100
Fusiformiseptata crocea MFLUCC 18-1415 F
13
Fusiformiseptata
Fungal Diversity (2020) 105:319–575
331
Fig. 2 (continued)
Nigrograna obliqua MRP
97/0.98
Nigrograna obliqua BW4
97/1.00
Nigrograna obliqua CBS 141475
Nigrograna obliqua CBS 141477
97/1.00
Nigrograna mycophila CBS 141478
99/1.00
Nigrograna mycophila CBS 141483
95/1.00
Nigrograna mycophila MF6
Nigrograna obliqua MF
Nigrograna cangshanensis MFLUCC 15-0253 F
94/1.00
Nigrograna thymi MFLU 17-0497
Nigrograna locuta-pollinis LC11691
100/1.00
Nigrograna locuta-pollinis LC11685
99/1.00
Nigrograna locuta-pollinis LC11690
Nigrograna aquatica MFLUCC 17-2318 F
Nigrograna rhizophorae MFLUCC 18-0397
100/--
Nigrogranaceae
Nigrograna samueliana NFCCI 4383
Nigrograna norvegica CBS 141485
Nigrograna fuscidula MF1a
81/1.00
91/1.00
Nigrograna fuscidula CBS 141476
Nigrograna fuscidula MF3
Nigrograna fuscidula CBS 141556
100/-94/1.00
Nigrograna fuscidula MF8
Nigrograna fuscidula MF9
Biatriospora mackinnonii E9303e
81/--
Biatriospora mackinnonii L3396
Biatriospora mackinnonii CBS 674.75
81/--
Biatriospora mackinnonii CBS 110022
Biatriospora marina CY 1228
Biatriospora peruviensis CCF 4485
Biatriospora antibiotica CCF 4378
Biatriospora carollii CCF 4484
Nigrograna yasuniana E8604b
84/--
Paradictyoarthrinium diffractum MFLUCC 13-0466
100/1.00
Paradictyoarthrinium tectonicola MFLUCC13-0465
87/0.99
100/1.00
78/0.99
100/1.00
Paradictyoarthriniaceae
Occultibambusa aquatica MFLUCC 11-0006 F
Occultibambusa jonesii GZCC 16-0117
Occultibambusa pustula MFLUCC 11-0502
Occultibambusa pustula MFLUCC 18-1028 F
Occultibambusa kunmingensis HKAS 102151 F
100/1.00
100/--
Occultibambusaceae
Seriascoma didymospora MFLUCC 11-0194
Seriascoma didymospora MFLUCC 11-0179
100/1.00
Seriascoma didymospora MFLUCC 18-1026 F
ibid., hat460-2 (HKAS 105066), living culture KUMCC
19-0083.
Notes: Our two new isolates MFLUCC 18-1249 and
KUMCC 19-0083 are identified as Aliquandostipite
khaoyaiensis based on their identical LSU, SSU, ITS
sequence data and morphology, although they lack a stalked
ascomata. The new isolates cluster with A. khaoyaiensis and
A. siamensis with high bootstrap support (Fig. 14), which
appears to be conspecific. However, our isolates must be A.
khaoyaiensis because of its typical sheath. Aliquandostipite
khaoyaiensis has monomorphic, pale brown ascospores with
well-developed sheath (Inderbitzin et al. 2001). While A.
siamensis has dimorphic, hyaline or brown ascospores without sheath (Pang et al. 2002). Aliquandostipite siamensis
13
332
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
Dendryphion fluminicola KUMCC 15-0321 F
98/-89/--
Dendryphion fluminicola DLUCC 0849 F
Dendryphion fluminicola MFLUCC 17-1689 F
Dendryphion fluminicola MFLUCC 17-0105 F
100/1.00
Dendryphion europaeum CPC 23231
96/1.00
Dendryphion europaeum CPC 22943
Dendryphion aquaticum MFLUCC 15-0257 F
100/1.00
Dendryphion submersum MFLUCC 15-0271
80/1.00
Dendryphion submersum DLUCC 0698 F
Dendryphion submersum KUMCC 15-0455 F
Dendryphion nanum MFLUCC 16-0975 F
99
Dendryphion nanum HKAS 84010 F
Dendryphion nanum HKAS 84012 F
Dendryphion nanum MFLUCC 17-1319 F
100/1.00
Dendryphion nanum MFLUCC 16-0987 F
100/1.00
Dendryphion nanum DLUCC 0745 F
Dendryphion nanum DLUCC 0807 F
79/0.99
Neotorula aquatica MFLUCC 15-0342 F
100/1.00
Neotorula submersa KUMCC 15-0280 F
Rostriconidium aquaticum MFLUCC 16-1113 F
100/1.00
Rostriconidium aquaticum KUMCC 15-0297 F
99/1.00
Torula ficus MFLUCC 16-1348 F
91/1.00
Torula ficus MFLUCC 16-1280 F
Torula ficus MFLUCC 16-1259 F
95/1.00
Torula ficus KUMCC 15-0428 F
100/1.00
Torula ficus KUMCC 16-0038
100/0.96
100/--
Torulaceae
90/--
Torula ficus KUMCC 16-0035
Torula ficus CBS 595.96
100/1.00
Torula aquatica DLUCC 0550 F
100/1.00
Torula aquatica MFLUCC 16-1115 F
100/1.00
Torula aquatica DLUCC 0699 F
75/--
Torula herbarum CPC 24114
Torula hollandica CBS 220.69
100/1.00
Torula chiangmaiensis KUMCC 16-0039
Torula pluriseptata MFLUCC 14-0437
100/-100/1.00
99/0.96
100/1.00
96/1.00
Torula chromolaenae KUMCC 16-0036
Torula mackenziei MFLUCC 13-0839
Torula masonii KUMCC 16-0033
Torula masonii DLUCC 0588 F
Torula masonii CBS 245.57
Torula acaciae CPC 29737
lacks ITS sequence data and needs fresh material to confirm
its phylogenetic difference with A. khaoyaiensis. This is a
new geographical record for A. khaoyaiensis from parts of
Thailand.
Aliquandostipite minuta Raja and Shearer, Mycoscience 43:
395 (2007); Fig. 5a–e
Freshwater distribution: USA (Raja and Shearer 2007;
Raja et al. 2009b)
Aliquandostipite manochii Sri-indr., Boonyuen, Suetrong,
K.L. Pang & E.B.G. Jones, Fungal Diversity 72: 103 (2015)
Freshwater distribution: Thailand (Liu et al. 2015)
Aliquandostipite separans (Abdel-Wahab & El-Sharouney)
J. Campb., Raja A. Ferrer, Sivichai & Shearer, Can. J. Bot.
85: 881 (2007)
13
Fungal Diversity (2020) 105:319–575
333
Fig. 2 (continued)
Roussoella nitidula MFLUCC 11-0182
100/1.00
90/0.99
Roussoella nitidula MFLUCC 11-0634
Roussoella aquatica MFLUCC 18-1040 F
Neoroussoella leucaenae KUMCC 18-0050 F
Roussoellaceae
Neoroussoella bambusae MFLUCC 11-0124
100/1.00
98/1.00
Neoroussoella bambusae MFLUCC 18-1048 F
Parathyridaria robiniae MFLUCC 14-1119
100/1.00
100/0.98
Parathyridaria ramulicola CBS 141479
Parathyridaria percutanea CBS 868.95
Thyridariaceae
Thyridaria acaciae CBS 138873
100/1.00
Thyridaria broussonetiae CBS 141481
Sporidesmioides thailandica MFLUCC 13-0840
100/1.00
Sporidesmioides thailandica KUMCC 16-0012
Pseudocoleodictyospora sukhothaiensis MFLUCC 12-0554
100/1.00
Pseudocoleodictyospora tectonae MFLUCC 12-0385
Pseudocoleodictyosporac
Flabellascoma cycadicola KT 2034
100/1.00
100/1.00
Flabellascoma minimum KT 2040
Flabellascoma fusiforme MFLUCC 18-1019 F
100/0.99
100/1.00
Lentistoma bipolare CBS 115370
Lentistoma aquaticum MFLUCC 18-1275 F
100/0.99
100/1.00
Pseudolophiostoma tropicum KT 3134
100/1.00
100/1.00
Lophiostomataceae
Pseudolophiostoma cornisporum KH 322
Lophiostoma vitigenum HH26931
Lophiostoma vitigenum HH26930
100/1.00
Pseudocapulatispora longiappendiculata MFLUCC 18-1027 F
100/--
Teichosporaceae
81/--
Sporormiaceae
100/0.99
Angustimassarina quercicola MFLUCC 15-0079
Angustimassarina rosarum MFLUCC 15-0080
Lentimurispora urniformis MFLUCC 18-0497
100/1.00
Lentimurisporaceae
Lignosphaeria thailandica MFLUCC 11-0376
Lignosphaeria fusispora MFLUCC 11-0377
100/1.00
76/--
Amorosiaceae
Lophiostoma macrostomoides GKM 1033
100/--
100/--
100/1.00
100/1.00 100/1.00
Thyridaria macrostomoides GKM 1159
Pleopunctum pseudoellipsoideum MFLUCC 19-0391
Pleopunctum pseudoellipsoideum HKAS 102147 F
Pleopunctum ellipsoideum MFLUCC 19-0390
Phaeoseptaceae
Phaeoseptum terricola MFLUCC 10-0102
100/1.00
100/--
Phaeoseptum mali MFLUCC 17-2108
Phaeoseptum aquaticum CBS 123113
100/1.00
Decaisnella formosa BCC 25616
Decaisnella formosa BCC 25617
100/--
Basionym: Patescospora separans Abdel-Wahab & ElSharouney, Mycol. Res. 106: 1033 (2002)
Freshwater distribution: Egypt (Pang et al. 2002)
Halotthiaceae
*Aliquandostipite siamensiae (Sivichai & E.B.G. Jones) J.
Campb., Raja A. Ferrer, Sivichai & Shearer, Can. J. Bot.
85: 879 (2007)
Basionym: Jahnula siamensiae Sivichai & E.B.G. Jones,
Mycol. Res. 106: 1037 (2002)
13
334
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
92/1.00
Hermatomycetaceae
Aquasubmersa japonica KT 2863
76/1.00
99/1.00
Aquasubmersa japonica KT 2862
Aquasubmersa japonica KT 2813 F
99/1.00
Aquasubmersaceae
Aquasubmersa mircensis MFLUCC 11-0401 F
100/--
Cryptocoryneaceae
Atrocalyx lignicola CBS 122364
Lophiotrema nucula CBS 627.86
Lophiotremataceae
82/0.97
Lophiotrema neoarundinaria KT 1034
Anteaglonium rubescens WU 36963
100/1.00
100/0.99
Anteaglonium rubescens WU 36960
Anteaglonium rubescens WU 36962
Anteaglonium thailandium MFLUCC 14-0816
Anteaglonium parvulum MFLUCC 14-0817
Anteaglonium parvulum MFLUCC 14-0821
78/--
Anteaglonium parvulum MFLUCC 14-0823
Anteaglonium globosum ANM 925.2
100/0.96
Anteaglonium globosum SMH5283
Anteaglonium brasiliense HUEFS 192250
Anteagloniaceae
Anteaglonium abbreviatum ANM 925a
93/1.00
Anteaglonium abbreviatum GKM1029
Anteaglonium gordoniae MFLUCC 17-2431
Flammeascoma bambusae MFLUCC 10-0551 F
87/--
Flammeascoma lignicola MFLUCC 10-0128b
Purpureofaciens aquatica MFLUCC 18-1241 F
Anteaglonium latirostrum GKML100Nb
Ernakulamia cochinensis MFLUCC 18-1237 F
78/--
Ernakulamia cochinensis PRC 3992
Ernakulamia xishuangbannaensis KUMCC 17-0187
91/1.00
Ernakulamia krabiensis MFLUCC 18-0237
Dothideomycetes sp. ARIZ FL2038
79
Shrungabeeja longiappendiculata 76463
100/1.00
100/1.00
Shrungabeeja longiappendiculata 76464
Tetraplosphaeriaceae
Shrungabeeja vadirajensis MFLUCC 17-2362 F
75/--
Tetraploa puzheheiensis KUMCC 20–0151 F
100/1.00
100/0.97
Tetraploa sp. CBS 996.70
Tetraploa yakushimensis CBS 125435
99/-90/--
Tetraploa yunnanensis MFLUCC 19-0319
Tetraploa sasicola KT 563
99/--
Tetraploa sp. 1 JCM 14424
100/--
Freshwater distribution: Thailand (Pang et al. 2002),
USA (Raja et al. 2009b)
Key to freshwater Aliquandostipite species
1. Ascospores with sheath………………………………2
1. Ascospores with narrow sheath or without sheath………
……………………………………………A. separans
2. Ascospores dimorphic……………………A. siamensiae
13
Hypsostromataceae
2.
3.
3.
4.
Ascospores monomorphic……………………………3
Ascospores without appendages………A. khaoyaiensis
Ascospores with appendages…………………………4
Ascospores with small apical appendages……………
…………………………………………A. crystallinus
4. Ascospores with numerous filamentous appendages……
………………………………………………A. minuta
Fungal Diversity (2020) 105:319–575
335
Fig. 2 (continued)
Pseudoastrosphaeriella bambusae MFLUCC 10-0885 F
Pseudoastrosphaeriella bambusae KUMCC 19-0095 F
Pseudoastrosphaeriella bambusae KUMCC 19-0093 F
100/1.00
Pseudoastrosphaeriella bambusae KUMCC 19-0091 F
86/0.98
Pseudoastrosphaeriella bambusae MFLUCC 11-0205 F
Pseudoastrosphaeriella longicolla MFLUCC 11-0171 F
90/0.98
Pseudoastrosphaeriella thailandensis MFLUCC 11-0144
Pseudoastrosphaeriella thailandensis MFLUCC 10-0553
Pseudoastrosphaerie
-llaceae
100/1.00
Pseudoastrosphaeriella thailandensis MFLUCC 14-0038
100/--
Pseudoastrosphaeriella africana MFLUCC 11-0176
Pseudoastrosphaeriella aquatica MFLUCC 18-0984 F
Pseudoastrosphaeriella aquatica MFLUCC 18-0991 F
100/1.00
Pseudoastrosphaeriella aquatica KUMCC 19-0096 F
Ulospora bilgramii AFTOL-ID 1598
Verruculina enalia BCC 18401
Testudinaceae
Angustospora nilensis MFLU 15-1511 F
Lepidosphaeria nicotiae AFTOL-ID 1576
Ligninsphaeria jonesii MFLUCC 15-0641
100/0.97
Ligninsphaeria jonesii GZCC 15-0080
98/--
Ligninsphaeriaceae
Amniculicola aquatica MFLUCC 16-0915 F
Amniculicola aquatica MFLUCC 16-1123 F
Amniculicola lignicola Ying 01 F
81/-89/1.00
Anguillospora longissima CCMF 10304
Anguillospora longissima CS869 F
Amniculicola parva CBS 123092 F
Amniculicola guttulata DLUCC 0538 F
90/1.00
85/1.00
Amniculicola guttulata MFLUCC 16-0907 F
Amniculicola guttulata MFLUCC 18-1038 F
Amniculicola guttulata MFLUCC 16-1297
91/0.98
Amniculicolaceae
Vargamyces aquaticus CBS 636.91
Vargamyces aquaticus HKUCC 10830
Amniculicola immersa CBS 123083 F
Pseudomassariosphaeria grandispora CBS 613.86
100/0.99
100/0.95
83/0.99
84/1.00
Pseudomassariosphaeria bromicola IT 1333
Murispora cardui MFLUCC 13-0761
Murispora medicaginicola MFLUCC 13-0762
Neomassariosphaeria typhicola CBS 123126 F
Jahnula Kirschst., Annls mycol. 34(3): 196 (1936)
Saprobic on submerged wood. Sexual morph: Ascomata semi-immersed to erumpent, become superficial with
base remaining immersed, solitary or clustered in small
groups, globose to subglobose, unilocular, brown to dark
brown, membranous, with a stalk-like strand or stoloniferous hyphae attached to the substratum, or covered by sparse
hair-like projections, with ostiolate papilla. Peridium variable in thickness, comprising a few layers of relatively
large, thin-walled, light brown cells of textura angularis.
Pseudoparaphyses cellular, persistent, filiform or hyphalike, hyaline, sparsely septate. Asci 8-spored, bitunicate,
fissitunicate, mostly cylindrical, short-pedicellate, with an
ocular chamber and sometimes with a faint ring. Ascospores
mostly uniseriate, ellipsoid-fusiform, apical cell slightly
larger, both cells tapering with rounded ends, reddish brown
or dark brown, l-septate, straight or curved (definition sensu
stricto from Hawksworth (1984), Hyde and Goh (1999b) and
Raja and Shearer (2006)). Asexual morph: Undetermined.
Type species: Jahnula aquatica (Kirschst.) Kirschst.
13
336
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
Lindgomyces rotundatus KH 114 F
Lindgomyces rotundatus KT 966 F
81/--
Lindgomyces rotundatus KT 1096 F
Lindgomyces rotundatus KT 1107 F
Lindgomyces cigarospora G619-2 F
100/0.99
Lindgomyces cigarospora G619 F
Lindgomyces cigarospora G619-3 F
Lindgomyces carolinensis G834 F
92/--
Lindgomyces carolinensis G833 F
Lindgomyces carolinensis G618-2 F
Lindgomyces carolinensis G618 F
98/0.97
Lindgomyces apiculatus KT 1144 F
Lindgomyces apiculatus KT 1108 F
99/0.97
Lindgomyces lemonweirensis A632-1a F
Lindgomyces lemonweirensis A632-1b F
Lindgomyces angustiascus G202-1a F
Lindgomyces angustiascus F60-1 F
Lindgomyces
Lindgomyces angustiascus A640-1a F
83/-90/1.00
100/0.96
Lindgomyces ingoldianus KH 100 F
Lindgomyces ingoldianus ATCC 200398 F
Lindgomyces sp. KH 241 F
Lindgomyces aquaticus MFLUCC 18-1416 F
96/1.00
Lindgomyces griseosporus CBS 123100 F
Lindgomyces griseosporus BJFC200094 F
100/1.00
89/--
Lindgomyces pseudomadisonensis KT 2742 F
Lindgomyces madisonensis G416a F
Lindgomyces madisonensis G416b F
100/1.00
Lindgomyces cinctosporus R56-3 F
Lindgomyces cinctosporus R56-1 F
Lindgomyces okinawaensis KT 3531 F
95/--
Lindgomyces breviappendiculatus KT 1215 F
Lindgomyces breviappendiculatus KT 1399 F
Notes: Jahnula is undoubtedly polyphyletic (Hyde et al.
2013, 2017, 2019; Huang et al. 2018) and Jahnula sensu
stricto was mentioned in Huang et al. (2018), and Hyde
et al. (2013). Hyde et al. (2013) suggested that taxonomic
changes are needed for taxa in Jahnula sensu lato, but more
jahnula-like species need to be collected and sequenced. We
accept three species, J. aquatica, J. granulosa K.D. Hyde
& S.W. Wong and J. rostrata Raja & Shearer in Jahnula
sensu stricto, which have been collected from freshwater
habitats (listed below). These three species share common characters in having dark ascomata with few appendages attached to the base, and ellipsoid-fusiform, brown,
13
l-septate ascospores with wider apical cell (Hawksworth
1984; Hyde and Wong 1999; Raja and Shearer 2006).
Asci of J. aquatica are cylindrical, those of J. granulosa
are obclavate and those of J. rostrata are clavate. Jahnula
aquatica has smooth-walled ascospores without a sheath,
while J. granulosa has granular-walled ascospores with a
thin mucilaginous sheath. Jahnula rostrata is similar to J.
granulosa by rough-walled ascospores, but differs in having
an irregularly striated pattern. These three species formed
a well-supported and stable clade in previous publications
(Huang et al. 2018; Hyde et al. 2019) and in this study
(Fig. 14).
Fungal Diversity (2020) 105:319–575
337
Fig. 2 (continued)
Clohesyomyces aquaticus MFLUCC 11-0092 F
100/1.00
Clohesyomyces aquaticus MFLUCC 18-1037 F
Clohesyomyces aquaticus MFLUCC 15-0979 F
92/1.00
83/--
Trematosphaeria hydrela CBS 880.70
96/1.00
87/0.99
Trematosphaeria hydrela HKUCC 10666
Aquimassariosphaeria kunmingensis KUMCC 18-1019 F
Aquimassariosphaeria typhicola CBS 609.86
Massariosphaeria sp. KT 667
Massariosphaeria sp. KT 797
99/1.00
Lolia aquatica CBS 22130 F
--/0.95
Lolia aquatica MF 644
--/1.00
Lindgomycetaceae
Lolia dictyospora CBS 22131 F
99/1.00
75/1.00
Hongkongmyces pedis HKU 35
Hongkongmyces snookiorum DAOMC 251900 F
84/1.00
Hongkongmyces aquaticus MFLUCC 18-1150 F
Hongkongmyces thailandicus MFLUCC 16-0406
Arundellina typhae MFLUCC 16-0310 F
100/1.00
Arundellina typhae MFLUCC 16-0309
Neolindgomyces pandanae MFLUCC 18-0245
100/--
Neolindgomyces submersa CBS 115553
Salsuginea ramicola CBS 125781
Salsuginea ramicola CBS 125780
100/0.96
Salsuginea ramicola KT 2597.2
Salsugineaceae
Salsuginea ramicola KT 2597.1
100/0.99
Fissuroma bambusae MFLUCC 11-0160
Fissuroma bambusae MFLUCC 11-0198
100/1.00
100/--
Fissuroma neoaggregata MFLUCC 13-0227
Fissuroma neoaggregata MFLUCC 10-0554
Fissuroma thailandicum MFLUCC 11-0189
92/-100/0.98
97/--
Fissuroma thailandicum MFLUCC 11-0206
Fissuroma aggregata KT 984
75/--
Fissuroma aggregata KT 767
98/1.00Fissuroma maculans MFLUCC10-0886
94/--
Fissuroma maculans MFLUCC 10-0887
Fissuroma taiwanense FU30861
92/--
100/--
Fissuroma taiwanense FU30862
Fissuroma caryotae MFLU 17-1253
99/--
Fissuroma calami MFLUCC 13-0836
Aigialaceae
Fissuroma calami MFLUCC 17-2030
99/1.00
100/1.00
Neoastrosphaeriella krabiensis MFLUCC 11-0025
Neoastrosphaeriella krabiensis MFLUCC 11-0022
Neoastrosphaeriella aquatica MFLUCC 18-0209 F
97/--
100/1.00
Aigialus grandis BCC 18419
Aigialus grandis BCC 20000
100/-87/-95/--
Aigialus mangrovis BCC 33564
Aigialus rhizophorae BCC 33572
Ascocratera manglicola HHUF 30032
Rimora mangrovei JK 5246A
13
338
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
98/--
Wicklowia aquatica F76-2 F
98/--
Wicklowia aquatica AF289-1 F
100/0.95
Wicklowia aquatica CBS 125634 F
Wicklowiaceae
Wicklowia submersa MFLUCC 18-0373 F
Astrosphaeriella bambusae MFLUCC 10-0095
Astrosphaeriella bambusae MFLUCC 13-0230
100/0.99
Astrosphaeriella stellata KT 998
Astrosphaeriella neostellata MFLUCC 11-0625
98/0.97
Astrosphaeriella thysanolaenae MFLUCC 11-0186
91/--
Astrosphaeriella thailandica MFLUCC 11-0191
100/--
Astrosphaeriella neofusispora MFLUCC 11-0161
Astrosphaeriella fusispora MFLUCC 10-0555
Pteridiospora javanica MFLUCC 11-0159
100/1.00
Pteridiospora javanica MFLUCC 11-0195
100/--
Pteridiospora bambusae MFLU 18-0071
Pteridiospora chiangraiensis MFLUCC 11-0162
Pithomyces vesuvius MTCC 12224
77/-100/1.00
Pithomyces flavus MTCC 12224
Pithomyces flavus LF 156
76
99/1.00
84/--
Pithomyces caryotae MFLUCC 13-0828
99/--
Pithomyces caryotae MFLUCC 13-0824
Pithomyces licualae MFLUCC 17-2031
Quercicola fusiformis MFUCC 18-0479
100/--
Quercicola guttulospora MFUCC 18-0481
100/1.00
Acrocordiopsis patilii BCC28167
Acrocordiopsis patilii BCC28166
Xenoastrosphaeriella tornata MFLUCC 11-0196
Aquatospora cylindrica MFLUCC 18-1287 F
Caryospora quercus MFLU 18-2151
100/1.00
Caryospora quercus MFLUCC 17-2342 F
Caryospora quercus MFLUCC 17-2323 F
99/0.99
83/--
Caryospora submersa MFLUCC 18-1283 F
100/1.00
100/0.99
Caryospora submersa MFLUCC 18-1409 F
Caryospora aquatica MFLUCC 18-1030 F
99/1.00
Caryospora aquatica MFLUCC 11-0008 F
“Caryospora minima”
Astrosphaeriellopsis caryotae MFLUCC 13-0832
87/0.99
100/1.00
Astrosphaeriellopsis caryotae MFLUCC 13-0830
Astrosphaeriellopsis caryotae MFLUCC 13-0833
Astrosphaeriellopsis bakeriana CBS 115556
Astrosphaeriellopsis bakeriana MFLUCC 11-0027
Acuminatispora palmarum MFLUCC 18-0461
Acuminatispora palmarum MFLUCC 18-0460
100/1.00
92/--
Acuminatispora palmarum MFLUCC 18-0462
Acuminatispora palmarum MFLUCC 18-0264
Zopfia rhizophila CBS 207.26
99/--
Delitschiaceae
100/0.96
Massariaceae
13
Astrosphaeriellaceae
Fungal Diversity (2020) 105:319–575
339
Fig. 2 (continued)
Hysteriales
100/0.99
Mytilinidiales
Pseudorobillarda eucalypti MFLUCC 12-0417
100/1.00
Pseudorobillarda sojae BCC 20495
100/--
Pseudorobillarda bolusanthi CBS 145072
87/--
Pseudorobillarda
ceae
Pseudorobillarda phragmitis CBS 398.61
Minutisphaera japonica JCM 18560
81/--
Minutisphaera japonica JCM 18562
99/0.95
Minutisphaera japonica JF08056
Minutisphaera japonica JCM 18561
99/1.00
100/1.00
Minutisphaera aspera G427-1b F
Minutisphaera aspera G427-1a F
94/0.99
95/1.00
Minutisphaera parafimbriatispora G156-4b
87/1.00
Minutisphaerales
Minutisphaera parafimbriatispora G156-4a
99/1.00
Minutisphaera parafimbriatispora G156-1b
100/1.00
Minutisphaera parafimbriatispora G156-1a
99/0.95
Minutisphaera fimbriatispora A242-7c F
98/0.96
Minutisphaera fimbriatispora A242-7d F
Minutisphaera fimbriatispora A242-8a
Minutisphaera fimbriatispora G155-1a F
Helicotruncatum palmigenum NBRC 32663
100/1.00
Helicotruncatum palmigenum MFLUCC 15-0993 F
83/-100/1.00
Tubeufia aquatica MFLUCC 16-1249
100/--
Tubeufiales
Tubeufia aquatica MFLUCC 15-0990 F
Neohelicomyces thailandicus MFLUCC 11-0005 F
100/1.00
Neohelicomyces dehongensis MFLUCC 18-1029 F
100/1.00
Acrospermales
77/-87/--
100/1.00
Muyocopronales
Dyfrolomyces tiomanensis MFLUCC 13-0440
86/-100/1.00
100/--
Macrovalsaria megalospora MFLU 15-3432
Macrovalsaria megalospora MFLU 15-3431
98/1.00
Stigmatodiscales
82/--
Strigulales
Kirschsteiniothelia submersa S-601 F
100/1.00
100/0.99
Kirschsteiniothelia submersa S-481 F
Kirschsteiniothelia submersa MFLUCC 15-0427 F
100/0.99
Kirschsteiniothelia rostrata MFLUCC 15-0619 F
Kirschsteiniothelia rostrata MFLUCC 16-1124 F
100
100/0.99
Kirschsteiniothelia aethiops S-783 F
Kirschsteiniothelia aethiops MFLUCC 16-1104 F
100/1.00
100/0.97
Kirschsteiniothelia aethiops MFLUCC 15-0424 F
Kirschsteiniotheli
ales
Kirschsteiniothelia aethiops CBS 109.53
100/1.00
Kirschsteiniothelia cangshanensis MFLUCC 16-1350 F
Kirschsteiniothelia aquatica MFLUCC 17-1685 F
100/0.96
99/--
Kirschsteiniothelia fluminicola MFLUCC 16-1263 F
Valsariales
97
The type species, Jahnula aquatica commonly occurs
in freshwater habitats as listed below. The lectotype of J.
aquatica was examined, illustrated and described by Hawksworth (1984). Jahnula aquatica is characterized by ascomata attached to the substratum by subiculum-like hyphae,
peridium uneven in thickness, cylindrical asci, elongateellipsoid to very broadly fusiform, reddish brown, smooth,
moderately thick-walled ascospores without a distinct
gelatinous sheath (Hawksworth 1984). The sequences of
two isolates R68-1 and R68-2 (not type) were obtained by
Raja and Shearer (2006), recognized and used for current
phylogenetic analyses (Suetrong et al. 2011a; Huang et al.
2018; Hyde et al. 2019).
Xylomyces chlamydosporus Goos et al. was considered to
be the asexual state of Jahnula aquatica based on fusiform
chlamydospores produced in culture (Sivichai et al. 2011).
However, the phylogenetic analyses did not support this conclusion because J. aquatica and X. chlamydosporus scattered
13
340
Fungal Diversity (2020) 105:319–575
Fig. 2 (continued)
97/1.00
Myriangiales
91/-100/1.00
82/--
Endosporium aviarium UAMH 10530
Endosporiuaceae
Endosporium populi-tremuloides UAMH 10529
99/--
100/1.00
Dothideales
100/1.00
Capnodiales
Homortomycetaceae
Homortomyces tamaricis MFLUCC 13-0441
100/-100/1.00
85/0.99
Botryosphaeria dothidea CMW 8000
Botryosphaeria dothidea CBS 115476
Botryosphaeria agaves MFLUCC 10 0051 LSU
85/--
Botryosphaeriales
Cophinforma atrovirens MFLUCC 11-0425
100/--
Bahusutrabeeja dwaya GUFCC 4904
Phyllosticta citricarpa CBS 102374
100/1.00
Asterinaceae
Strigula nemathora MPN72
100/1.00
Collemopsidiales
100/1.00
Eremomycetaceae
Jahnula granulosa SS1567
84/-82/--
Jahnula rostrata F4-3
Jahnula aquatica R68-1 F
85/--
Megalohypha aqua-dulces AF005-2a F
100/0.99
100/1.00
Jahnulales
Megalohypha aqua-dulces AF005-2b F
Ascagilis sunyatsenii UBC-F13876
100/1.00
Monoblastiales
95/--
Asterinaceae
100/1.00
Cladoriella rubrigena CBS 124760
100/--
Cladoriella eucalypti CPC 10953
98/1.00
Abrothallales
99/1.00
Lichenotheliaceae
85/--
Patellariales
100/1.00
100/1.00
Natipusillales
Natipusilla decorospora AF236-1A F
Natipusilla limonensis AF286-1A F
Natipusilla naponensis AF217-1B F
Natipusilla bellaspora PE91-1b F
100/1.00
Zeloasperisporiales
100/1.00
Venturiaceae
100/1.00
Phaeotrichales
96/1.00
Microthyriales
100/1.00
Endocarpon pallidulum AFTOL-ID 661
OUTGROUP
Capronia pilosella AFTOL-ID 657
0.3
in different clades within Jahnulales (Campbell et al. 2007;
Suetrong et al. 2011a; this study, Fig. 14).
We introduce two new genera Neojahnula and Pseudojahnula to accommodate two known species Jahnula australiensis K.D. Hyde and J. potamophila K.D. Hyde & S.W.
Wong, respectively. Seven species, i.e. J. bipolaris, J. guttulaspora, J. queenslandica, J. seychellensis, J. submersa, J.
sunyatsenii and J. thailandensis, which are phylogenetically
distant from J. aquatica, are transferred to Ascagilis K.D.
Hyde. Nine species, i.e. J. apiospora, J. appendiculata, J.
bipileata, J. dianchia, J. morakotii, J. poonythii, J. purpurea, J. sangamonensis and J. systyla, are placed in Jahnula
sensu lato.
For convenience and practicality, we have excluded Jahnula systyla K.D. Hyde & S.W. Wong from the key, because
we believe it may be phylogenetically related to Aliquandostipite, pending future phylogenetic studies (Fig. 6).
13
List of freshwater Jahnula sensu stricto species
*Jahnula aquatica (Plöttner & Kirschst.) Kirschst., Ann.,
Mycol. 34: 196 (1936); Fig. 7
Basionym: Melanopsamma aquatica Kirschst., Krypt.-Fl.
Brandenburg (Leipzig) 7(2): 226 (1911)
Freshwater distribution: China (Ho 1998), France
(Fournier et al. 2010), Germany (Kirschstein 1936; Hawksworth 1984), Malaysia (Ho et al. 2001), South Africa (Hyde
and Wong 1999), Thailand (Sivichai et al. 2011), USA (Raja
and Shearer 2006; Raja et al. 2009b)
*Jahnula granulosa K.D. Hyde & S.W. Wong, Nova Hedwig. 68: 497 (1999)
Freshwater distribution: China (Cai et al. 2002a), South
Africa (Hyde and Wong 1999), Thailand (Suetrong et al.
2011a)
Fungal Diversity (2020) 105:319–575
341
Fig. 3 Aliquandostipite spp.
(Material examined: USA, Tennessee, Great Smoky Mountains
National Park, Elkmont Campground, a small stream between
Jakes Creek and Little River, on
submerged, decorticated wood,
5 July 2002, H.A. Raja & N.
Hamburger, R76-1; Florida,
Big Cypress National Preserve,
Cypress Swamp Loop Road, on
submerged decorticated wood,
22 March 2005, H.A. Raja, J.L.
Crane & A.N. Miller, F89-1).
a–d, f A. khaoyaiensis (F89-1).
a Stalked ascoma. b Ascospore
with slug-like gelatinous
sheath in nigrosin. c, d Asci.
f Ascospore with thick-walled
gelatinous sheath. e, g–i A.
crystallinus (R76-1). e Ascoma.
g, h Asci. i Ascospores emerging from the ascus to form a
large, C-shaped to fusiform,
gelatinous sheath. Scale bars:
a = 100 µm, b–d = 20 µm, e =
200 µm, f–i = 20 µm
*Jahnula rostrata Raja & Shearer, Mycologia 98: 325
(2006)
Facesoffungi number: FoF09237; Fig. 8
Freshwater distribution: China (this study), USA (Raja
and Shearer 2006; Raja et al. 2009b)
Saprobic on submerged wood. Sexual morph: Ascomata
320–420 × 310–330 μm, superficial, solitary to scattered,
subglobose or pyriform, brown to black, rough-walled, with
ostiolate papilla. Neck 48–75 × 74–98 μm, composed of subglobose cells diverging from the ostiole. Peridium 25–40
μm wide, two-layered, outer layer composed of large, dark
brown, thick-walled cells of textura angularis, inner layer
composed of compressed, hyaline, thin-walled cells of
textura angularis. Pseudoparaphyses 2.5–3.5 μm wide, cellular, hypha-like, hyaline, septate, unbranched. Asci 135–200
× 21–32 μm ( x̄ = 160 × 28 μm, n = 15), 8-spored, bitunicate, fissitunicate, clavate, short pedicellate, with an ocular
chamber. Ascospores 28–40 × 11–17 μm ( x̄ = 35 × 13.5
μm, n = 15), overlapping biseriate, ellipsoidal or broadly
fusiform, with rounded or occasionally acute ends, upper
cell slightly broader than lower cell, hyaline to brown when
young, dark brown at maturity, 1-septate, constricted at the
septum, with thick and dark band at septum when mature,
rough-walled, with minute striated pattern, guttulate, with or
without sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colonies irregular,
reaching 35 mm in 30 days at 25 °C, dark from above and
below, dry, hairy, dense in the centre, becoming sparse at
the edge.
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Fig. 4 Aliquandostipite khaoyaiensis a–h, l, m (MFLU 18-1573,
new geographical record), i–k (MFLU 18-1554, new geographical record). a, b, i Ascomata on host surface. c Vertical section of
ascoma. d Structure of peridium. e Pseudoparaphyses. f, j Bituni-
cate asci. g, h, k Ascospores. l, m Colony on PDA (left-front, rightreverse). Scale bars: c = 100 μm, d = 30 μm, e, g, h = 20 μm, f, j, k
= 50 μm
Material examined: CHINA, Yunnan Province, Kunming,
Liangwang National Wetland Park, on submerged wood in
a stream, 19 March 2019, C.X. Li, L15 (MFLU 20-0435).
Notes: Our collection MFLU 20-0435 is identified as Jahnula rostrata based on very similar morphology and only
one and one nucleotide difference in LSU and SSU sequence
data between MFLU 20-0435 and F4-3, respectively. They
form a well-supported clade in our phylogenetic analysis
(Fig. 14). MFLU 20-0435 has thinner asci than the holotype
(135–200 × 21–32 μm vs. 152–190 × 32–40 μm) (Raja and
Shearer 2006). Our collection is a new geographical record
for J. rostrata from China. Jahnula rostrata (F4-3) lacks ITS
sequence data, we supplement ITS sequence for this species
in this study.
Key to freshwater Jahnula sensu stricto species
13
1.
1.
2.
2.
Ascospores smooth-walled…………………J. aquatica
Ascospores rough-walled………………………………2
Ascospores granular-walled………………J. granulosa
Ascospores with an irregularly striated pattern………
………………………………………………J. rostrata
List of freshwater Jahnula sensu lato species
Jahnula apiospora A. Carter, Raja & Shearer, Mycoscience
49: 326 (2008); Fig. 6d, i
Freshwater distribution: Canada (Raja et al. 2008)
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Fig. 5 Aliquandostipitaceae spp. (Material examined: USA, Florida,
Monroe County, freshwater swamp at Big Cypress National Preserve, on submerged decorticated wood, 17 February 2006, H.A. Raja
& J.L. Crane, ILL 40108 = F117-1, holotype; PANAMA, Colon,
Soberania National Park, Juan Grande, on submerged, decorticated
wood, 12 January 2003, A. Ferrer, AF005). a–e Aliquandostipite
minuta (F117-1). a Squash mount of ascoma. b, c Asci. d Extended
endoascus releasing ascospore. e Ascospore showing fusiform sheath
and filamentous appendages. f–j Megalohypha aqua-dulces (AF005).
f Ascoma on substrate. g Ascus. h, i Ascospores. j Broad hyphae in
PDA. Scale bars: a–e, g–i = 20 µm, f, j = 200 µm
*Jahnula appendiculata Pinruan, K.D. Hyde & E.B.G.
Jones, Sydowia 54: 243 (2002); Fig. 9
Freshwater distribution: Peru (Shearer et al. 2015), Thailand (Pinruan et al. 2002; Sivichai and Boonyuen 2010)
Jahnula poonythii K.D. Hyde & S.W. Wong, Nova Hedwig.
68: 499 (1999)
Freshwater distribution: China (Cai et al. 2002a; Luo
et al. 2004), Mauritius (Hyde and Wong 1999), Mexico
(Gonzalez and Chavarria 2005)
*Jahnula bipileata Raja & Shearer, Mycologia 98: 321
(2006); Fig. 6c, j
Freshwater distribution: USA (Raja and Shearer 2006)
Jahnula morakotii Sivichai & Boonyuen, Mycotaxon 112:
476 (2012)
Freshwater distribution: Thailand (Sivichai and Boonyuen 2010)
Jahnula purpurea J. Fourn., Raja & Shearer, Mycokeys 9:
30. (2015)
Freshwater distribution: Martinique (Fournier et al. 2015)
*Jahnula sangamonensis Shearer & Raja, Mycologia 98:
327 (2006)
Freshwater distribution: USA (Raja and Shearer 2006;
Raja et al. 2009b)
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Fig. 6 Aliquandostipitaceae spp. (Material examined: USA, Florida,
on submerged wood, July 2006, J.L. Crane & H.A. Raja, F111; Costa
Rica, La Selva Biological Station, Sura 60, on submerged, corticated
wood, CMP, A492; Florida, Apalachicola National Forest, swampy
area of Whitehead Lake, on submerged, decorticated wood, 13 July
2004, HAR and CB, F49-1 (ILL), holotype; CANADA, Prince
Edward Island, on submerged decorticated wood, in a small creek
beside dirt road near Bonshaw and Trans Canada Highway, 13 Octo-
ber 2007, A. Carter, AC-706, ILL40554, holotype). a, e, h Pseudojahnula potamophila (F111). a Ascomata attached to the wood with
broad hyphae. e, h Ascospores. b Section of ascoma of Ascagilis
seychellensis (A492). c, j Jahnula bipileata (F49-1). c Ascoma. j
Ascospores. d, i Jahnula apiospora (AC-706). d Structure of peridium. i One-septate apiosporous ascospore. f, g Ascospores of Ascagilis bipolaris showing bipolar appendages. Scale bars: a, c = 200 µm,
b, d–j = 20 µm
Jahnula systyla K.D. Hyde & S.W. Wong, Nova Hedwig.
68: 506 (1999)
Freshwater distribution: Australia (Hyde and Wong
1999), China (Ho 1998), Malaysia (Ho et al. 2001)
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345
Fig. 7 Jahnula aquatica (Material examined: USA, Illinois,
Salt Fork Association Lake
(SFA Lake), a manmade private
lake, on submerged, decorticated wood, 15 October 2002,
HAR, R68-1). a Squash mount
of ascoma. b Ascus. c Pseudoparaphyses. d, e Ascospores.
Scale bars: a = 100 µm, b, c, e
= 20 µm, d = 10 µm
Key to freshwater Jahnula sensu lato species
1. Ascospores without modification (no appendages/sheath/
caps)……………………………………………………2
1. Ascospores equipped with modification (appendages/
sheath/caps)……………………………………………6
2. Ascospores asymmetric……………………J. apiospora
2. Ascospores symmetric…………………………………3
3. Ascomata stain substrate purple……………J. purpurea
3. Ascomata not as above…………………………………4
4. Ascomata with a short but prominent neck, endoasci
extend up to 500 µm long in water………………………
………………………………………J. sangamonensis
4. Ascomata with a short papilla, endoasci not extending……………………………………………………5
5. Ascomata with stalk-like strands attached to the base,
with a few sparse hair-like projections…………………
……………………………………………J. poonythii
5. Ascomata without any appendages………J. dianchia
6. Ascospores without unfurling bipolar appendages…
………………………………………………J. bipileata
6. Ascospores with long unfurling bipolar appendages……………………………………………………7
7. Ascospores 18–20 × 5–6 μm………………J. morakotii
7. Ascospores 48–55 × 23–26 μm………J. appendiculata
Ascagilis K.D. Hyde, Aust. Syst. Bot. 5(1): 109 (1992)
Saprobic on submerged wood. Sexual morph: Ascomata
semi-immersed or superficial, solitary, scattered or gregarious, globose or subglobose, coriaceous, black, ostiolate.
Peridium comprising several layers of thin-walled, brown,
angular cells. Pseudoparaphyses numerous, cellular, hyaline,
thin, septate, branched. Asci 8-spored, clavate, obclavate,
broadly or narrowly cylindrical, fissitunicate, sometimes
expanding 4–5 times longer than original length, with an
ocular chamber and ring. Ascospores bi- to tri-seriate, 1-septate, brown, mostly with a refractive mucilaginous pad at
each end. Asexual morph: Undetermined.
Type species: Ascagilis bipolaris K.D. Hyde
Notes: Ascagilis was introduced to accommodate a jahnula-like species, A. bipolaris, characterized by clavate to
cylindrical asci with an ocular chamber and ring and fusiform to ellipsoidal, 1-septate, brown ascospores with a pad at
each end (Hyde 1992b). Ascagilis was suppressed since the
type species A. bipolaris was transferred to Jahnula (Hyde
and Wong 1999) and they thought the fusiform to ellipsoidal
ascospores with bipolar pads of Ascagilis was insufficient
to separate these two genera. Jahnula has been shown to
be polyphyletic in previous publications (Hyde et al. 2013,
2017, 2019; Huang et al. 2018) and in this study. The clade
comprising A. bipolaris and six other species is phylogenetically distant from Jahnula sensu stricto (Fig. 14). Additionally, most species including our new species, A. submersa
and A. thailandensis, have a refractive mucilaginous pad
at each end, which is lacking in Jahnula sensu stricto. All
species in this clade cluster with strong bootstrap support
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Fig. 8 Jahnula rostrata (MFLU 20-0435, new geographical
record). a Appearance of black ascoma on host substrate. b Structure
of ostiole. c Pseudoparaphyses. d Vertical section of ascoma. e Struc-
ture of peridium. f–i Ascospores. j–m Bitunicate asci. n Germinated
ascospore. o, p Colony on PDA (left-front, right-reverse). Scale bars:
b, e–n = 30 μm, c = 10 μm, d = 50 μm
(Fig. 14). We therefore, resurrect Ascagilis and transfer four
species and introduce two new species in the genus (see list
below). The mucilaginous pad of ascospores is treated as a
noticeable characteristic of Ascagilis.
Synonymy: Jahnula bipolaris (K.D. Hyde) K.D. Hyde,
Nova Hedwigia 68(3–4): 494 (1999)
Freshwater distribution: Australia (Hyde 1992b; Hyde
and Wong 1999), China (Tsui et al. 2000; Tsui and Hyde
2004), Costa Rica (Raja and Shearer 2006), Malaysia (Ho
et al. 2001), Peru (Shearer et al. 2015), Thailand (Suetrong
et al. 2011a), USA (Raja et al. 2009b)
List of freshwater Ascagilis species
*Ascagilis bipolaris K.D. Hyde, Aust. Syst. Bot. 5(1): 111
(1992); Fig. 6f, g
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347
Fig. 9 Jahnula appendiculata (Material examined: PERU, on submerged wood, S. Zelski & H.A. Raja, PE0010). a Stalked ascoma.
b Broad hypha attached to the ascomatal sides and bases. c Young
ascus. d, e Ascospores with broad gelatinous sheath and polar
appendages indicated by arrows. Scale bars: a = 50 µm, b–e = 20 µm
*Ascagilis guttulaspora (Qing Tian, Y.Z. Lu & K.D. Hyde)
W. Dong, Doilom & K.D. Hyde, comb. nov.
Index Fungorum number: IF557893; Facesoffungi number: FoF09228
Basionym: Jahnula guttulaspora Qing Tian, Y.Z. Lu &
K.D. Hyde, Fungal Diversity 87: 6 (2017)
Freshwater distribution: China (Hyde et al. 2017)
*Ascagilis seychellensis (K.D. Hyde & S.W. Wong) W.
Dong, Doilom & K.D. Hyde, comb. nov.
Index Fungorum number: IF557895; Facesoffungi number: FoF09230
Basionym: Jahnula seychellensis K.D. Hyde & S.W.
Wong, Nova Hedwig. 68: 504 (1999); Fig. 6b
Freshwater distribution: Brazil (Barbosa et al. 2013),
Costa Rica (Raja and Shearer 2006), Seychelles (Hyde and
Wong 1999)
*Ascagilis queenslandica (Dayarathne, Fryar & K.D. Hyde)
W. Dong, Doilom & K.D. Hyde, comb. nov.
Index Fungorum number: IF557894; Facesoffungi number: FoF09229
Basionym: Jahnula queenslandica Dayarathne, Fryar &
K.D. Hyde, Fungal Diversity 96: 91 (2019)
Freshwater distribution: Australia (Hyde et al. 2019)
*Ascagilis submersa W. Dong, H. Zhang & K.D. Hyde, sp.
nov.
Index Fungorum number: IF557896; Facesoffungi number: FoF09238; Fig. 10
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Fig. 10 Ascagilis submersa
(MFLU 18-1527, holotype). a
Ascomata on host with associated appendages (arrow). b
Vertical section of ascoma. c
Structure of peridium. d Pseudoparaphyses. e–h Bitunicate
asci. i Ascospores. j Ascospore
in Indian Ink. k Germinated
ascospore. l, m Colony on PDA
(left-front, right-reverse). Scale
bars: b = 50 μm, c–k = 20 μm
Etymology: in reference to the submerged habitats of the
fungus
Holotype: MFLU 18-1527
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 160–250 μm diam., black, solitary
or scattered, superficial, globose or subglobose, with several
long, brown to dark brown appendages which are procumbent on the substrate surface, membranous. Peridium 30–35
μm thick, composed of 4–5 layers of thick-walled, olive to
dark brown, compressed cells of textura angularis, with
13
2 layers of flattened, compressed, hyaline cells inwardly.
Pseudoparaphyses 3 μm diam., numerous, cellular, hyphalike, hyaline, septate, embedded in a gelatinous matrix. Asci
110–160 × 35–42 μm (x̄ = 135 × 38 μm, n = 10), 8-spored,
bitunicate, thick-walled, broadly obclavate, with thick-walled
apex and well-developed ocular chamber, completely filled
by ascospores, short pedicel observed only when young.
Ascospores 37.5–41 × 17–19 μm ( x̄ = 40 × 18 μm, n = 20),
overlapping biseriate and uniseriate near the apex, straight
or slightly curved, hyaline when young, pale brown to brown
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when mature, 1-septate, constricted at the septum, ellipsoidal, guttulate, thin-walled, smooth, with bipolar, helmetshaped appendages. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 10 days at 25 °C, dark brown from above
and below, surface rough, with sparse mycelium, dry, edge
entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
20180513-1 (MFLU 18-1527, holotype), ex-type living culture MFLUCC 18-1143; ibid., 20180513-2 (HKAS 104996,
isotype), ex-type living culture KUMCC 19-0013.
Notes: Ascagilis submersa clusters with A. queenslandica, A. sunyatsenii and A. thailandensis (Fig. 14).
Ascagilis submersa is similar to A. sunyatsenii in having
ellipsoidal, pale brown ascospores with bipolar helmetshaped appendages (Inderbitzin et al. 2001). However, A.
sunyatsenii has saccate, ovoid to elongate asci and longer
ascospores ((39–)46–52 × 16–23 μm) contrasting with
broadly obclavate asci and shorter ascospores (37.5–41 ×
17–19 μm) in A. submersa (Inderbitzin et al. 2001). Ascagilis submersa has olive peridium and ascospores, which are
unique in the genus and different from the dark pigmented
peridium and light brown ascospores of A. sunyatsenii.
Additionally, A. sunyatsenii has two types of ascomata, sessile and stalked, while A. submersa has only sessile ascomata and with several appendages which are procumbent on
the substrate surface (Inderbitzin et al. 2001). Because of
scarcity of some nucleic acid genes, only ITS sequence can
be compared. There are two nucleotide differences in ITS
sequence data, but only crossing 207 nucleotides. The distinct morphology of A. submersa provides strong evidence
to be a new species in Ascagilis
*Ascagilis sunyatsenii (Inderb.) W. Dong, Doilom & K.D.
Hyde, comb. nov.
Index Fungorum number: IF557897; Facesoffungi number: FoF09239
Basionym: Aliquandostipite sunyatsenii Inderb., Am. J.
Bot. 88(1): 57 (2001)
Synonymy: Jahnula sunyatsenii (Inderb.) K.L. Pang,
E.B.G. Jones & Sivichai, Mycol. Res 106: 1037 (2002)
Freshwater distribution: China (Inderbitzin et al. 2001;
Pang et al. 2002)
*Ascagilis thailandensis W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557898; Facesoffungi number: FoF09240; Fig. 11
Etymology: in reference to Thailand, where the holotype
was collected
Holotype: MFLU 18-1514
349
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 280–320 μm high, 200–250 μm
diam., black, scattered, superficial with base immersed in
host tissue, conical, membranous, coriaceous, with short
ostiolate papilla. Peridium 30–40 μm thick, composed of
4–5 layers of thin-walled, pale brown, large cells of textura
angularis. Pseudoparaphyses 3.5 μm diam., numerous, cellular, hypha-like, hyaline, septate. Asci 125–185 × 24–30 μm
(x̄ = 162 × 27.5 μm, n = 5), 8-spored, bitunicate, broadly or
narrowly cylindrical, with an ocular chamber, short pedicellate. Ascospores 33–48.5 × 11–16 μm ( x̄ = 41 × 13 μm, n
= 30), biseriate or uniseriate above ascus center and overlapping biseriate near the center or base, brown, 1-septate,
slightly constricted at the septum, irregularly fusiform,
mostly curved, tapering at apical cell and slightly flattened
or rounded at the basal cell, guttulate, thin-walled, smooth,
with small, inconspicuous pads at one or both apices. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 5 days at 25 °C, olive brown from above, brown
from below, surface rough, with sparse mycelium, dry, edge
entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
20180524 (MFLU 18-1514, holotype), ex-type living culture MFLUCC 18-1149; ibid., Chestnut Hill, on submerged
wood in a stream, 10 May 2018, W. Dong, hat454-1 (MFLU
18-1546), living culture MFLUCC 18-1247; ibid., hat454-2
(HKAS 105022), living culture KUMCC 19-0037.
Notes: Ascagilis thailandensis clusters with A. queenslandica, A. submersa and A. sunyatsenii with low bootstrap
support (Fig. 14). The phylogenetically closest species A.
sunyatsenii only has ITS sequence data (323 nucleotides)
in GenBank, which has three nucleotide differences with A.
thailandensis. Due to the limited sequence data, A. sunyatsenii and A. thailandensis appear to represent the same species
in our phylogenetic tree (Fig. 14). However, their morphological characteristics strongly support them to be the different species. Ascagilis sunyatsenii has two types of ascomata
(sessile and stalked), saccate, ovoid to elongate asci and
ellipsoidal ascospores with two conspicuous helmet-shaped
appendages (Inderbitzin et al. 2001). Whereas, A. thailandensis has sessile ascomata, cylindrical asci and irregularly
fusiform ascospores with small, inconspicuous pads. Ascagilis thailandensis can be easily distinguished from the other
two phylogenetically related species A. queenslandica and
A. submersa by ascospore shape, size and appendages (Hyde
et al. 2019).
Ascagilis thailandensis is morphologically similar to
A. guttulaspora in having nearly fusiform, smooth-walled,
guttulate ascospores with tapering apical cell and slightly
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Fig. 11 Ascagilis thailandensis
(MFLU 18-1514, holotype).
a, b Ascomata on host tissue.
c Vertical section of ascoma. d
Structure of peridium. e–h Bitunicate asci. i Pseudoparaphyses.
j–m Ascospores. n Germinated
ascospore. o, p Colony on PDA
(left-front, right-reverse). Scale
bars: c, n = 50 μm, d–m = 20
μm
flattened basal cell. However, A. guttulaspora has smaller
ascospores (29–33 × 8.5–10.5 μm vs. 33–48.5 × 11–16 μm)
and without pads (Hyde et al. 2017). The ascomata of A.
guttulaspora are attached to the substratum by wide, brown
hyphae, while they are lacking in A. thailandensis (Hyde
et al. 2017). Our phylogenetic analysis supports them to be
the different species (Fig. 14).
Key to freshwater Ascagilis species
1. Ascospores without mucilaginous pads………………2
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1. Ascospores with mucilaginous pads at one or both
ends……………………………………………………3
2. Ascospores aseptate…………………A. queenslandica
2. Ascospores 1-septate…………………A. guttulaspora
3. Ascomata sessile………………………………………4
3. Ascomata sessile and stalked…………………………6
4. Ascospores with inconspicuous pads at one or both
ends……………………………………A. thailandensis
4. Ascospores with conspicuous bipolar pads……………5
5. Ascospores 37.5–41 × 17–19 μm…………A. submersa
5. Ascospores 42–52 × 16–23 μm……………A. bipolaris
6. Ascospores monomorphic………………A. sunyatsenii
6. Ascospores dimorphic…………………A. seychellensis
Neojahnula W. Dong, H. Zhang & K.D. Hyde, gen. nov.
Index Fungorum number: IF557823; Facesoffungi number: FoF07688
Etymology: named refers to its morphological similarity
to Jahnula
Saprobic on submerged wood. Sexual morph: Ascomata
scattered or clustered in small groups, superficial, subglobose or obpyriform, brown to black, membranous, with
scarcely projecting papilla, ostiolate, with few setae. Peridium thin, one-layered. Pseudoparaphyses numerous, cellular,
hypha-like, hyaline, sparsely septate. Asci 8-spored, bitunicate, fissitunicate, cylindrical, sessile or short pedicellate,
with an ocular chamber and faint apical ring. Ascospores
uni- to bi-seriate, elongate, ellipsoidal, tapering towards the
apices, brown, 1-septate, upper cell slightly wider than lower
cell. Asexual morph: Undetermined.
Type species: Neojahnula australiensis (K.D. Hyde) W.
Dong, H. Zhang & K.D. Hyde
Notes: Jahnula australiensis K.D. Hyde was collected
from submerged wood in freshwater in Australia (Hyde
1993a). It was initially placed in Jahnula because the
peridium comprises massive pseudoparenchymatous cells,
which is the remarkable feature of Jahnula (Hyde 1993a).
However, J. australiensis was shown to not belong to Jahnula sensu stricto and clustered basal to two asexual genera
Brachiosphaera Nawawi and Speiropsis Tubaki with strong
bootstrap support (Prihatini et al. 2008; Hyde et al. 2017;
Huang et al. 2018; this study, Fig. 14). Jahnula australiensis
is not supported in Jahnula because it has few setae surrounding the ascomata, a thin peridium comprising single
row of cells, asci with an ocular chamber and faint apical
ring and uni- to bi-seriate ascospores, these features are different from Jahnula. As suggested by Hyde et al. (2013), we
transfer J. australiensis to a new genus, Neojahnula based
on phylogenetic analyses and morphology.
351
List of freshwater Neojahnula species
*Neojahnula australiensis (K.D. Hyde) W. Dong, H. Zhang
& K.D. Hyde, comb. nov.
Index Fungorum number: IF557899; Facesoffungi number: FoF09241; Fig. 12
Basionym: Jahnula australiensis K.D. Hyde, Aust. Syst.
Bot. 6(2): 161 (1993)
Freshwater distribution: Australia (Hyde 1993a), Brunei
(Ho et al. 2001), China (Ho et al. 2001), Peru (Shearer et al.
2015), Thailand (Suetrong et al. 2011a; this study) USA
(Raja et al. 2009b)
Saprobic on submerged wood. Sexual morph: Ascomata
80–110 μm high, 100–130 μm diam., scattered or gregarious,
superficial, subglobose or lenticulate when compressed, black,
membranous, with scarcely projecting papilla, ostiolate, with
few hyaline to pale brown hyphae. Peridium thin, 5–10 μm
thick, composed of one row of large, thin-walled, brown to dark
brown cells of textura angularis. Pseudoparaphyses 3 μm diam.,
numerous, cellular, hypha-like, hyaline, sparsely septate. Asci
135–175 × 17–19 μm (x̄ = 155 × 17.5 μm, n = 5), 8-spored,
bitunicate, fissitunicate, cylindrical, sessile or short pedicellate, with an ocular chamber and faint apical ring. Ascospores
23–27.5 × 6–7.5 μm (x̄ = 25.5 × 7 μm, n = 15), uni- to bi-seriate, elongate, ellipsoidal, tapering towards the apices, rounded at
the base, brown, 1-septate, constricted at the septum, upper cell
slightly wider than lower cell, guttulate, mostly curved, smooth,
thin-walled, without sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 20
mm in 25 days at 25 °C, reddish brown from above, dark brown
to black from below, surface rough, with dense mycelium, fluffy,
raised as a annulus around the margin, dry, edge entire.
Material examined: THAILAND, Prachuap Khiri Khan
Province, on submerged wood in a stream, 30 July 2015, K.D.
Hyde, 66D (MFLU 15-2711), living culture MFLUCC 15-0975.
Notes: Our collection MFLUCC 15-0975 clusters with
two strains of Neojahnula australiensis (SS3613 and
SS0665) with high bootstrap support (Fig. 14). SS3613 only
has ITS and SS0665 only has SSU sequence data in GenBank, which are identical with MFLUCC 15-0975. The morphology of SS3613 and SS0665 cannot be compared as they
were not provided in Prihatini et al. (2008) and Pang et al.
(2002), respectively. The sequence data from ex-type strain
of N. australiensis is lacking, but our collection MFLUCC
15-0975 has identical morphological characteristics with the
holotype (BRIP 19208), except for longer asci (135–175 ×
17–19 μm vs. 90–140 × 14–18 μm) (Hyde 1993a). Based on
morphological characteristics and available sequence data,
MFLUCC 15-0975 is identified as N. australiensis.
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Fig. 12 Neojahnula australiensis (MFLU 15-2711). a
Ascomata on host surface. b
Vertical section of ascoma. c
Pseudoparaphyses. d Structure
of peridium. e, f Bitunicate
asci. g Ocular chamber and
faint apical ring. h Ascospore.
i, j Colony on PDA (up-front,
down-reverse). Scale bars: b,
e, f = 20 μm, c = 15 μm, d = 5
μm, g, h = 10 μm
Pseudojahnula W. Dong, H. Zhang & K.D. Hyde, gen. nov.
Index Fungorum number: IF557824; Facesoffungi number: FoF07689
Etymology: named refers to its morphological similarity
to Jahnula
Saprobic on submerged wood. Sexual morph: Ascomata solitary or gregarious, erumpent, become superficial
with base remaining immersed, subglobose, obpyriform or
almost conical, hyaline and then metallic grey, coriaceous,
with ostiolate papilla, with algal associations. Peridium
comprising several layers of relatively large, hyaline, thinwalled, angular cells and covered with sparse hyaline hairs.
Pseudoparaphyses numerous, trabeculate, filamentous,
13
hyaline, septate, unbranched between the asci, branching
and anastomosing above. Asci 8-spored, bitunicate, fissitunicate, obclavate, pedicellate, with an ocular chamber and faint
ring. Ascospores bi- to tri-seriate near the base, overlapping
uniseriate near the apex, l-septate, light brown, ellipsoidfusiform, surrounded by a mucilaginous sheath, which is
wavy in outline. Asexual morph: Undetermined.
Type species: Pseudojahnula potamophila (K.D. Hyde &
S.W. Wong) W. Dong, H. Zhang & K.D. Hyde
Notes: Jahnula potamophila is morphologically closest to
Ascagilis bipolaris in ascospore size, but differs by narrow
mucilaginous sheath which is wavy in outline (Hyde and
Goh 1999b; Hyde and Wong 1999). Both species cluster
Fungal Diversity (2020) 105:319–575
distantly in Jahnulales (Fig. 14). The phylogenetic placement of J. potamophila is unstable, it clustered with Jahnula
sensu stricto clade with low bootstrap support (Huang et al.
2018), but had good affinity in Hyde et al. (2017). With Megalohypha aqua-dulces and our new sequences added in the
order, J. potamophila clustered with M. aqua-dulces with
moderate bootstrap support (Hyde et al. 2019; this study,
Fig. 14).
Jahnula potamophila has hyaline to metallic grey ascomata, a several-layered peridium comprising hyaline cells
and covered with sparse hyaline hairs, obclavate asci with an
ocular chamber and faint ring, and ascospores surrounded by
a wavy mucilaginous sheath (Hyde and Wong 1999). These
characteristics differ from Jahnula, therefore, Pseudojahnula is introduced to accommodate J. potamophila.
List of freshwater Pseudojahnula species
*Pseudojahnula potamophila (K.D. Hyde & S.W. Wong)
W. Dong, H. Zhang & K.D. Hyde, comb. nov.
Index Fungorum number: IF557900; Facesoffungi number: FoF09242; Fig. 6a, e, h
Basionym: Jahnula potamophila K.D. Hyde & S.W.
Wong, Nova Hedwigia 68(3–4): 499 (1999)
Freshwater distribution: Australia (Hyde and Wong 1999),
Costa Rica (Raja and Shearer 2006), USA (Raja et al. 2009b)
Megalohypha A. Ferrer & Shearer, Ferrer, Sivichai &
Shearer, Mycologia 99: 456 (2007)
Saprobic on submerged wood. Sexual morph: Ascomata
superficial, uniloculate, globose to obpyriform, hyaline, membranous, with ostiolate papilla, stalked or sessile. Peridium
composed of large, hyaline, thin-walled cells. Pseudoparaphyses numerous, trabeculate, filamentous, hyaline, septate,
branched, anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, broadly clavate or fusiform, short pedicellate, with or without an ocular chamber. Ascospores irregularly arranged, ellipsoidal, acutely tapered at apices, straight,
brown to dark brown, 1-septate, symmetrical, rough-walled,
longitudinally striate, without appendages or gelatinous
sheath (Ferrer et al. 2007). Asexual morph: Undetermined.
Type species: Megalohypha aqua-dulces A. Ferrer & Shearer
Notes: Megalohypha is monotypic with M. aqua-dulces
occurring on submerged decorticated wood in Panama and
Thailand (Ferrer et al. 2007). Megalohypha can be easily
distinguished from Aliquandostipite and Jahnula based on
its pale to dark brown, 1-septate, rough-walled ascospores
with longitudinally sulcate striations and without appendages or gelatinous sheath (Ferrer et al. 2007). It formed a
moderately supported clade with Pseudojahnula (Hyde et al.
2019) and this study (Fig. 14).
List of freshwater Megalohypha species
353
*Megalohypha aqua-dulces A. Ferrer & Shearer, Mycologia 99(3): 458 (2007); Fig. 5f–j
Freshwater distribution: Panama (Ferrer et al. 2007),
Thailand (Ferrer et al. 2007)
Key to freshwater asexual genera of Aliquandostipitaceae
1. Chlamydospores long, multi-septate……………………
………………………………………………Xylomyces
1. Conidia tetraradiate with 4–8 appendages………………
…………………………………………Brachiosphaera
Brachiosphaera Nawawi, Descals, Nawawi & Webster,
Trans. Br. Mycol. Soc. 67: 213 (1976)
Saprobic on submerged wood, wet wood and river foam.
Sexual morph: Undetermined. Asexual morph: Colonies
effuse, mycelium mostly immersed. Hyphae varying in width,
septate, hyaline at first, turning olivaceous brown with age. Conidiophores macronematous, erect, unbranched, hyaline, septate,
of variable length. Conidiogenous cells holoblastic, sympodial.
Conidia acrogenous, tetraradiate, clustered in chains, ellipsoid
or round-shaped, with 4–10 conidial arms, each 1–4-septate
(Nawawi and Webster 1976; Suetrong et al. 2011a).
Type species: Brachiosphaera tropicalis Nawawi
Notes: Brachiosphaera species are commonly recorded in
freshwater habitats as listed below. The genus is characterized by tetraradiate conidia with a round central cell with
4–10 arms, each of them 1–4-septate (Nawawi and Webster
1976; Suetrong et al. 2011a). The conidia of B. tropicalis
consist of a brown, spherical body and furnished with 4–5
arms, which are longer than 1.5 times diam. of the central
part (Nawawi and Webster 1976; Suetrong et al. 2011a).
Currently, B. tropicalis is phylogenetically related to Jahnulales (Fig. 14). Sequence data for B. jamaicensis is necessary
to shed light on its phylogenetic affinities.
List of freshwater Brachiosphaera species
Brachiosphaera jamaicensis (J.L. Crane & Dumont)
Nawawi, Trans. Br. mycol. Soc. 67(2): 216 (1976)
Basionym: Actinospora jamaicensis J.L. Crane &
Dumont, Can. J. Bot. 53(9): 843 (1975)
Freshwater distribution: Jamaica (Crane and Dumont
1975), China (Chang 1994), Poland (Suetrong et al. 2011a)
*Brachiosphaera tropicalis Nawawi, Trans. Br. mycol. Soc.
67(2): 213 (1976); Fig. 13c–f
Freshwater distribution: China (Chang 1994; Cai et al.
2002a), Malaysia (Nawawi and Webster 1976), Panama
(Campbell et al. 2007), Peru (Shearer et al. 2015), Puerto
Rico (Nieves-Rivera and Santos-Flores 2005), Thailand
(Tubaki et al. 1983; Suetrong et al. 2011a), USA (Raja et al.
2009b), Venezuela (Smits et al. 2007)
13
354
Fungal Diversity (2020) 105:319–575
Fig. 13 Aliquandostipitaceae
spp. (Material examined:
USA, Tennessee, Great Smoky
Mountains National Park, on
submerged decorticated wood,
21 June 2005, H.A. Raja, A.N.
Miller & E.B. Lickey, H58-4;
PANAMA, Colon Province,
Barro Colorado Island, small
river, January 2003, on submerged decorticated wood, A.
Ferrer, E192-1). a, b Conidia
of Xylomyces chlamydosporus
(H58-4). c–f Brachiosphaera
tropicalis (E192-1). c–e Conidia
in PDA. f Broad hyphae in
PDA. Scale bars: a–d = 200
µm, e = 50 µm, f = 100 µm
Key to freshwater Brachiosphaera species
1. Conidia with 4–5 arms……………………B. tropicalis
1. Conidia with 6–10, up to 10–13 arms…………………
…………………………………………B. jamaicensis
Xylomyces Goos, Brooks & Lamore, Mycologia 69(2): 282
(1977)
Saprobic on submerged wood, leaves in freshwater or rotten leaves, root in terrestrial habitats. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate thin,
effuse, reddish brown. Mycelium immersed and superficial,
composed of branched, septate, fuscous hyphae. Stroma absent.
Conidiophores and conidia absent. Chlamydospores abundant,
13
broadly fusiform, brown to blackish, intercalary, solitary or catenate, occasionally branched, straight or curved, uniform in colour or sometimes end cells paler, with thick septate, distinctly
constricted at the septa, thick-walled (Goos et al. 1977).
Type species: Xylomyces chlamydosporus Goos, R.D.
Brooks & Lamore
Notes: The chlamydospores of Xylomyces are often found
on submerged wood in freshwater habitats (Goos et al. 1977;
Goh et al. 1997). The type species X. chlamydosporus was
shown to belong in Aliquandostipitaceae based on molecular
data (Campbell et al. 2007; Huang et al. 2018). Freshwater
species X. aquaticus (Dudka) K.D. Hyde & Goh and X. elegans Goh et al. clustered in Pleosporales, therefore they were
excluded from Xylomyces (Prihatini et al. 2008; Suetrong et al.
Fungal Diversity (2020) 105:319–575
355
Fig. 14 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU and ITS sequence data for
species of Jahnulales. Bootstrap
values for maximum likelihood
equal to or greater than 70% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree
is rooted to Cladosporium
allicinum AFTOL-ID 1591
(Capnodiales)
Ascagilis thailandensis MFLUCC 18-1247 F
Ascagilis thailandensis MFLUCC 18-1149 F
Ascagilis sunyatsenii UBC-F13876
--/1.00
Ascagilis queenslandica MFLU 18-1692
91/-Ascagilis submersa MFLUCC 18-1143 F
99/0.99
Ascagilis bipolaris BCC 3390
Ascagilis guttulaspora MFLUCC 17-0244
Ascagilis seychellensis SS2113.2
100/1.00
Ascagilis seychellensis A492 F
Xylomyces chlamydosporus SS0807
100/1.00
Xylomyces chlamydosporus SS2917
84/0.96
Xylomyces chlamydosporus H58-4 F
Jahnula appendiculata BCC11445
100/1.00
Jahnula appendiculata BCC11400
Jahnula bipileata F49-1 F
82
99/1.00
Jahnula bipileata AF220-1
99/1.00
Jahnula sangamonensis F81-1
Jahnula sangamonensis A482-1B F
100/1.00
Jahnula dianchia KUMCC 17-0039
100/1.00
Jahnula dianchia KUMCC 17-0034
96/1.00 Speiropsis pedatospora SS2229
Speiropsis pedatospora SS2236
97/1.00
Brachiosphaera tropicalis SS2523
Brachiosphaera tropicalis E192-1
100/1.00 96/-- Neojahnula australiensis MFLUCC 15-0975 F
Neojahnula australiensis SS3613
95/0.99
Neojahnula australiensis SS0665
99/1.00 Jahnula rostrata F4-3
--/0.95
Jahnula rostrata MFLU 20-0435 F
93/0.99
Jahnula granulosa SS1567
89/0.98
Jahnula aquatica R68-2
Jahnula aquatica R68-1 F
80/1.00
Ascagilis
Xylomyces
Jahnula
sensu lato
Speiropsis
Brachiosphaera
Neojahnula*
Jahnula
sensu stricto
100/1.00
Megalohypha aqua-dulces AF005-2b F
Megalohypha
Megalohypha aqua-dulces AF005-2a F
Pseudojahnula potamophila F111-1 F
Pseudojahnula*
Aliquandostipite khaoyaiensis KUMCC 19-0083 F
95/1.00
Aliquandostipite khaoyaiensis CBS 118232
96/0.99 Aliquandostipite khaoyaiensis MFLUCC 18-1249 F
Aliquandostipite siamensiae SS81.02
Aliquandostipite
100/1.00
Aliquandostipite crystallinus F83-1 F
93/0.97
76/-Aliquandostipite crystallinus A514-1
Aliquandostipite separans CY2787
100/1.00
Manglicola guatemalensis BCC20157
Manglicola
Manglicola guatemalensis BCC20156
Cladosporium allicinum AFTOL-ID 1591
Outgroup
71/0.99
0.04
2011a; Tanaka et al. 2015). Five species are accepted in the
genus, however, only X. chlamydosporus has been confirmed
with molecular data. The species can be distinguished by chlamydospores size, septation and ornamentation of wall (see key
below). This genus is in need of additional molecular studies to
establish if other described species are phylogenetically related
to X. chlamydosporus within Jahnulales (Fig. 14).
List of freshwater Xylomyces species
Freshwater distribution: Brazil (Oliveira et al. 2015)
*Xylomyces chlamydosporus Goos, R.D. Brooks & Lamore [as ‘chlamydosporis’], Mycologia 69(2): 282 (1977);
Fig. 13a, b
Freshwater distribution: Australia (Hyde and Goh 1997),
Brunei (Goh et al. 1997; Fryar et al. 2004), China (Luo et al.
2004), China (Goh et al. 1997; Tsui et al. 2001c), Seychelles
(Hyde and Goh 1998b), USA (Goos et al. 1977; Lamore and
Goos 1978; Raja et al. 2009b)
Xylomyces acerosisporus M.S. Oliveira, Malosso & R.F.
Castañeda, Mycotaxon 130: (2015)
13
356
Xylomyces giganteus Goh, W.H. Ho, K.D. Hyde & K.M.
Tsui, Mycol. Res. 101(11): 1324 (1997)
Freshwater distribution: Australia (Goh et al. 1997),
China (Jiang et al. 2008), South Africa (Hyde et al. 1998)
Xylomyces punctatus Goh, W.H. Ho, K.D. Hyde & K.M.
Tsui, Mycol. Res. 101(11): 1328 (1997)
Freshwater distribution: China (Goh et al. 1997; Tsui
et al. 2001c)
Xylomyces pusillus Goh, W.H. Ho, K.D. Hyde & K.M. Tsui,
Mycol. Res. 101(11): 1328 (1997)
Freshwater distribution: China (Goh et al. 1997; Tsui
et al. 2001c; Cai et al. 2002a)
Key to freshwater Xylomyces species
1.
1.
2.
2.
3.
Chlamydospores > 25 µm wide………………………2
Chlamydospores < 20 µm wide………………………3
Chlamydospores 5–9-septate………X. chlamydosporis
Chlamydospores 6–26-septate……………X. giganteus
Chlamydospores with distinct punctate wall……………
……………………………………………X. punctatus
3. Chlamydospores lack distinct punctate wall……………4
4. Chlamydospores 42–56 × 7–11 µm…………X. pusillus
4. Chlamydospores 95–180 × 8–10 µm……………………
………………………………………X. acerosisporus
Fungal Diversity (2020) 105:319–575
branched, septate, brown to dark brown, smooth-walled. Conidiogenous cells holoblastic, monoblastic, terminal, constricted at
delimiting septa. Conidia broadly ellipsoid-obovoid, fusiform to
obclavate, rounded at both ends, initially 1-septate, later becoming 2-septate, occasionally 3-septate, constricted and darkly
pigmented at the septa, reddish brown to dark brown, grayish
brown, smooth-walled (Hyde et al. 2013; Su et al. 2016b).
Type species: Kirschsteiniothelia atra (Corda) D.
Hawksw.
Notes: Kirschsteiniothelia is characterized by superficial, subglobose to globose ascomata, cylindric-clavate asci and ellipsoidal, dull green, olive-brown to dark brown ascospores (Boonmee
et al. 2012). The asexual morph of Kirschsteiniothelia has been
connected to Dendryphiopsis, typified by D. atra (Corda) S.
Hughes with molecular data (Boonmee et al. 2012; Hyde et al.
2013; Su et al. 2016b). For convenience, Wijayawardene et al.
(2014) proposed to use Kirschsteiniothelia over Dendryphiopsis
and named K. atra as the type species. All Kirschsteiniothelia species formed a well-supported clade and were placed in
Kirschsteiniotheliaceae by Boonmee et al. (2012). Six freshwater species with molecular data have been accepted in the genus
and all of which are asexual morphs (see list below). All freshwater species have unbranched conidiophores and mostly clavate
conidia (slender conidia in K. fluminicola Z.L. Luo et al.), except
K. aethiops (Sacc.) D. Hawksw. producing branched conidiophores and cylindrical conidia (Su et al. 2016b).
List of freshwater Kirschsteiniothelia species
Kirschsteiniotheliales Hern.-Restr., R.F. Castañeda, Gené
& Crous, Stud. Mycol. 86: 72 (2017)
Kirschsteiniotheliaceae Boonmee & K.D. Hyde, Mycologia 104(3): 705 (2012)
Kirschsteiniothelia D. Hawksw., J. Linn. Soc., Bot. 91: 182
(1985)
Synonymy: Dendryphiopsis S. Hughes, Can. J. Bot. 31:655
(1953)
Saprobic mostly on dead wood in terrestrial or submerged
wood in freshwater habitats. Sexual morph: Ascomata superficial, solitary or clustered, subglobose to globose, membranaceous, dark brown to black, with a central papilla. Peridium
composed of several layers of cells of textura angularis. Pseudoparaphyses numerous, trabeculate, filiform, hyaline. Asci
8-spored, bitunicate, fissitunicate, cylindric-clavate, long pedicellate, apically rounded, with an ocular chamber. Ascospores
biseriate, ellipsoidal, dull green, olive brown to dark brown
at maturity, 1–2-septate, smooth-walled (Hawksworth 1985;
Boonmee et al. 2012). Asexual morph: Hyphomycetous. Conidiophores macronematous, mononematous, erect, gregarious,
elongate and thick-walled, straight and slightly curved, apically
13
*Kirschsteiniothelia aquatica Z.L. Luo, K.D. Hyde & H.Y.
Su, Mycosphere 9(4): 759 (2018)
Freshwater distribution: China (Bao et al. 2018)
*Kirschsteiniothelia atra (Corda) D. Hawksw., Fungal
Diversity 69:37 (2014); Fig. 15a–h
Basionym: Dendryphion atrum Corda, Icon. fung.
(Prague) 4: 33 (1840)
Synonymy: Amphisphaeria aethiops Sacc., Syll. fung.
(Abellini) 1: 722 (1882)
Dendryphiopsis atra (Corda) S. Hughes, Can. J. Bot. 31:
655 (1953)
Sphaeria aethiops Berk. & Curtis, Grevillea 4: 143 (1876)
Kirschsteiniothelia aethiops (Sacc.) D. Hawksw., J. Linn.
Soc., Bot. 91(1 -2): 185 (1985)
Freshwater distribution: China (Su et al. 2016b)
*Kirschsteiniothelia cangshanensis Z.L. Luo, D.F. Bao,
K.D. Hyde & H.Y. Su, Mycosphere 9(4): 760 (2018)
Freshwater distribution: China (Bao et al. 2018)
*Kirschsteiniothelia fluminicola Z.L. Luo, K.D. Hyde &
H.Y. Su, Mycosphere 9(4): 760 (2018)
Freshwater distribution: China (Bao et al. 2018)
Fungal Diversity (2020) 105:319–575
357
Fig. 15 Kirschsteiniothelia spp. (Material examined: CHINA, Yunnan
Province, Dali, Cangshan Mountain, Huadianba, saprobic on decaying
wood submerged in stream, May 2014, Y. Su, S-144, HKAS 84022;
ibid., on decaying wood submerged in Wanhua Stream, March 2014,
X.Y. Liu, S-043, HKAS 93066, holotype). a–h Kirschsteiniothelia
atra (HKAS 84022). a, b Colonies on the substratum. c Conidiophores
with conidia. d, e Conidiogenous cells and conidia. f–h Conidia. i–o
Kirchsteiniothelia submersa (HKAS 93066). i, j Conidiophores with
conidia. k Conidiogenous cells with conidia. l–o Conidia. Scale bars: c
= 130 μm, d, e = 50 μm, f–h, l–o = 20 μm, i, j = 100 μm, k = 30 μm
*Kirschsteiniothelia rostrata J Yang & K.D. Hyde, Fungal
Diversity 87: 45 (2017)
Freshwater distribution: China (Bao et al. 2018), Thailand (Hyde et al. 2017)
*Kirschsteiniothelia submersa H.Y. Su & K.D. Hyde, Fungal Diversity (2016); Fig. 15i–o
Freshwater distribution: China (Su et al. 2016b)
13
358
Key to freshwater Kirschsteiniothelia species
1. Conidiophores branched…………………………K. atra
1. Conidiophores unbranched……………………………2
2. Conidia slender, solitary to short-catenate………………
…………………………………………K. fluminicola
2. Conidia not slender, solitary……………………………3
3. Conidia with sheath……………………………………4
3. Conidia without sheath…………………………………5
4. Conidia 33–43 μm long………………K. cangshanensis
4. Conidia 80–150 μm long……………………K. rostrata
5. Conidiophores 114–151 × 7–8 μm………K. aquatica
5. Conidiophores 220–280 × 6–7 μm………K. submersa
Minutisphaerales Raja, Oberlies, Shearer & A.N. Mill.,
Mycologia 107(4): 854 (2015)
Acrogenosporaceae Jayasiri & K.D. Hyde, Mycosphere
9(4): 809 (2018)
Acrogenospora M.B. Ellis, Dematiaceous Hyphomycetes
(Kew): 114 (1971)
Saprobic on decaying wood, bark, culms of bamboo in
freshwater or terrestrial habitats. Sexual morph: Hysterothecia laterally compressed, with a prominent sunken slit, solitary
to gregarious, erect and elevated, presenting an almost stipitate appearance, thick-walled. Pseudoparaphyses branched.
Asci 8-spored, cylindrical, fusoid or obovate, obtuse at the
apex, short pedicellate, thin-walled. Ascospores oval, aseptate,
hyaline or moderately pigmented (Mason 1941; Jayasiri et al.
2018). Asexual morph: Hyphomycetous. Colonies effuse, dark
brown to black, glistening, hairy. Mycelium mostly immersed,
consist of septate, thin-walled, smooth, hyaline to pale brown
hyphae. Conidiophores macronematous, mononematous, solitary, erect, subulate or cylindrical, straight or slightly flexuous,
pale brown to dark brown, septate, unbranched, smooth. Conidiogenous cells holoblastic, monoblastic, integrated, terminal or
intercalary, with percurrent proliferations, cylindrical. Conidia
solitary, dry, acrogenous, simple, spherical, subspherical, olive
to brown, aseptate, truncate at the base, smooth or verrucose
(Ellis 1971; Hyde et al. 2019).
Type species: Acrogenospora sphaerocephala (Berk. &
Broome) M.B. Ellis
Notes: Acrogenospora is a holomorphic genus, characterized by laterally compressed hysterothecia with a prominent
sunken slit, oval, aseptate, hyaline ascospores and spherical,
olive to brown conidia (Hyde et al. 2019). Acrogenospora is
used to suppress Farlowiella Sacc. (Rossman et al. 2015) and
their sexual-asexual connections were linked based on molecular data (Jayasiri et al. 2018). Thus, Acrogenospora is the only
genus in Acrogenosporaceae (Jayasiri et al. 2018; Hyde et al.
2019). The sequence data of the type species A. sphaerocephala
were generated based on a freshwater collection from Thailand
13
Fungal Diversity (2020) 105:319–575
(Hyde et al. 2019). Bao et al. (2020) introduced seven new species collected from a small area of Yunnan Province, China,
which indicated Acrogenospora is a speciose genus. 20 species
were reported in Acrogenospora and 13 have been confirmed
with molecular data (Bao et al. 2020). Freshwater Acrogenospora species are morphologically very similar and the sequence
data are main evidence to separate them.
List of freshwater Acrogenospora species
*Acrogenospora aquatica D.F. Bao, Z.L. Luo, K.D. Hyde &
H.Y. Su, Frontiers in Microbiology 11: 3 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
*Acrogenospora basalicellularispora D.F. Bao, Z.L. Luo,
K.D. Hyde & H.Y. Su, Frontiers in Microbiology 11: 8 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
*Acrogenospora ellipsoidea D.M. Hu, L. Cai & K.D. Hyde,
Sydowia 62(2): 194 (2010)
Freshwater distribution: Yunnan, China (Hu et al. 2010a)
*Acrogenospora guttulatispora D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Frontiers in Microbiology 11: 8 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
*Acrogenospora obovoidspora D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Frontiers in Microbiology 11: 8 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
*Acrogenospora olivaceospora D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Frontiers in Microbiology 11: 10 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
Acrogenospora ovalis Goh, K.D. Hyde & C.K.M. Tsui [as
‘ovalia’], Mycol. Res. 102(11): 1312 (1998)
Freshwater distribution: Hong Kong (China) (Goh et al.
1998b)
*Acrogenospora sphaerocephala (Berk. & Broome) M.B.
Ellis, Dematiaceous Hyphomycetes (Kew): 114 (1971)
Basionym: Monotospora sphaerocephala Berk. &
Broome, Ann. Mag. nat. Hist., Ser. 3 3: 361 (1859)
Synonymy: Halysium sphaerocephalum (Berk. &
Broome) Vuill., Bull. Séanc. Soc. Sci. Nancy, Sér. 3 11:
167 (1911)
Monosporella sphaerocephala (Berk. & Broome) S.
Hughes, Can. J. Bot. 31: 654 (1953)
Monotosporella sphaerocephala (Berk. & Broome) S.
Hughes, Can. J. Bot. 36: 787 (1958)
Freshwater distribution: China (Goh and Hyde 1999; Tsui
et al. 2000), Philippines (Cai et al. 2003a), Seychelles (Hyde
and Goh 1998b), Thailand (Sivichai et al. 2000; Hyde et al.
2019), USA (Lamore and Goos 1978; Shearer and Crane 1986)
Fungal Diversity (2020) 105:319–575
*Acrogenospora submersa D.F. Bao, Z.L. Luo, K.D. Hyde
& H.Y. Su, Frontiers in Microbiology 11: 12 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
*Acrogenospora subprolata Goh, K.D. Hyde & C.K.M.
Tsui, Mycol. Res. 102(11): 1314 (1998)
Freshwater distribution: Australia (Goh et al. 1998b),
China (Goh et al. 1998b; Tsui et al. 2001c; Bao et al. 2020),
Seychelles (Goh et al. 1998b), South Africa (Goh et al.
1998b), UK (Goh et al. 1998b)
*Acrogenospora thailandica J. Yang & K.D. Hyde, Fungal
Diversity 96: 78 (2019)
Freshwater distribution: Thailand (Hyde et al. 2019)
*Acrogenospora verrucispora Hong Zhu, L. Cai & K.Q.
Zhang [as ‘verrucospora’], Mycotaxon 92: 384 (2005)
Freshwater distribution: Yunnan, China (Zhu et al. 2005;
Bao et al. 2020)
*Acrogenospora yunnanensis D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Frontiers in Microbiology 11: 18 (2020)
Freshwater distribution: Yunnan, China (Bao et al. 2020)
Key to freshwater Acrogenospora species
1.
Conidiophores 850–950 × 3.5–8 µm…………………
…………………………………………A. thailandica
1. Conidiophores < 800 µm long………………………2
2. Conidiophores have wide range of length, 100–730 ×
7.2–10.5 µm………………………A. sphaerocephala
2. Conidiophores not as above…………………………3
3. Conidia with a small, hyaline basal cell………………4
3. Conidia without basal cell……………………………5
4. Conidiophores 202–250 × 7.5–9.5 µm………………
……………………………………………A. aquatica
4. Conidiophores 259–395 × 8–12 µm…………………
………………………………A. basalicellularispora
5. Conidia hyaline when young, darker when
mature………………………………………………6
5. Conidia colour almost consistent……………………8
6. Conidiophores 163–223 × 6.7–10 µm…………………
……………………………………………A. submersa
6. Conidiophores > 260 µm long………………………7
7. Conidiophores 7.5–8.6 µm wide………………………
………………………………………A. guttulatispora
7. Conidiophores 8.6–12 µm wide………A. yunnanensis
8. Conidia verrucose……………………A. verrucispora
8. Conidia smooth………………………………………9
9. Conidia ellipsoidal………………………A. ellipsoidea
9. Conidia not as above…………………………………10
10. Conidia 24–33 × 18–22 µm…………………A. ovalia
10. Conidia > 32 µm long………………………………11
359
11. Conidia 39–46 × 30–39 µm……………A. subprolata
11. Conidia < 39 µm long………………………………12
12. Conidiophores 209–277 × 7.5–10 µm………………
………………………………………A. obovoidspora
12. Conidiophores 102–172 × 5.8–9 µm…………………
………………………………………A. olivaceospora
Minutisphaeraceae Raja, Oberlies, Shearer & A.N. Mill.,
Mycologia 107(4): 854 (2015)
Minutisphaera Shearer, A.N. Mill. & A. Ferrer, Mycologia
103(2): 415 (2011)
Saprobic on submerged wood. Sexual morph: Ascomata
scattered or clustered in groups, erumpent to superficial, uniloculate, globose to subglobose, brown, with central, rounded
ostiole surrounded by black hairs. Peridium thin, composed
of textura angularis to globulosa cells. Pseudoparaphyses
sparse in young ascomata, becoming abundant with age,
cellular, hyaline, septate, simple or branched, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
ovoid to obclavate, lacking a pedicel, rounded at the apex.
Ascospores irregularly arranged, clavate, mostly straight,
hyaline to pale brown, 1(–3)-septate, constricted at the septum, multiguttulate, asymmetrical, smooth- or rough-walled
when mature, with sheath and filamentous appendages (Ferrer et al. 2011). Asexual morph: Undetermined.
Type species: Minutisphaera fimbriatispora Shearer, A.N.
Mill. & A. Ferrer
Notes: All species reported within Minutisphaera were collected from freshwater habitats (Ferrer et al. 2011; Raja et al.
2013b, 2015; Bao et al. 2019a). The ascospores of Minutisphaera are unusual in being asymmetrical and often equipped
with a sheath and filamentous appendages (Ferrer et al. 2011;
Raja et al. 2015). The shape and size of ascospores, and features of appendages and sheaths are used for distinguishing
all species within the genus (see key below).
In the analysis of Raja et al. (2015) BLAST search of
ITS sequences revealed that two isolates, Pleosporales
sp. 39g (JX244063) and Didymosphaeria sp. TS_04_050
(HQ713763), shared 99% similarity with Minutisphaera
aspera. Both strains were isolated as fungal endophytes in
previous studies (Grünig et al. 2011). Thus, it was hypothesized that members of Minutisphaerales may lead a dual life
as saprobes on submerged wood and as fungal endophytes of
trees living in close proximity to rivers and streams. However, additional studies are needed to shed light on the dual
ecological life style of Minutisphaera spp.
List of freshwater Minutisphaera species
*Minutisphaera aquaticum D.F. Bao, L. Xu & H.Y. Su,
Asian Journal of Mycology 2(1): 309 (2019)
Freshwater distribution: Thailand (Bao et al. 2019a)
13
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Fungal Diversity (2020) 105:319–575
Fig. 16 Minutisphaera spp. (Material examined: USA, North Carolina:
Rockingham County, Piedmont Plateau, Big Beaver Island Creek, Madison, on submerged decorticated wood, 26 April 2013, H.A. Raja & N.H.
Oberlies, G427-1, ILLS 72340, holotype; ibid., G156-4, ILLS 72342,
paratype). a–c M. aspera (G427-1). a Ascoma on wood. b Ascus. c
Ascospores. d–f M. parafimbriatispora (G156-4). d Ascoma in culture.
e Asci. f Ascospore with gelatinous sheath constricted at the mid-septum. Scale Bars: a = 100 µm, b, c = 20 µm, d = 50 µm, e, f = 10 µm
*Minutisphaera aspera Raja, Oberlies, Shearer & A.N.
Mill, Mycologia 107(4): 854 (2015); Fig. 16a–c
Freshwater distribution: USA (Raja et al. 2015)
1.
2.
2.
3.
*Minutisphaera fimbriatispora Shearer, A.N. Mill. & A.
Ferrer, Mycologia 103(2): 415 (2011)
Freshwater distribution: USA (Ferrer et al. 2011; Raja
et al. 2013b)
*Minutisphaera japonica Kaz. Tanaka, Raja & Shearer,
Mycologia 105(4): 966, (2013); Fig. 17
Freshwater distribution: Japan (Raja et al. 2013b), France
(Raja et al. 2015)
*Minutisphaera parafimbriatispora Raja, Oberlies, Shearer
& A.N. Mill, Mycologia 107(4): 855 (2015); Fig. 16d–f
Freshwater distribution: USA (Raja et al. 2015)
Key to freshwater Minutisphaera species
1. Ascospores with sheath, but lacking appendages………2
13
Ascospores with sheath and appendages………………4
Ascospores with rough-walled………………M. aspera
Ascospores smoth-walled………………………………3
Ascospores broadly fusiform, with acute apex…………
……………………………………………M. japonica
3. Ascospores fusiform to clavate, with rounded apex……
…………………………………………M. aquaticum
4. Ascospores 24–36 × 6–8 µm………M. fimbriatispora
4. Ascospores 18–25 × 4–7 µm……………………………
…………………………………M. parafimbriatispora
Natipusillales Raja, Shearer, A.N. Mill. & K.D. Hyde, Fungal Diversity 63: 9 (2013)
Natipusillaceae Raja, Shearer & A.N. Mill., Mycologia
104(2): 570 (2012)
Natipusilla A. Ferrer, A.N. Mill. & Shearer, Mycologia
103(2): 417 (2011)
Saprobic on submerged wood. Sexual morph: Ascomata
scattered or aggregated, erumpent to superficial, uniloculate,
Fungal Diversity (2020) 105:319–575
361
Fig. 17 Minutisphaera japonica (Material examined: FRANCE,
Ariège, Vernajoul, ruisseaude Vernajoul, on submerged wood of
Fraxinus excelsior, 2 July 2007, J. Fournier, JF07132, associated with
Pseudohalonectria lutea and Aniptodera sp., leg det JF). a Ascomata
on the wood surface. b Squash mount of ascoma, note dark-walled
hyphae on the peridial wall. c Longitudinal section of apothecioid
ascoma. d–f Asci. g, h Ascospores. Scale bars: a = 200 µm, b, d, f, h
= 20 µm, c = 50 µm, e = 30 µm, g = 10 µm
globose to subglobose, hyaline to brown, lacking ostioles,
occasionally with minute papilla. Peridium thin, membranous, glabrous, forming a textura angularis in surface view.
Pseudoparaphyses sparse or absent. Asci 8-spored, bitunicate, fissitunicate, globose to obclavate, lacking a pedicel,
with or without an ocular chamber. Ascospores overlapping
tri- to multi-seriate, arranged in a group, fusiform or clavate,
straight, the upper cell often shorter and broader than the
lower one, mostly 1-septate, hyaline, occasionally becoming
3-septate, pale brown at age, guttulate when young, smooth,
thin-walled, with or without a gelatinous sheath (Ferrer et al.
2011). Asexual morph: Undetermined.
Type species: Natipusilla decorospora A. Ferrer, A.N.
Mill. & Shearer
Notes: All species described in this genus were reported
from freshwater habitats in tropical and subtropical regions
(Ferrer et al. 2011; Raja et al. 2012). Phylogenetic analysis
using combined SSU and LSU sequence dataset placed this
genus in a distinct clade within Dothideomycetes. Therefore,
a novel family, Natipusillaceae (Raja et al. 2012), and new
order Natipusillales (Hyde et al. 2013) were established for
Natipusilla.
Ascominuta Ranghoo & K.D. Hyde was thought to be
an earlier name of Natipusilla (Hyde et al. 2013) due to the
bitunicate, globose asci, sheathed ascospores and freshwater
habitat (Ranghoo and Hyde 1999; Hu et al. 2010a; Ferrer
et al. 2011; Raja et al. 2012). However, Ascominuta differs
from Natipusilla in the numbers of ascospores per asci (4 vs.
8) and colour of ascomata (hyaline to brown vs. dark brown to
black). Only LSU sequence (AF132335) of A. lignicola Ranghoo & K.D. Hyde (generic type) was deposited in Genbank,
and it cannot be aligned with Natipusilla. Further molecular
and morphological data are needed to clarify these two genera
(Raja et al. 2012). All Natipusilla species are surrounded by
a complex gelatinous sheath, except N. naponensis A. Ferrer
et al. lacks a sheath (Ferrer et al. 2011; Raja et al. 2012).
List of freshwater Natipusilla species
*Natipusilla decorospora A. Ferrer et al., Mycologia
103(2): 417 (2011)
Freshwater distribution: Ecuador (Ferrer et al. 2011)
*Natipusilla limonensis A. Ferrer, A.N. Mill. & Shearer,
Mycologia 103(2): 417 (2011); Fig. 18
Freshwater distribution: Costa Rica (Ferrer et al. 2011),
Peru (Shearer et al. 2015)
*Natipusilla naponensis A. Ferrer, A.N. Mill. & Shearer,
Mycologia 103(2): 420 (2011)
Freshwater distribution: Ecuador (Ferrer et al. 2011)
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Fungal Diversity (2020) 105:319–575
Fig. 18 Natipusilla limonensis (Material examined: PERU, Cusco,
Camanti, Stream at Quince Mil Trail 1, on submerged wood, 26 May
2010, S. Zelski & H.A. Raja, PE0003). a Section of ascoma. b Glo-
bose to subglobose asci. c–f Ascospores. Arrows indicating complex
gelatinous sheath. Scale bars: a = 50 µm, b–f = 20 µm
*Natipusilla bellaspora Raja, Shearer & A.N. Mill., Mycologia 104(2): 570 (2012)
Freshwater distribution: Peru (Raja et al. 2012; Shearer
et al. 2015)
Acrocalymma Alcorn & J.A.G. Irwin, Trans. Br. mycol. Soc.
88(2): 163 (1987)
Saprobic on submerged wood, leaf litter or other plants.
Sexual morph: (from culture) Ascomata scattered, erumpent
becoming superficial, uniloculate, globose, covered with
light gray, gnarled, warted hairs, black, coriaceous, with
ostiolate papilla. Neck central, terete, composed of 2–5 layers of brown, polygonal cells around ostiole with hyaline
periphyses. Peridium thin, composed of polygonal, brown,
pseudoparenchyma cells. Pseudoparaphyses numerous, septate, anastomosing. Asci bitunicate, 8-spored, cylindrical,
short pedicellate, apically rounded, with an distinct ocular
chamber. Ascospores uni- to bi-seriate, narrowly fusiform,
straight or slightly curved, hyaline first, pale reddish brown
with lighter coloured end cells, 1–3-septate, constricted at
first submedian septum, basically symmetrical, smooth to
rough-walled, thin-walled, with 4 guttules, with a thin sheath
(Shoemaker et al. 1991). Asexual morph: Coelomycetous.
Pycnidia scattered or aggregated, semi-immersed to superficial, unilocular, globose, brown to black, with ostiolate
papilla or rostrate. Conidiophores reduced. Conidiogenous
cells holoblastic, determinate, discrete, cylindrical, lageniform to ampulliform, hyaline, smooth. Conidia cylindrical
Key to freshwater Natipusilla species
1.
1.
2.
2.
3.
Ascospores without sheath………………N. naponensis
Ascospores with complex gelatinous sheaths…………2
Ascospores 40–48 × 10–13 µm…………N. bellaspora
Ascospores < 40 µm long………………………………3
Ascospores 30–37 × 6–9 µm, with a single scythe
shaped appendage at the apex……………N. decrospora
3. Ascospores 35–40 × 9–11 µm, without appendage…
……………………………………………N. limonensis
Pleosporales Luttrell ex M.E. Barr
Acrocalymmaceae Crous & Trakun., IMA Fungus 5(2):
404 (2014)
13
Fungal Diversity (2020) 105:319–575
to fusoid with subobtuse apex, straight, hyaline at first,
becoming pigmented with age, 0–3-septate, not constricted
at septa, smooth, thin-walled, with or without helmet-shaped
appendages (Zhang et al. 2012a).
Type species: Acrocalymma medicaginis Alcorn & J.A.G.
Irwin
Notes: Acrocalymma is well-studied because all species
have sequences in GenBank. Acrocalymma species are easy
to produce pycnidia in culture (Trakunyingcharoen et al.
2014). The sexual morph was linked to Massarina walkeri
Shoemaker et al. which formed in the pure culture of Acrocalymma medicaginis (Shoemaker et al. 1991). However,
these two species are phylogenetically distinct (Trakunyingcharoen et al. 2014). Therefore, Trakunyingcharoen et al.
(2014) transferred M. walkeri to Acrocalymma and treated
A. medicaginis and A. walkeri as distinct species. The type
material of A. medicaginis was re-examined and described
by Li et al. (2020). Acrocalymma aquatica is the first species
reported from freshwater habitats, and it is characterized
by cylindrical to fusoid, 0(–1)-septate conidia with an apical, mucilaginous, helmet-shaped appendage (Zhang et al.
2012a). In this study, we report a new freshwater species A.
bipolare, which is collected from Egypt.
List of freshwater Acrocalymma species
*Acrocalymma aquatica H. Zhang & K.D. Hyde, Cryptog.
Mycol. 33(3): 337 (2012); Fig. 19
Freshwater distribution: Thailand (Zhang et al. 2012a)
*Acrocalymma bipolare Abdel-Aziz & Abdel-Wahab, sp.
nov.
Index Fungorum number: IF557901; Facesoffungi number: FoF09243; Fig. 20
Etymology: in reference to the polar mucoid appendages
of the conidia
Holotype: CBS H-22673
Saprobic on submerged wood in freshwater habitats.
Sexual morph: Undetermined. Asexual morph: Pycnidia
175–230 µm diam., globose or subglobose, erumpent to
363
superficial, solitary, dark brown to black, without ostiolate
papilla. Peridium 8–17 µm thick, composed of 8–11 layers
of dark brown to black, thick-walled, cells of textura angularis, inwardly with narrow lumens, hyaline and thin-walled
cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 2.5–7 × 2–3.5 μm, holoblastic, determinate,
discrete, cylindrical or lageniform, hyaline, formed from the
inner cells of the pycnidial wall. Conidia 9–12 × 3–5 µm
( x̄ = 10.8 × 4.3, n = 50), cylindrical to fusiform, straight,
unicellular, hyaline, thin-walled, guttulate, with rounded
apex and slightly narrow, truncate base, with mucoid polar
appendages that filled with oil droplets, appendages elongate in water to form filaments. Apical appendages 4–6 μm
diam., rounded. Lower appendages 1–2 μm in length, sometimes split in the middle.
Culture characteristics: Colonies on PDA up to 22 mm
diam. after 2 weeks at 23 °C, with grey to dark brown aerial
and immersed mycelium; reverse dark–brown.
Material examined: EGYPT, Sohag City, River Nile,
26°33′32ʺN, 31°41′44ʺE, on decaying wood submerged in
River Nile water, 14 August 2012, F.A. Abdel-Aziz, (CBS
H-22673, holotype), ex-type living culture MD1321.
Notes: Acrocalymma bipolare clusters with the type species A. medicaginis and A. pterocarpi Jayasiri et al. with
high bootstrap support (Fig. 2). Morphologically, Acrocalymma bipolare is different from A. medicaginis by its
appendages that are filled with oil droplets and become filaments when mounted in water, while A. medicaginis processes globose to hemispherical or helmet-shaped appendages which do not form filaments in water (Alcorn and Irwin
1987). In addition, A. bipolare has shorter conidia than A.
medicaginis (9–12 × 3–5 µm vs. 11–21 × 3.5–5 µm). The
phylogenetically close species A. pterocarpi did not produce
asexual morph in culture, thus their morphology can not be
compared (Jayasiri et al. 2019).
Acrocalymma bipolare only has LSU sequence data, and
it has four and six nucleotide differences with A. medicaginis
and A. pterocarpi, respectively, which indicates them to be
Fig. 19 Acrocalymma aquatica
(Material examined: THAILAND, Chiang Mai Province,
Doi Inthanon, on submerged
wood, 16 November 2010, H.
Zhang, d67, MFLU 11-1113,
holotype). a Pycnidia on substrate surface. b–d Conidia with
single apical appendage. Scale
bars: b–d = 5 μm
13
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Fungal Diversity (2020) 105:319–575
Fig. 20 Acrocalymma bipolare
(CBS H-22673, holotype). a
Vertical section of a pycnidium.
b Section through the peridium.
c–f Variously shaped conidia
with mucoid polar appendages.
Note the apical appendages
formed long filament (arrowed
in d, stained in toluidine blue).
Scale bars: a = 50 μm, b, c =
10 μm, d–f = 5 μm
different species. This is also supported by our phylogenetic
analysis (Fig. 2).
Acrocalymma bipolare differs from the other freshwater
species A. aquatica in having shorter conidia (9–12 × 3–5
μm vs. 12–17 × 3–4 μm) and bipolar appendages compared
with a single appendage in A. aquatica (Zhang et al. 2012a).
Key to freshwater Acrocalymma species
1. Conidia 12–17 × 3–4 μm, with a single, helmet-shaped
appendage…………………………………A. aquatica
1. Conidia 9–12 × 3–5 μm, with bipolar appendages filled
with oil droplets, appendages elongate in water to form
filaments……………………………………A. bipolare
Aigialaceae Suetrong, Sakay., E.B.G. Jones, Kohlm.,
Volkm.-Kohlm. & C.L. Schoch, Stud. Mycol. 64: 166 (2009)
Key to freshwater genera of Aigialaceae
1. Asci cylindro-clavate, ascospores often hyaline,
smooth………………………………………Fissuroma
13
1. Asci obclavate, ascospores often brown, verrucose………………………………Neoastrosphaeriella
Fissuroma J.K. Liu, Phook., E.B.G. Jones & K.D. Hyde,
Fungal Diversity 51(1): 145 (2011)
Saprobic on submerged wood, dead stem of bamboo or
other plant materials. Sexual morph: Ascomata scattered,
rarely clustered, solitary to gregarious, immersed beneath
host epidermis, becoming raised, uniloculate, rarely biloculate joined at the base, hemispherical domes, glabrous, black,
coriaceous or carbonaceous, opening with a slit-like ostiole.
Peridium thick, uneven in thickness, poorly developed at
the base, thick at sides towards the apex, composed of dark
brown pseudoparenchymatous cells, with host cells, plus
fungal tissue, arranged in a textura angularis to textura epidermoidea, carbonaceous at slit-like opening. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, embedded
in a gelatinous matrix and anastomosing between and above
the asci. Asci 8-spored, bitunicate, fissitunicate, obclavate to
cylindrical, pedicellate, apically rounded, with a small ocular chamber. Ascospores overlapping uni- to tri-seriate, fusiform, straight to slightly curved, hyaline, 1-septate, slightly
constricted at the septum, smooth, thin-walled, surrounded
Fungal Diversity (2020) 105:319–575
by a mucilaginous sheath (Liu et al. 2011). Asexual morph:
Coelomycetous, reported as pleurophomopsis-like from culture of Fissuroma aggregata (KT 767). Conidiomata globose to subglobose. Conidiophores cylindrical, simple to
branched, septate. Conidiogenous cells phialidic. Conidia
globose, hyaline (Tanaka and Harada 2005).
Type species: Fissuroma maculans (Rehm) J.K. Liu,
E.B.G. Jones & K.D. Hyde
Notes: Fissuroma comprises 12 species in Index Fungorum (2020). Eight species have been confirmed with
molecular data (Liu et al. 2011; Phookamsak et al. 2015b;
Tennakoon et al. 2018; Wanasinghe et al. 2018a). However,
Phookamsak et al. (2015b) indicated that F. aggregata (I.
Hino & Katum.) Phook. et al. is a species complex which
might comprise at least three distinct species based on their
phylogenetic analyses. After re-examination of the holotype
specimen of F. aggregata (≡ Melanopsamma aggregata),
Phookamsak et al. (2015b) considered the collection MFLU
11-0146 of Liu et al. (2011) as a distinct species from F.
aggregata. Therefore, F. neoaggregatum Phook. & K.D.
Hyde was introduced based on the specimen MFLU 11-0146
(Phookamsak et al. 2015b). The asexual morph was developed in the culture of F. aggregata (KT 767) (Tanaka and
Harada 2005). However, the real identity of KT 767 needs
confirmation to separate F. aggregata KT 984, as shown in
the phylogenetic tree (Fig. 2). We provide a description and
illustration for F. neoaggregatum collected from submerged
wood in Thailand.
List of freshwater Fissuroma species
*Fissuroma neoaggregatum Phook. & K.D. Hyde [as ‘neoaggregata’], Fungal Diversity 74: 158 (2015); Fig. 21
Freshwater distribution: Thailand (this study)
Saprobic on submerged wood. Sexual morph: Ascostromata dark brown to black, solitary to gregarious, immersed
in host epidermis, visible as raised, dome-shaped areas
on the host surface, hemisphaerical to depressed conical,
with flattened base, uniloculate, coriaceous, ostiolate, with
slit-like opening. Peridium of unequal thickness, poorly
developed at the base. Pseudoparaphyses dense, trabeculate, filiform, embedded in a hyaline gelatinous matrix. Asci
140–155 × 16–21 μm ( x̄ = 149 × 19 μm, n = 5), 8-spored,
bitunicate, obclavate, with furcate to truncate pedicel, apically rounded, with an ocular chamber. Ascospores 36–43 ×
6–8 μm (x̄ = 40.5 × 7 μm, n = 10), overlapping biseriate and
uniseriate at the apex, hyaline, fusiform, with acute ends,
1-septate, constricted at the septum, smooth-walled, guttulate, surrounded by a gelatinous sheath. Asexual morph:
Undetermined.
Material examined: THAILAND, Chiang Rai Province, on submerged wood, 18 January 2010, H. Zhang, a45
(MFLU 11-1140), living culture MFLUCC 10-0917.
365
Notes: Our new collection MFLUCC 10-0917 is identified as Fissuroma neoaggregatum based on nearly identical
ITS sequence data with ex-type strain MFLUCC 10-0554
(two nucleotide differences) and morphological characters.
Our collection has shorter asci (140–155 × 16–21 μm vs.
155–197 × 15–18) μm) and ascospores (36–43 × 6–8 μm
vs. 39–54 × 7–9 μm) than the holotype (Phookamsak et al.
2015b). Moreover, our collection shows a broad, mucilaginous sheath in Indian Ink, while the holotype has only a
thin sheath (Phookamsak et al. 2015b). Other sequences are
expected for MFLUCC 10-0917 to further clarify this species. This is a new habitat record for F. neoaggregatum from
freshwater.
Neoastrosphaeriella J.K. Liu, E.B.G. Jones & K.D. Hyde,
Fungal Diversity 51(1): 148 (2011)
Saprobic on decaying twigs, petiole and submerged wood
in terrestrial and freshwater habitats. Sexual morph: Ascomata scattered, immersed to semi-immersed, with a slit-like
ostiole, black. Peridium uneven in thickness, carbonaceous,
composed of dark brown, thick-walled cells. Pseudoparaphyses trabeculate, hyaline, anastomosing, embedded in a
gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
obclavate to cylindrical, pedicellate, with a small ocular chamber. Ascospores fusiform, hyaline to dark brown,
mostly verrucose when mature, 1-septate, surrounded by
a mucilaginous sheath (Liu et al. 2011). Asexual morph:
Undetermined.
Type species: Neoastrosphaeriella krabiensis J.K. Liu,
E.B.G. Jones & K.D. Hyde
Notes: Neoastrosphaeriella was established by Liu
et al. (2011) to accommodate species having ascomata
with slit-like ostioles, obclavate asci and brown, verrucose ascospores. Neoastrosphaeriella aquatica D.F. Bao
et al. was the only species found in freshwater habitats and
differed from the others by hyaline to greyish brown, verrucose ascospores and distinct molecular characters (Bao
et al. 2019b). Neoastrosphaeriella nested well in Aigialaceae
(Liu et al. 2011; Wanasinghe et al. 2018a; Bao et al. 2019b;
Zhang et al. 2020; this study, Fig. 2).
List of freshwater Neoastrosphaeriella species
*Neoastrosphaeriella aquatica D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Phytotaxa 391(3): 201 (2019)
Freshwater distribution: Thailand (Bao et al. 2019b)
Amniculicolaceae Y. Zhang ter, C.L. Schoch, J. Fourn.,
Crous & K.D. Hyde, Stud. Mycol. 64: 95 (2009)
Key to freshwater genera of Amniculicolaceae
1. Asexual morph……………………………Vargamyces
1. Sexual morph…………………………………………2
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Fig. 21 Fissuroma neoaggregatum (MFLU 11-1140, new
habitat record). a Appearance
of ascomata on wood. b Asci
and pseudoparaphyses. c–e
Asci. Note the bitunicate ascus
in e. f Pseudoparaphyses. g–i
Ascospores. (i in Indian Ink). j
Front view of colony on PDA.
Scale bars: b–f = 50 μm, g–i =
20 μm
2. Ascospores hyaline, 1-septate……………Amniculicola
2. Ascospores pigmented, multi-septate…………………3
3. Ascospores golden yellow to brown, dictyoseptate……
………………………………………………Murispora
3. Ascospores reddish brown, phragmoseptate………
…………………………………Neomassariosphaeria
Amniculicola Y. Zhang ter & K.D. Hyde, Mycological
Research 112 (10): 1189 (2008)
Saprobic on submerged wood or in plant humus. Sexual
morph: Ascomata solitary to gregarious, immersed to nearly
13
superficial, black, uniloculate, subglobose to conical, glabrous, ostiolate, with or without two tuberculate flared lips
surrounding a slit-like ostiole, sometimes with a flattened
base not easily removed from the substrate, usually staining
the woody substrate purple. Peridium 2-layered, outer layer
composed of heavily pigmented thick-walled cells of textura
angularis, inner layer composed of hyaline thin-walled cells
of textura angularis. Pseudoparaphyses dense, trabeculate,
filiform, persistent, hyaline, embedded in mucilage, anastomosing between and above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical to narrowly fusoid, short
Fungal Diversity (2020) 105:319–575
pedicellate. Ascospores mostly uniseriate, fusoid, hyaline,
septate, symmetrical, smooth, thin-walled, surrounded by
a hyaline, gelatinous sheath (Zhang et al. 2008c, 2009b,
2012b). Asexual morph: Hyphomycetous. Conidiophores
usually simple. Conidia curved or sigmoid, tapering to the
end, septate, hyaline (illustration and description see Ingold
(1942)).
Type species: Amniculicola lignicola Y. Zhang ter & K.D.
Hyde
Notes: The freshwater genus Amniculicola is characterized by staining the woody substrate purple (Zhang et al.
2008c, 2009b). However, two other species, A. aquatica
Z.L. Luo et al. and A. guttulata et al., did not produce pigmentation on the host substrate (Hyde et al. 2019; this
study, Fig. 22). A comprehensive account of Amniculicola
was provided by Zhang et al. (2008c, 2012b). Amniculicola
immersa Y. Zhang ter, J. Fourn., A. lignicola and A. parva Y.
Zhang ter, J. Fourn. were collected from Europe (Denmark
and France), A. longissima (Sacc. & P. Syd.) Nadeeshan &
K.D. Hyde was collected from Latin America (Costa Rica),
while A. aquatica and A. guttulata were collected from Asia
(China). Amniculicola longissima is the only asexual morphic species in the genus and confirmed by phylogenetic
analyses (Hyde et al. 2019; this study, Fig. 2).
The paraphyletic nature of Amniculicola was shown in
Wanasinghe et al. (2015), as Amniculicola species clustered in three different sister clades. This nature is not well
resolved as one asexual genus Vargamyces Tóth nested
within Amniculicola in Hyde et al. (2019) and this study
(Fig. 2).
List of freshwater Amniculicola species
*Amniculicola aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
Fungal Diversity 96: 13 (2019)
Freshwater distribution: China (Hyde et al. 2019)
*Amniculicola guttulata Z.L. Luo, K.D. Hyde & H.Y. Su,
Fungal Diversity 96: 13 (2019); Fig. 22
Freshwater distribution: China (Hyde et al. 2019, this
study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 160–180 μm high, 170–190 μm
diam., black, solitary, scattered or occasionally gregarious,
superficial, subglobose to conical, with ostiolate papilla.
Peridium 35–45 μm thick, comprising several layers of hyaline to pale brown, thin-walled, large cells of textura angularis or irregular cells. Pseudoparaphyses 2.2 μm diam.,
dense, trabeculate, filiform, persistent, hyaline, embedded
in mucilage, anastomosing between and above the asci. Asci
105–141(–256) × 8.5–10.5 μm (x̄ = 158.5 × 10 μm, n = 10),
8-spored, bitunicate, fissitunicate, cylindrical to narrowly
fusiform, with a twisted, bifurcate, short pedicel, 7.5–18.5
μm long, apically rounded or slightly obtuse, endoascus
367
elongate to 256 μm long when mature, with a small inconspicuous apical apparatus barely seen in water. Ascospores
21.5–27 × 5.2–6.2 μm ( x̄ = 23.7 × 5.6 μm, n = 15), uniseriate or overlapping uni-seriate, straight or slightly curved,
hyaline, 1-septate, obscure when young, visible when germinated, deeply constricted at the septa, slightly constricted
at a quarter, the upper cell slightly broader than the lower
one, broadly fusiform, with two prominent big guttules in
the middle cell and four gradually smaller ones beside, thinwalled, smooth, with a thin, hyaline, bipolar narrowed, mucilaginous sheath which is 1–2 μm thick. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 25 mm diam. in 25 days at 25 °C, grey to white from
above, pale grey from below, surface rough, fluffy, with
dense mycelium, dry, raised, edge entire, with white margin.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H55A-1 (MFLU 18-1160), living culture MFLUCC
18-1038; ibid., H55A-2 (HKAS 101736), living culture
KUMCC 18-0088.
Notes: Our new collection MFLUCC 18-1038 is identified as Amniculicola guttulata based on morphological
characters; identical LSU and TEF sequence data and phylogenetic analysis (Fig. 2). The asci in the holotype are up
to 127 μm long, while in our collection they are up to 256
μm long when mature (Hyde et al. 2019).
*Amniculicola immersa Y. Zhang ter, J. Fourn., Crous &
K.D. Hyde, Persoonia 23: 50 (2009)
Freshwater distribution: Denmark (Zhang et al. 2009b)
*Amniculicola lignicola Y. Zhang ter & K.D. Hyde, Mycological Research 112 (10): 1189 (2008)
Freshwater distribution: France (Zhang et al. 2008c)
*Amniculicola longissima (Sacc. & P. Syd.) Nadeeshan &
K.D. Hyde, IMA Fungus 7(2): 301 (2016)
Freshwater distribution: Leicester, UK (Ingold 1942)
*Amniculicola parva Y. Zhang ter, J. Fourn., Crous & K.D.
Hyde, Persoonia 23: 52 (2009)
Freshwater distribution: France (Zhang et al. 2009b)
Key to freshwater Amniculicola species
1.
1.
2.
2.
3.
3.
4.
Asexual morph……………………………A. longissima
Sexual morph…………………………………………2
Ascomata superficial…………………………………3
Ascomata immersed………………………A. immersa
Asci longer than 130 μm……………………A. lignicola
Asci shorter than 130 μm………………………………4
Substrate stained purple………………………A. parva
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Fungal Diversity (2020) 105:319–575
◂Fig. 22 Amniculicola guttulata (MFLU 18-1160). a Appearance of
black ascomata on host. b Vertical section of ascoma. c Structure of
peridium. d Pseudoparaphyses. e–h Bitunicate asci. i–n Ascospores.
o Germinated ascospore. p Colony on PDA (from front and reverse).
Scale bars: b = 50 μm, c, e–h = 20 μm, d, i–o = 10 μm
4. Substrate natural colour………………………………5
5. Peridium 35–50 µm thick, ascospores 24–32 × 6–8
µm…………………………………………A. aquatica
5. Peridium 27–35 µm thick, ascospores 23–27 × 5–7
µm…………………………………………A. guttulata
Murispora Yin. Zhang, C.L. Schoch, J. Fourn., Crous &
K.D. Hyde, Studies in Mycology 64: 95 (2009)
Saprobic on submerged wood or dead stems or branches
of plant materials. Sexual morph: Ascomata scattered to
gregarious, immersed, uniloculate, lenticular, slightly protruding at the apex, opening through a small rounded pore,
with ostiolate papilla, stain substrate purple. Peridium thin,
composed of a few layers cells of textura angularis, thicker
at the apex with pseudoparenchymatous cells. Pseudoparaphyses cellular, embedded in mucilage. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate, short pedicellate. with
a small ocular chamber. Ascospores biseriate to uniseriate
near the base, fusoid with narrowly rounded ends, slightly
curved, golden yellow or hyaline, turning brown when
senescent, muriform, with one, rarely two longitudinal septa
in all cells except end cells, 7–9-transversely septate, basically symmetrical, smooth or finely verruculose, surrounded
by a wide mucilaginous sheath (Zhang et al. 2009c, 2012b).
Asexual morph: Coelomycetous. Pycnidia solitary, mainly
immersed, uniloculate, dark brown to black. Peridium reddish brown cells of textura angularis, with inner most layer
thin, hyaline. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells phialidic, determinate, hyaline, smooth,
formed from the inner most layer of pycnidial wall. Conidia
ellipsoidal, hyaline, aseptate, smooth-walled, thin-walled
(Wanasinghe et al. 2015).
Type species: Murispora rubicunda (Niessl) Y. Zhang ter,
J. Fourn. & K.D. Hyde
Notes: Zhang et al. (2009c) introduced Murispora based
on Pleospora rubicunda Niessl which clustered in a robust
clade with Amniculicola species in Amniculicolaceae.
Wanasinghe et al. (2015) introduced another six species
to Murispora from terrestrial habitats and they found the
asexual morph of M. hawksworthii Wanas. et al. from single
ascospore isolates. Murispora species stain substrate purple
(Zhang et al. 2009c, 2012b; Wanas. et al. 2015; Bao et al.
2019d) and this character should be a common feature to
the genus. Four freshwater species have been reported in
Murispora, three of which were collected from China (see
list below).
369
List of freshwater Murispora species
*Murispora aquatica D.F. Bao, Z.L. Luo, K.D. Hyde &
H.Y. Su, Phytotaxa 416(1): 5 (2019)
Freshwater distribution: China (Bao et al. 2019d)
*Murispora cicognanii Wanas., Camporesi, E.B.G. Jones &
K.D. Hyde, Cryptog. Mycol. 36(4): 437 (2015)
Freshwater distribution: China (Hyde et al. 2019)
*Murispora fagicola Wanas., Camporesi, E.B.G. Jones &
K.D. Hyde, Cryptog. Mycol. 36(4): 429 (2015)
Freshwater distribution: China (Bao et al. 2019d)
*Murispora rubicunda (Niessl) Yin. Zhang, J. Fourn. &
K.D. Hyde, Studies in Mycology 64: 96 (2009); Fig. 23
Basionym: Pleospora rubicunda Niessl, Verh. nat. Ver.
Brünn 14: 191 (1876)
Synonymy: Sphaeria rubicunda Schwein., Transactions
of the American Philosophical Society 4 (2): 222 (1832)
Massariosphaeria rubicunda (Niessl) Crivelli, Über
die heterogene Ascomycetengattung Pleospora Rabh.: 144
(1983)
Karstenula rubicunda (Niessl) M.E. Barr, N. Amer. Fl.,
Ser. 2 (New York) 13: 52 (1990)
Freshwater distribution: Austria (Magnes and Hafellner
1991), France (Zhang et al. 2012b)
Key to freshwater Murispora species
1. Ascospores > 31 μm long………………………………2
1. Ascospores < 31 μm long………………………………3
2. Ascospores more than 10 transverse septa………………
…………………………………………M. cicognanii
2. Ascospores 7–9 transverse septa…………M. rubicunda
3. Ascospores 21–25 × 8–10 μm……………M. aquatica
3. Ascospores 27–31 × 11–13 μm……………M. fagicola
Neomassariosphaeria Y. Zhang ter, J. Fourn. & K.D. Hyde,
Stud. Mycol. 64: 96 (2009)
Saprobic on submerged culm of Phragmites. Sexual
morph: Ascomata scattered or in small groups, deeply
immersed, lenticular or depressed ellipsoidal, with ostiolate
papilla, stain substrate purple, visible as black, slit-like spot
on the surface of substrate. Peridium comprising several layers of rounded to angular, brown to dark brown, thick-walled
cells. Pseudoparaphyses numerous, trabeculate, filiform,
hyaline, embedded in a matrix. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate to broadly clavate, with an ocular
chamber and short pedicel. Ascospores bi- to tri-seriate, narrowly to broadly fusoid, straight or curved, multi-septate,
reddish brown, minutely verruculose, usually with a thin
gelatinous sheath. Asexual morph: Undetermined.
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◂Fig. 23 Murispora rubicunda (Material examined: FRANCE, Haute-
Garonne, Avignonet, on submerged dead herbaceous stem, 16 January 2007, M. Delpont, IFRD 2017). a Herbarium label and specimen
of Murispora rubicunda. b Appearance of ascomata on host surface.
c Vertical section through ascoma. d Peridium. e, f Asci. g Apex
of ascus. h Pedicel. j Pseudoparaphyses. i, k Ascospores with wide
sheath in Indian Ink. Scale bars: c = 100 μm, d = 20 μm, e, f = 25
μm, g–k = 10 μm
Type species: Neomassariosphaeria typhicola Y. Zhang
ter, J. Fourn. & K.D. Hyde
Notes: Neomassariosphaeria was established in Amniculicolaceae to accommodate Massariosphaeria typhicola (P.
Karst.) Leuchtm (Zhang et al. 2009c). Later, it was transferred to Lindgomycetaceae based on phylogenetic analysis (Ariyawansa et al. 2015). However, we found that CBS
123126 is a problematic strain, which possesses two different LSU sequences and clusters in different families, i.e.
FJ795504 (Zhang et al. 2009c) in Lindgomycetaceae and
GU301844 (Schoch et al. 2009) in Amniculicolaceae. We
substantiate GU301844 is the real sequence of Neomassariosphaeria typhicola (P. Karst.) Y. Zhang ter et al., while
FJ795504, listed in Zhang et al. (2009c), is invalid (pers.
comm. Zhang Y.). Ariyawansa et al. (2015) only included
the invalid sequence FJ795504 in their phylognetic tree,
which resulted in the classification of Neomassariosphaeria
in Lindgomycetaceae.
Neomassariosphaeria often stains the woody substrate
purple, and has reddish brown, multi-septate ascospores
with an irregular, hyaline, gelatinous sheath, which morphologically fit well with the familial concept of Amniculicolaceae (Zhang et al. 2009c; Hyde et al. 2013). Based
on morphology and our phylogenetic tree (Fig. 2), we follow Zhang et al. (2009c), to retain Neomassariosphaeria
in Amniculicolaceae. Accession number FJ795504 bearing
CBS123126 is treated as an invalid sequence.
Neomassariosphaeria typhicola (IFRD 2018) was considered as a synonym of Massariosphaeria typhicola (Zhang
et al. 2009c). Although they are similar in the ascomata that
staining the substrate purple, re-examination of IFRD 2018
(Fig. 24) shows that it is different from M. typhicola in having
lenticular ascomata with an elongate papilla on the substrate,
dark brown to black, reddish brown ascospores (Leuchtmann
1984). Phylogeny also placed them in different families, i.e.
N. typhicola in Amniculicolaceae, M. typhicola in Lindgomycetaceae (Zhang et al. 2009c; this study, Fig. 2). We
therefore, treat Neomassariosphaeria typhicola and Massariosphaeria typhicola as different species in this study.
List of freshwater Neomassariosphaeria species
*Neomassariosphaeria typhicola Y. Zhang ter, J. Fourn. &
K.D. Hyde, Stud. Mycol. 64: 96 (2009)
Facesoffungi number: FoF09244; Fig. 24
371
Saprobic on submerged culm of Phragmites. Sexual
morph: Ascomata 150–250 μm high, 300–500 μm diam.,
scattered or gregarious, immersed, lenticular or depressed
ellipsoidal, with ostiolate papilla, stain the substrate purple,
visible as a black, slightly protruding elongate papilla on
the substrate. Peridium 20–35 μm thick, comprising several
layers of rounded to angular, brown to dark brown, thickwalled cells. Pseudoparaphyses 1–2 μm wide, numerous,
uneven in width or tuberculed, trabeculate, hyaline, septate, embedded in hyaline to reddish matrix. Asci 90–120 ×
14–20 μm ( x̄ = 97 × 16.5 μm, n = 10), 8-spored, bitunicate,
fissitunicate, cylindro-clavate to broadly clavate, apically
rounded, with a well-developed ocular chamber and short
pedicel. Ascospores 30–40 × 6–8 μm ( x̄ = 34.5 × 6.5 μm,
n = 20), bi- to tri-seriate, narrowly to broadly fusoid, with
subacute or rounded ends, straight or curved, 7-septate, constricted at the septa, dark brown to black or reddish brown,
minutely verruculose, somewhat thick-walled, usually with a
thin, broadly fusoid or irregular, gelatinous sheath. Asexual
morph: Undetermined.
Material examined: DENMARK, Sjaeland, Frederikskilde, Suserup Skove, Tystrup Lake, on submerged culm
of Phragmites, 25 May 2007, leg. & det. J. Fournier (IFRD
2018).
Notes: We re-examined the specimen Neomassariosphaeria typhicola IFRD 2018 collected by Zhang et al. (2009c)
to confirm its generic placement in Amniculicolaceae. The
sequence data (LSU: GU301844, SSU: GU296174, RPB2:
GU371795) and morphological characters of N. typhicola
support it as a distinct genus in Amniculicolaceae. The
reddish brown, phragmoseptate ascospores of Neomassariosphaeria differs from hyaline, 1-septate ascospores of
Amniculicola and golden yellow to brown, dictyoseptate
ascospores of Murispora (see key to freshwater genera of
Amniculicolaceae).
Vargamyces Tóth, Acta bot. hung. 25: 403 (1980) [1979]
Saprobic on submerged wood or leaves. Sexual morph:
Undetermined. Asexual morph: Mycelium mostly
immersed, composed of septate, hyaline to pale brown,
smooth hyphae. Conidiophores micronematous to semimacronematous, solitary, erect, straight to flexuous, cylindrical, subhyaline to pale brown, with percurrent proliferations. Conidiogenous cells integrated, terminal, cylindrical
to cupuliforme, hyaline to pale brown. Conidia solitary, fusiform, truncate at the base, apically rounded, paler toward
the ends, multi-septate, smooth-walled. Microconidia blastic
growing on undifferentiated hyphae, solitary, terminal, lateral or intercalary, globose to obovoid, unicellular, hyaline,
smooth (Hernández-Restrepo et al. 2017).
Type species: Vargamyces aquaticus (Dudka) Tóth
Notes: The monotypic genus Vargamyces was introduced
by Tóth (1979) to accommodate V. aquaticus which was
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◂Fig. 24 Neomassariosphaeria typhicola (IFRD 2018). a Herbarium
packet and specimen. b Ascomata immersed in host substrate. c
Vertical section of ascoma. d Structure of peridium. e Pseudoparaphyses. f, g Mature asci. h Immature ascus. i An ocular chamber in
mature ascus. j Verruculose ascospores. k Ascospore in Indian Ink.
l Ascospore in ascus. m Ascospore with a thin, gelatinous sheath. n
Mature ascospore. Scale bars: c = 100 μm, d, f–h = 20 μm, e = 3 μm,
i–n = 10 μm
collected from submerged leaves and wood (Dudka 1966;
Hyde and Goh 1999a). Vargamyces aquaticus was transferred to Xylomyces based on detachable, fusiform, dematiaceous and multiseptate chlamydospores (Hyde and Goh
1999a). However, Hernández-Restrepo et al. (2017) showed
that V. aquaticus clustered in Amniculicolaceae, which is
phylogenetically distant from the type species of Xylomyces,
X. chlamydosporus (this study, Fig. 2). Therefore, Vargamyces is distinct and the only asexual genus in Amniculicolaceae, as listed in Wijayawardene et al. (2018, 2020).
The lectotype (Fig. 1 in Dudka (1966)) and epitype (CBS
H-22992, culture ex-epitype CBS 636.91) were also designated by Hernández-Restrepo et al. (2017).
List of freshwater Vargamyces species
*Vargamyces aquaticus (Dudka) Tóth, Acta Mus. Silesiae,
Ser. A 25(3–4): 403 (1980) [1979]
Basionym: Camposporium aquaticum Dudka, Ukr. Bot.
Zh. 23:91 (1966)
Synonymy: Sporidesmium ontariense Matsush., Matsush.
Mycol. Mem. 3: 16 (1983)
Xylomyces aquaticus (Dudka) K.D. Hyde & Goh, Mycol.
Res. 103(12): 1573 (1999)
Repetophragma ontariense (Matsush.) W.P. Wu, Fungal
Diversity Res. Ser. 15: 82 (2005)
Freshwater distribution: England (Hyde and Goh 1999a),
Spain (Hernández-Restrepo et al. 2017), Ukraine (Dudka
1966), USA (Tóth 1979)
Anteagloniaceae K.D. Hyde, J.K. Liu & A. Mapook, Fungal
Diversity 63: 33 (2013)
373
Saprobic on submerged bamboo or dead wood. Sexual
morph: Ascostromata solitary to gregarious, erumpent to
superficial, visible as dull, black, dark at the basal region
and orange brown at the apex, conical to lenticular, with
a flattened base, uni- to bi-loculate, glabrous, coriaceous,
ostiolate, with pore-like opening. Peridium thick-walled, of
unequal thickness, poorly developed at the base, composed
of several layers of dark, pseudoparenchymatous cells, with
fungal tissue intermixed with the host cells. Pseudoparaphyses dense, trabeculate, filiform, hyaline, septate. Asci
8-spored, bitunicate, fissitunicate, clavate to cylindricclavate, short pedicellate, apically rounded with well-developed ocular chamber. Ascospores uni- to bi-seriate, fusiform,
hyaline, becoming brown when released from asci, septate,
smooth-walled, with or without mucilaginous sheath (Liu
et al. 2015). Asexual morph: Undetermined.
Type species: Flammeascoma bambusae Phook. & K.D.
Hyde
Notes: Flammeascoma was introduced to accommodate
F. bambusae, which was collected from submerged bamboo
(Liu et al. 2015). Flammeascoma is similar to familial type
Anteaglonium Mugambi & Huhndorf in the shape of asci
and ascospores (Mugambi and Huhndorf 2009, Liu et al.
2015). However, Flammeascoma is unique in producing reddish orange pigmentation at the apex of ascostromata, while
Anteaglonium lacks this character and always sits on a thin
darkened crust with or without sparse, dark brown, septate
subiculum (Mugambi and Huhndorf 2009; Ariyawansa et al.
2015; Liu et al. 2015).
Flammeascoma appears to be polyphyletic as F. bambusae (type) clusters with Anteaglonium gordoniae Jayasiri
et al. and F. lignicola Boonmee & K.D. Hyde clusters with
our new genus Purpureofaciens, but without bootstrap support (Fig. 2). More species are expected to clarify whether
Flammeascoma is polyphyletic.
List of freshwater Flammeascoma species
*Flammeascoma bambusae Phookamsak & K.D. Hyde,
Fungal Diversity 72(1): 64 (2015); Fig. 25
Freshwater distribution: Thailand (Liu et al. 2015)
Key to freshwater sexual genera of Anteagloniaceae
1. Ascomata orange-pigmented at the apex, asci clavate to
cylindric-clavate, ascospores hyaline, fusiform…………
…………………………………………Flammeascoma
2. Ascomata stain the substrate purple, asci cylindrical,
ascospores olivaceous, ellipsoidal………………………
………………………………………Purpureofaciens
Flammeascoma Phook. & K.D. Hyde, Fungal Diversity
72(1): 63 (2015)
Purpureofaciens W. Dong, H. Zhang & K.D. Hyde, gen.
nov.
Index Fungorum number: IF557804; Facesoffungi number: FoF08717
Etymology: referring to staining the substrate purple
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata black, scattered or gregarious,
semi-immersed to superficial, conical, flattened at the basal
region, stain the substrate purple and produce reddish pigmentation at the apex, coriaceous, with a large, protuberant, ostiolate papilla. Peridium of unequal thickness, poorly
developed at the base, composed of reddish brown, large
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Fig. 25 Flammeascoma bambusae (Material examined: THAILAND, Chiang Mai Province, Chom Tong District, Doi Inthanon, on
dead stem of submerged bamboo (Bambusae), 5 September 2009, R.
Phookamsak, RP0013, MFLU 11-0143, holotype). a Appearance of
ascostromata on the host surface. b Section through an ascostroma. c
Peridium. d–f Bitunicate asci. g Pseudoparaphyses. h–m Ascospores.
Scale bars: b = 200 μm, c–f = 50 μm, g–m = 20 μm
cells of textura angularis. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, embedded in a hyaline gelatinous
matrix. Asci 8-spored, bitunicate, cylindrical, pedicellate,
apically rounded. Ascospores uniseriate, curved, olivaceous,
septate, ellipsoidal with acute ends, guttulate, thin-walled,
smooth, surrounded by a thin mucilaginous sheath. Asexual
morph: Undetermined.
Type species: Purpureofaciens aquatica W. Dong, H.
Zhang & K.D. Hyde
Notes: Purpureofaciens is unique by semi-immersed
to superficial, conical ascomata staining woody substrate
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purple, cylindrical asci, ellipsoidal, curved, olivaceous
ascospores with acute ends and white to reddish colonies
in culture (Mugambi and Huhndorf 2009; Liu et al. 2015).
The reddish culture is also observed in a strain of Anteaglonium parvulum (MFLUCC 14-0821), which has similar
shape of ascospores, but the characters of ascomata, asci and
ascospores can easily separate them (Jayasiri et al. 2016).
Flammeascoma has reddish pigmentation at the apex of
ascomata, which resembles Purpureofaciens. However, the
conical ascomata with a large, protuberant, ostiolate papilla,
cylindrical asci and ellipsoidal, olivaceous ascospores of
Purpureofaciens are different from those of Flammeascoma
(Liu et al. 2015). Purpureofaciens clusters with Flammeascoma species with low bootstrap support (Fig. 2) and morphology separates them as distinct genera. Purpureofaciens
comprises one freshwater species P. aquatica which was
collected from Thailand.
List of freshwater Purpureofaciens species
*Purpureofaciens aquatica W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557902; Facesoffungi number: FoF08718; Fig. 26
Etymology: in reference to aquatic habitat of the fungus
Holotype: MFLU 18-1569
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 200–300 μm high, 300–400 μm
diam., dark brown to black, scattered or gregarious, semiimmersed to superficial, conical, flattened at the basal
region, stain the substrate purple and produce reddish pigmentation at the apex, coriaceous, with a large, protuberant, ostiolate papilla, 110–130 μm long, 110–130 μm wide.
Peridium unequal thickness, poorly developed at the base,
up to 45 μm at the sides and 15 μm at the base, composed of
reddish brown, large cells of textura angularis. Pseudoparaphyses 2 μm diam., numerous, trabeculate, filiform, hyaline,
embedded in a hyaline gelatinous matrix. Asci 135–150 ×
8.5–11 μm (x̄ = 143 × 9.7 μm, n = 10), 8-spored, bitunicate,
cylindrical, pedicellate, apically rounded. Ascospores 15–22
× 7–10 μm ( x̄ = 18.5 × 8 μm, n = 20), uniseriate, curved,
olivaceous, 1-septate, constricted at the septum, ellipsoidal
with acute ends, with two prominent guttules when immature, thin-walled, smooth, surrounded by a thin mucilaginous sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 50 mm diam. in 25 days at 25 °C, annulate, white to
reddish to purplish red from center to edge, surface rough,
dry, raised, edge entire.
Material examined: THAILAND, Satun Province, on
submerged wood in a stream, 10 May 2018, W. Dong,
hat266-1 (MFLU 18-1569, holotype), ex-type living culture MFLUCC 18-1241; ibid. hat266-2 (HKAS 105016,
isotype), ex-type living culture KUMCC 19-0032.
375
Aquasubmersaceae A. Hashim. & Kaz. Tanaka, Persoonia
39: 56 (2017)
Aquasubmersa K.D. Hyde & H. Zhang, Cryptog. Mycol.
33(3): 340 (2012)
Saprobic on submerged or dead wood. Sexual morph:
Ascomata scattered to grouped, immersed to semi-immersed,
uniloculate, subglobose, glabrous, dark brown to black, with
ostiolate papilla. Peridium composed of flattened, thinwalled cells of textura angularis. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, septate. Asci 8-spored,
bitunicate, cylindrical, short pedicellate, apically rounded,
with an ocular chamber. Ascospores biseriate, broadly fusiform with rounded ends, straight, hyaline, with one mostly
median septum, with gelatinous sheath (Ariyawansa et al.
2015). Asexual morph: Coelomycetous. Pycnidia solitary
to scattered, semi-immersed to superficial, subglobose to
ellipsoidal, dark brown to black, ostiolate, minute or without papilla. Ostiole circular, dark-brown, central. Peridium
composed of polygonal, hyaline to brown cells. Conidiophores reduced. Conidiogenous cells holoblastic, determinate, lageniform, hyaline, smooth. Conidia ellipsoidal, some
with papillate base, hyaline, aseptate, smooth, thin-walled,
without sheath or appendage (Zhang et al. 2012a).
Type species: Aquasubmersa mircensis H. Zhang & K.D.
Hyde
Notes: Aquasubmersa was established to accommodate a
freshwater coelomycetous species, A. mircensis (Zhang et al.
2012a). It was placed in the family Lophiotremataceae in previous studies (Ariyawansa et al. 2015; Doilom et al. 2017). A
comprehensive study on Lophiotremataceae based on morphological observations and multigene analyses showed that
Aquasubmersa should be transferred to a newly established
family Aquasubmersaceae (Hashimoto et al. 2017). The second species, A. japonica A. Hashim. & Kaz. Tanaka with
sexual morph, was also collected from freshwater habitats
and conidiomata were produced in culture (Ariyawansa et al.
2015). The asexual morph of A. japonica matches the generic
concept of Aquasubmersa. It differs from A. mircensis by
pycnidia with minute papilla, thicker peridium and conidia
lacking papillate bases. The type material of A. mircensis was
re-examined by Li et al. (2020) and this study.
List of freshwater Aquasubmersa species
*Aquasubmersa japonica A. Hashim. & Kaz. Tanaka, Fungal Diversity 75: 87 (2015)
Freshwater distribution: Japan (Ariyawansa et al. 2015)
*Aquasubmersa mircensis H. Zhang & K.D. Hyde, Cryptog. Mycol. 33(3): 340 (2012); Fig. 27
Freshwater distribution: Thailand (Zhang et al. 2012a)
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376
Fig. 26 Purpureofaciens aquatica (MFLU 18-1569, holotype). a–c
Appearance of black ascomata on host surface. d, e Vertical section
of ascomata. f Structure of peridium. g Pseudoparaphyses. h, i Bituni-
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cate asci. j–l Ascospores. m Germinated ascospores. n, o Colony on
PDA (left-front, right-reverse). Scale bars: d, e = 100 μm, f–i, m =
20 μm, j–l = 10 μm
Fungal Diversity (2020) 105:319–575
377
Fig. 27 Aquasubmersa mircensis (Material examined: THAILAND,
Chiang Mai Province, Mushroom Research Centre, on submerged
wood, 21 April 2011, H. Zhang, m3, MFLU 11-1001, holotype). a
Pycnidium on wood surface. b Section of pycnidium. c Peridium with
conidiogenous cells. d Conidiogenous cells with conidia. e, f Conidia.
Scale bars: b = 100 µm, c = 20 µm, d = 15 µm, e, f = 10 µm
Key to freshwater Aquasubmersa species
1. Pycnidia up to 215 μm high, 115–195 μm diam., with
minute papilla, peridium > 20 µm thick, conidia without
papillate base……………………………………A. japonica
1. Pycnidia 130–170 μm high, 150–250 μm diam., without papilla, peridium < 20 μm thick, conidia with papillate
base……………………………………………A. mircensis
superficial, hemisphaerical to conical, base flattened, carbonaceous, with ostiolate papilla. Ostiole central, relatively
broad, circular, black. Peridium uneven in width, carbonized, composed of black amorphous layer, whose cells are
obscured. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, sparsely septate. Asci 8-spored, bitunicate,
clavate to narrowly ellipsoidal, with a short pedicel, apically
rounded. Ascospores 2–4-seriate, hyaline, septate, subcylindrical, thin-walled, smooth. Asexual morph: Undetermined.
Type species: Aquatospora cylindrica W. Dong, H. Zhang
& K.D. Hyde
Notes: Aquatospora clusters with Acrocordiopsis Borse &
K.D. Hyde and Xenoastrosphaeriella Jayasiri et al. with low
bootstrap support (Fig. 2). Acrocordiopsis is a problematic
genus, which was placed in Salsugineaceae by Hyde et al.
(2013) and Caryosporaceae by Ariyawansa et al. (2015).
Acrocordiopsis was referred to Salsugineaceae (Wijayawardene et al. 2018, 2020) based on some common morphology between Acrocordiopsis and Salsuginea K.D. Hyde
(Borse and Hyde 1989; Hyde 1991). Acrocordiopsis clustered distantly from Salsuginea in the phylogenetic tree of
Zhang et al. (2018) and close to Aquatospora in our study
(Fig. 2). Xenoastrosphaeriella differs from Aquatospora
in having paler end cells of ascospores (Hawksworth and
Boise 1985; Phookamsak et al. 2015b; Jayasiri et al. 2019).
Aquatospora shares similar characters with Astrosphaeriellaceae members in having large, erumpent to superficial,
hemisphaerical or conical, carbonaceous ascomata with
nearly flattened bases (Borse and Hyde 1989; Ariyawansa
et al. 2015; Phookamsak et al. 2015b). However, the unique
combination of morphology of clavate to narrowly ellipsoidal asci, hyaline, cylindrical ascospores and molecular characters warrant Aquatospora as a new genus (more details see
key to freshwater genera of Astrosphaeriellaceae).
Astrosphaeriellaceae Phook. & K.D. Hyde, Fungal Diversity 74: 161 (2015)
Key to freshwater genera of Astrosphaeriellaceae
1.
1.
2.
2.
3.
3.
4.
4.
Asci mostly broadly clavate……………………………2
Asci cylindrical………………………………………3
Ascospores hyaline, cylindrical…………Aquatospora
Ascospores mostly dark brown to black, fusiform to
broad-ellipsoid or biconic…………………Caryospora
Producing hyphomycetous asexual morph………………
………………………………………………Pithomyces
Producing coelomycetous asexual mor ph or
unknown………………………………………………4
Ascospores brown to reddish brown, with paler end
cells………………………………Xenoastrosphaeriella
Ascospores hyaline to brown, without paler end
cells……………………………………Astrosphaeriella
Aquatospora W. Dong, H. Zhang & K.D. Hyde, gen. nov.
Index Fungorum number: IF557903; Facesoffungi number: FoF09245
Etymology: referring to aquatic habitat of this fungus
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata black, scattered or gregarious,
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Fungal Diversity (2020) 105:319–575
◂Fig. 28 Aquatospora cylindrica (MFLU 18-1534, holotype). a
Appearance of black ascomata on host. b, c Vertical section of
ascoma. d Structure of peridium. e Pseudoparaphyses. f–h Bitunicate asci. i–k Ascospores. k Ascospore in Indian Ink. l Germinated
ascospore. m, n Colony on PDA (left-front, right-reverse). Scale bars:
c = 200 μm, d, f–l = 50 μm, e = 20 μm
List of freshwater Aquatospora species
*Aquatospora cylindrica W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557904; Facesoffungi number: FoF09246; Fig. 28
Etymology: referring to cylindrical ascospores of this
fungus
Holotype: MFLU 18-1534
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 600–650 μm high, 800–900 μm
diam., black, scattered or gregarious, superficial, hemisphaerical to conical, base flattened, carbonaceous, with
ostiolate papilla. Ostiole central, relatively broad, circular,
black. Peridium up to 155 μm at the sides and 65 μm at
the base, carbonized, composed of black amorphous layer,
whose cells are obscured. Pseudoparaphyses 0.5 μm diam.,
numerous, trabeculate, filiform, hyaline, sparsely septate.
Asci 175–270 × 50–65 μm ( x̄ = 225 × 58 μm, n = 10),
8-spored, bitunicate, clavate to narrowly ellipsoidal, straight
or slightly curved, with a short pedicel, apically rounded.
Ascospores 70–85 × 20–25 μm ( x̄ = 77 × 23 μm, n = 10),
2–4-seriate, cylindrical, slightly tapering at two sides, 1-septate, constricted at the septum, hyaline, straight, with four
prominent big guttules, thin-walled, smooth, without sheath.
Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching
40 mm in 35 days at 25 °C, brown to white from above, dark
brown from below, surface rough, dry, raised, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat146-1 (MFLU 18-1534, holotype), ex-type living culture MFLUCC 18-1287; ibid., hat146-2 (HKAS 105042,
isotype), ex-type living culture KUMCC 19-0060.
Astrosphaeriella Syd. & P. Syd., Annls mycol. 11(3): 260
(1913)
Saprobic or parasitic on bamboo, palm or stout grasses in
freshwater or terrestrial habitats. Sexual morph: Ascostromata solitary to gregarious, erumpent to superficial, conical
or mammiform, at maturity with stellate appearance from
above, dark opaque, uniloculate, glabrous, brittle, carbonaceous, ostiolate. Peridium relatively thick, poorly developed
at the base, composed of thick, opaque and melanized cells,
arranged in a textura angularis with palisade-like cells at
the rim. Pseudoparaphyses dense, trabeculate, filiform,
hyaline, anastomosing, embedded in a hyaline gelatinous
379
matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical
to cylindric-clavate, subsessile to short-pedicellate, apically
rounded with ocular chamber. Ascospores overlapping unito bi-seriate, hyaline to brown, fusiform with rounded to
acute ends, septate, smooth, with or without appendages or
mucilaginous sheaths (Phookamsak et al. 2015b). Asexual
morph: Coelomycetous. Conidiomata pycnidial, scattered
or solitary, immersed to superficial, conical or hemisphaerical to globose. Pycnidial walls composed of several layers of
dark brown to black, textura angularis to textura intricata of
cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic or phialidic, integrated, cylindrical or cylindric-clavate or ampulliform, septate or aseptate,
hyaline, smooth. Conidia globose to subglobose, or oblong,
aseptate, hyaline, smooth (Phookamsak et al. 2015b).
Type species: Astrosphaeriella fusispora Syd. & P. Syd.
Notes: Astrosphaeriella is characterized by carbonaceous, conical to mammiform ascostromata with ruptured,
reflexed, stellate, host remnants and hyaline to brown, fusiform ascospores (Liu et al. 2011; Phookamsak et al. 2015b).
Astrosphaeriella aquatica K.D. Hyde, A. seychellensis K.D.
Hyde & Goh and A. thailandensis J. Ren et al. were only
found from freshwater habitats (Hyde 1994; Hyde and
Frohlich 1998; Hyde and Goh 1998b; Ren et al. 2013). Other
freshwater Astrosphaeriella species were found from both
freshwater and terrestrial habitats (see list below). Astrosphaeriella aquatica resembles A. seychellensis and A. thailandensis in having hyaline, fusiform ascospores, but differs
in having a wide spreading mucilaginous sheath which is
drawn out at the poles.
Phookamsak et al. (2015b) transferred Astrosphaeriella
sensu lato, which have coriaceous ascomata with short to
long neck and striate ascospores, to a newly established family Pseudoastrosphaeriellaceae. Astrosphaeriella papuana
Aptroot has striate ascospores which are typical characters
of Pseudoastrosphaeriellaceae (Aptroot 1995; Hyde and
Frohlich 1998; Phookamsak et al. 2015b). However, molecular data for A. papuana is unavailable, thus this species is
retained in Astrosphaeriellaceae. Astrosphaeriella papuana
might be transferred to Pseudoastrosphaeriellaceae when the
type specimen is recollected and recircumscribed.
Astrosphaeriella stellata (Pat.) Sacc. was treated as an
earlier name of A. fusispora and followed by some studies
(Hyde and Frohlich 1998; Hyde et al. 2000; Tanaka et al.
2009; Liu et al. 2011). Liu et al. (2011) named one collection
MFLUCC 10-0555 as A. stellata, which was later considered
as a reference specimen of A. fusispora (Phookamsak et al.
2015b). Therefore, A. stellata isolated from freshwater habitats is probably a collection of A. fusispora.
The asexual morph was developed in the culture of Astrosphaeriella bambusae Phookamsak& K.D. Hyde (Phookamsak et al. 2015b).
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380
List of freshwater Astrosphaeriella species
Astrosphaeriella aquatica K.D. Hyde, Mycol. Res. 98(7):
719 (1994)
Freshwater distribution: Papua New Guinea (Hyde 1994),
Ecuador (Hyde and Frohlich 1998)
Astrosphaeriella exorrhiza Boise, Sydowia 38: 117 (1986)
[1985]
Freshwater distribution: Ecuador (Hyde and Frohlich
1998)
Astrosphaeriella maquilingiana (Rehm) K.D. Hyde & J.
Fröhl., Sydowia 50(1): 103 (1998)
Basionym: Trematosphaeria maquilingiana Rehm, Leafl.
of Philipp. Bot. 8: 2952 (1916)
Synonymy: Trematosphaeria maquilingiana var. schizostachyi Rehm, Leafl. of Philipp. Bot. 8: 2952 (1916)
Trematosphaeria maquilingiana Rehm, Leafl. of Philipp.
Bot. 8: 2952 (1916) var. maquilingiana
Freshwater distribution: Australia (Vijaykrishna and
Hyde 2006)
Astrosphaeriella papuana Aptroot, Nova Hedwigia 60(3–
4): 333 (1995)
Freshwater distribution: Papua New Guinea (Hyde and
Frohlich 1998), Philippines (Cai et al. 2003a)
Astrosphaeriella seychellensis K.D. Hyde & Goh, S. Afr.
J. Bot. 64: 332 (1998)
Freshwater distribution: Seychelles (Hyde and Goh
1998b)
*Astrosphaeriella stellata (Pat.) Sacc., Syll. fung. (Abellini)
24(2): 938 (1928); Fig. 29
Basionym: Amphisphaeria stellata Pat., Bull. Soc. mycol.
Fr. 29: 223 (1913)
Synonymy: Astrosphaeriella stellata var. palmicola F. San
Martín & P. Lavín, Lavin & Lavín, Acta Bot. Mexicana 46:
22 (1999)
Astrosphaeriella bambusella Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 129: 168 (1920)
Astrosphaeriella fuscomaculans W. Yamam., Sci. Rep.
Hyogo Univ. Agric., Ser. 2, Agr. Biol. 1: 63 (1954)
Microthelia fuscomaculans (W. Yamam.) E. Müll., Beitr.
Kryptfl. Schweiz 11(no. 2): 286 (1962)
Freshwater distribution: China (Tsui et al. 2001d; Wong
and Hyde 2001; Luo et al. 2004; Hu et al. 2013), Philippines
(Cai et al. 2003a)
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Astrosphaeriella thailandensis J. Ren, C.Y. Jie, Y.L. Jiang,
K.D. Hyde & Yong Wang bis, Sydowia 65(1): 33–43 (2013)
Freshwater distribution: Thailand (Ren et al. 2013)
Key to freshwater Astrosphaeriella species
1. Ascospores surrounded by sheath or with appendage……………………………………………………2
1. Ascospores without sheath or appendage………………6
2. Ascospores sometimes slightly verrucose………………
………………………………………A. maquilingiana
2. Ascospores smooth-walled……………………………3
3. Ascospores with minute appendages at both ends…
……………………………………………A. exorrhiza
3. Ascospores sur rounded by a mucilaginous
sheath…………………………………………………4
4. Ascospores pigmented………………………A. stellata
4. Ascospores hyaline……………………………………5
5. Ascospores 30–42 × 7–8 μm, sheath drawn out at the
poles………………………………………A. aquatica
5. Ascospores 16–22 × 4–4.5 μm, sheath appears as remnants in drided material………………A. seychellensis
6. Ascospores with striate ornamentation…………………
……………………………………………A. papuana
6. Ascospores smooth-walled……………A. thailandensis
Caryospora De Not., Micr. Ital. Novi 9: 7 (1855)
Saprobic on submerged wood, decaying fruit pericarp or old
stone of Amygdalus persica. Sexual morph: Ascomata pseudothecioid, medium to large, scattered, erumpent to superficial,
uniloculate, broadly conical or hemispherical, with flattened
base, brown to black, carbonaceous, with ostiolate papilla.
Ostiole relatively broad, central, circular. Peridium thicker at
sides and thinner at base, strongly carbonized. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, sparsely septate.
Asci (2)8-spored, bitunicate, fissitunicate, broadly cylindrical
to clavate or obclavate, pedicellate. Ascospores large, bi- to triseriate, fusiform to broad-ellipsoid or biconic, straight, usually
hyaline when young, becoming pigmented at maturity, septate,
most symmetrical, smooth or rough, often with polar germ
pores at each end, thin- to thick-walled, with or without sheath.
Asexual morph: Undetermined.
Type species: Caryospora putaminum (Schwein.) Fuckel
Notes: Caryospora was placed in Caryosporaceae based
on sequence data of C. aquatica and one putative strain of C.
minima (Ariyawansa et al. 2015). Caryospora clustered with
Acrocordiopsis with weak bootstrap support (Ariyawansa
et al. 2015). Jayasiri et al. (2019) placed Caryospora in
Astrosphaeriellaceae based on the similarity of carbonaceous ascostromata and trabeculate pseudoparaphyses with
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381
Fig. 29 Astrosphaeriella stellata (Material examined: VIETNAM,
Hanoi, on Bamboo (sur Bambou), 17 April 1911, M. Duport 451, FH
00290284, Pat 552, holotype). a Herbarium label and specimen. b
Ascostromata on host surface. c Section through ascostroma. d Sec-
tion through peridium. e Pseudoparaphyses. f–h Bitunicate asci. i
Ocular chamber. j–p Ascospores. Scale bars: d, e, i–p = 20 μm, f–h
= 50 μm
other genera in Astrosphaeriellaceae. However, Astrosphaeriellaceae members, including Caryospora, did not form a
well-supported clade (Jayasiri et al. 2019; this study, Fig. 2).
Zopfia rhizophila Rabenh. (Zopfiaceae) also clustered in
Astrosphaeriellaceae, but with low bootstrap support (Jayasiri et al. 2019; this study, Fig. 2). In this study, we follow
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Jayasiri et al. (2019) and accept Caryospora in Astrosphaeriellaceae based on morphology.
Caryospora obclavata was reported from freshwater
habitat and characterized by obclavate asci with ellipsoidal
to biconic ascospores uniseriately arranged at the apex and
base (Raja and Shearer 2008). The taxonomic placement of
this species needs to be confirmed with molecular data. We
introduce one new species C. submersa based on morphology and phylogeny. Additionally, C. aquatica and C. quercus
are redescribed and reported as new geographical record in
Thailand and new habitat record in freshwater, respectively.
List of freshwater Caryospora species
*Caryospora aquatica H. Zhang, K.D. Hyde & Ariyawansa,
Fungal Diversity 75: 54 (2015); Figs. 30, 31
Freshwater Distribution: China (this study), Thailand
(Ariyawansa et al. 2015)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 270–310 μm high, 250–330 μm
diam., black, scattered or gregarious, superficial, hemisphaerical, flattened at the base, carbonaceous, ostiolate, always
covered with mass of black, glistening ascospores. Peridium
50–60 μm at the sides, poorly developed at the base, easily
cracked, two-layered, outer layer strongly carbonized, amorphous, composed of several layers of black cells that cannot
be differentiated and often occluded, inner layer composed
of several layers of hyaline cells of textura angularis. Pseudoparaphyses 2 μm diam., numerous, trabeculate, filiform,
hyaline, septate. Asci 160–210(–270) × 45–58 μm ( x̄ = 190
× 52 μm, n = 10), 8-spored, bitunicate, narrowly to broadly
clavate, with short pedicel, apically rounded. Ascospores
35–48 × 18–28 μm ( x̄ = 42 × 22 μm, n = 40), biseriate,
straight, broadly fusiform, hyaline to pale brown, 1-septate
when young, becoming irregularly diamond-shaped, dark
brown to black, 3-septate when mature, with rounded ends,
constricted and darker at the median septum, not constricted
and inconspicuous at two terminal septa, asymmetric, with
two large central cells and two small end cells, guttulate,
thick-walled, smooth, sometimes with polar germ pores at
each end, surrounded by an irregular, mucilaginous sheath.
Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 15 mm diam. in 50 days at 25 °C, grey from above,
black from below, surface rough, dry, with dense mycelium,
raised, edge entire.
Material examined: CHINA, Yunnan Province, on submerged wood in a stream, 25 November 2017, G.N. Wang,
H3A-1 (MFLU 18-1202), living culture MFLUCC 18-1030;
ibid., H3A-2 (HKAS 101729), living culture KUMCC
18-0079.
Notes: Our new collection MFLUCC 18-1030 clusters
with Caryospora aquatica H. Zhang et al. (MFLUCC
11-0008) and one putative strain of C. minima Jeffers with
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high bootstrap support (Fig. 2). MFLUCC 18-1030 has
identical LSU sequence data with C. aquatica MFLUCC
11-0008 and seven (including three gaps) nucleotide differences in ITS sequence data between the two isolates, which
indicate them to be the same species. Morphologically, our
collection has very similar morphological characters with
C. aquatica (Figs. 30, 31), except the slightly thinner asci
(45–58 μm wide vs. 60–80 μm wide) and irregular sheaths
(Ariyawansa et al. 2015). Given this, we identify our collection MFLUCC 18-1030 as C. aquatica. This is a new
geographical record of C. aquatica in China.
Caryospora minima was initially collected from putrescent putamina of Amygdalus persica in Maryland, USA (Jeffers 1940). Later, several freshwater collections of C. minima were reported in China, Philippines and South Africa
(see freshwater distribution of this species), but they were
not confirmed by molecular data. Cai and Hyde (2007b)
provided LSU (EU196550) and SSU (EU196551) sequence
data for C. minima without strain code, description, illustration and herbarium information. Thus, these two sequences
are doubtful. A specimen (IFRD 083-010) under name C.
minima is illustrated in this study (Fig. 32). However, this
species should be treated with caution based on the holotype
BPI 71132. It would be good to establish an epitype for C.
minima with reliable sequence data.
*Caryospora minima Jeffers, Mycologia 32(4): 561 (1940);
Fig. 32
Freshwater Distribution: China (Tsui et al. 2000; Luo
et al. 2004), Philippines (Cai et al. 2003a), South Africa
(Hyde et al. 1998)
Caryospora obclavata Raja & Shearer, Mycologia 100(3):
479 (2008)
Freshwater Distribution: USA (Raja and Shearer 2008)
*Caryospora quercus Jayasiri, E.B.G. Jones & K.D. Hyde,
Mycosphere 10(1): 34 (2019); Fig. 33
Freshwater Distribution: Thailand (this study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 400–450 μm high, 410–460
μm diam., black, scattered, semi-immersed to superficial,
conical, flattened at the base, carbonaceous, ostiolate, 70
μm diam., often covered with mass of black, glistening
ascospores. Peridium 60–70 μm at the sides, poorly developed at the base, easily cracked, two-layered, outer layer
strongly carbonized, composed of several layers of black
cells that cannot be differentiated and often occluded, inner
layer composed of 8–10 layers of hyaline, compressed
cells of textura angularis. Pseudoparaphyses 2 μm diam.,
numerous, trabeculate, filiform, hyaline, distantly septate.
Asci (165–)205–270(–305) × (58–)65–80 μm ( x̄ = 235 ×
70 μm, n = 10), 8-spored, bitunicate, narrowly to broadly
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383
Fig. 30 Caryospora aquatica (MFLU 18-1202, new
geographical record). a–c
Ascomata on host surface. d, f
Vertical section of ascomata. e
Structure of peridium. g Pseudoparaphyses. h–k Bitunicate
asci. l–q Immature and mature
ascospores. r, s Ascospores in
Indian Ink. t, u Colony on PDA
(up-front, down-reverse). Scale
bars: d = 100 μm, e–g, l–s = 20
μm, h–k = 40 μm
clavate, indistinctly pedicellate which become tapered and
disappeared with age, 30–60 μm long, apically rounded.
Ascospores 47–60 × 19–33 μm ( x̄ = 54 × 27 μm, n = 30),
overlapping bi- to tri-seriate, straight, broadly fusiform to
ellipsoidal, hyaline to pale brown, 1-septate when young,
becoming irregularly diamond-shaped, dark brown to black,
3-septate when mature, with acute ends, constricted and
darker at the median septum, not constricted and inconspicuous at two terminal septa, asymmetric, with two large
central cells and two small end cells, thick-walled, smooth,
sometimes with polar germ pores at each end, surrounded by
a large, ellipsoidal, gelatinous, mucilaginous sheath, 3–37
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Fig. 31 Caryospora aquatica (Material examined: THAILAND, Chiang Rai Province, on submerged wood, 18 January 2010, H. Zhang,
MFLU 11-1083, holotype). a Ascomata on the host surface. b Sec-
tion of an ascoma on wood. c Section of ascoma. d, e Different stages
of asci. f Ascospores. Scale bars: a = 700 µm, b, c = 350 µm, d, e =
60 µm, f = 50 µm
μm thick, sheath two-layered and large when young, becoming one-layered and smaller after mature. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 5 mm diam. in 20 days at 25 °C, grey from above and
brown from below, surface rough, dry, raised, edge entire.
Material examined: THAILAND, Chiang Mai Province,
on submerged wood in a stream, 1 September 2017, X.D.
Yu, 1A (MFLU 17-1705, holotype), ex-type living culture
MFLUCC 17-2342; ibid., G.N. Wang, 4.2 (MFLU 17-1668),
living culture MFLUCC 17-2323.
Notes: Our new collections MFLUCC 17-2342 and
MFLUCC 17-2323 cluster with Caryospora quercus with
strong bootstrap support (Fig. 2). They have identical LSU
sequence data and there are two nucleotide differences in
SSU sequence data between MFLUCC 17-2342 and MFLU
18-2151 (type). No other genes can be compared.
Our new collections have overlapping ascospore size
(47–60 × 19–33 μm vs. 41–54 × 18–28 μm) and similar
morphological characters with C. quercus, i.e. broadly fusiform to ellipsoidal and hyaline to pale brown when young,
becoming irregularly diamond-shaped and dark brown to
black when mature, with acute ends, with one primary
median septum and two inconspicuous septa near both
ends, asymmetric, and with a sheath (Jayasiri et al. 2019).
However, the ascospore sheaths in our collections are up
to 37 μm thick, while they are 5.5 μm thick in the holotype of C. quercus. This might be a result of only measuring the mature ascospores in Jayasiri et al. (2019), because
they are 37 μm thick in young ascospores and 3 μm thick in
mature ascospores in our collections. Additionally, our collections have longer and wider asci ((165–)205–270(–305)
× (58–)65–80 μm vs. (69–)110–147 × 30–35 μm) than the
holotype of C. quercus. It is very likely that Jayasiri et al.
(2019) only measured the immature asci as shown in their
photo plate. We identify our collections to be C. quercus
based on current phylogenetic analysis (Fig. 2), single gene
comparison and morphology. We supplement ITS sequence
data for C. quercus. This is a new habitat record for C.
quercus from freshwater in Thailand.
13
*Caryospora submersa W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Fungal Diversity (2020) 105:319–575
385
Fig. 32 Caryospora minima
(Material examined: CHINA,
Hong Kong, on submerged
wood in Lam Tsuen River, 1
December 1996, C. Tsui, IFRD
083-010). a Herbarium label
and specimen of Caryospora
minima. b Ascomata on host
surface. c, d Vertical section
through ascoma. e Structure of
peridium. f Pseudoparaphyses.
g, h Bitunicate asci. i Ocular chamber. j Ascospore. k
Ascospore in Indian Ink. Scale
bars: c = 100 μm, e, g, h = 20
μm, f, j–k = 10 μm, i = 5 μm
Index Fungorum number: IF557905; Facesoffungi number: FoF09248; Figs. 34, 35
Etymology: in reference to submerged habitat of the
fungus
Holotype: MFLU 18-1539
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 400–450 μm high, 450–540 μm
diam., black, scattered or gregarious, superficial, conical,
flattened at the base, carbonaceous, with ostiolate papilla.
Peridium 45–85 μm at the sides, poorly developed at the
base, easily cracked, strongly carbonized, composed of
13
386
Fungal Diversity (2020) 105:319–575
Fig. 33 Caryospora quercus
(MFLU 17-1705, new habitat
record). a, b Ascomata on
host surface. c Vertical section of ascoma. d Structure of
peridium. e–g, i–k Bitunicate
asci. h Pseudoparaphyses. l
Immature ascospore, with twolayered sheath (arrows). m–o
Mature ascospores. p Germinated ascospore. q Ascospore in
Indian Ink. r, s Colony on PDA
(left-front, right-reverse). Scale
bars: c = 100 μm, d, h, m–o =
20 μm, e–g, i–l, p, q = 50 μm
several black, amorphous layers of cells that cannot be differentiated and often occluded. Pseudoparaphyses 2 μm
diam., numerous, trabeculate, filiform, hyaline, sparsely
septate. Asci 130–180 × 17–26 μm ( x̄ = 160 × 21 μm, n =
20), 8-spored, bitunicate, narrowly obclavate, with rounded
apex, short pedicellate, with a well-developed ocular chamber. Ascospores 38–50 × 8–12 μm ( x̄ = 44 × 9 μm, n =
20), mostly uniseriate near the apex, becoming biseriate
towards the base, straight or slightly curved, hyaline, 1-septate, constricted at the septum, narrowly fusiform, guttulate,
13
thin-walled, smooth, surrounded by a large, oblong or ellipsoidal, mucilaginous sheath, 40–80 μm wide, sheath becoming thin and dumbbell-like in dried specimen, 1.5–2.5 μm
wide at the sides and 5–15 μm wide at two ends. Asexual
morph: Undetermined.
Culture characteristics: On PDA, colony circular or ellipsoidal, reaching 10 mm diam. in 10 days at 25 °C, grey to
black from above, dark brown from below, surface rough,
dry, raised, edge entire.
Fungal Diversity (2020) 105:319–575
387
Fig. 34 Caryospora submersa (MFLU 18-1539, holotype). a, b
Ascomata on host surface. c Vertical section of ascoma. d Structure
of peridium. e–g Bitunicate asci. h Asci embedded in pseudoparaphyses. i–k Ascospores. l Ascospore from fresh specimen mounted
in Indian Ink. m Germinated ascospore. n Colony on PDA (up-front,
down-reverse). Scale bars: c = 100 μm, d, l, m = 50 μm, e–k = 20
μm
Material examined: THAILAND, Songkhla Province,
Chestnut hill, on submerged wood in a stream, 10 May 2018,
W. Dong, hat458-1 (MFLU 18-1539, holotype), ex-type
living culture MFLUCC 18-1283; ibid., hat458-2 (HKAS
105039, isotype), ex-type living culture KUMCC 19-0056;
Satun Province, on submerged wood in a stream, 10 May
2018, W. Dong, hat286-1 (MFLU 18-1541), living culture
MFLUCC 18-1409; ibid., hat286-2 (HKAS 105070).
Notes: Caryospora submersa is similar to other Caryospora species in having superficial, dark, carbonaceous
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388
Fig. 35 Caryospora submersa (MFLU 18-1541). a, b Ascomata on
host surface. c Vertical section of ascoma. d Vertical section of ostiolate papilla. e Pseudoparaphyses. f–h Bitunicate asci. i, j Ascospores.
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k, l Ascospores from dried specimen mounted in Indian Ink. m Colony on PDA (left-front, right-reverse). Scale bars: c = 100 μm, d =
50 μm, e–l = 20 μm
Fungal Diversity (2020) 105:319–575
ascomata with flattened bases. Caryospora submersa is very
different from other species by obclavate asci and narrowly
fusiform, hyaline, thin-walled ascospores contrasting with
broadly cylindrical to clavate asci and broadly fusiform,
ellipsoidal or biconic, mostly pigmented, relatively thickwalled ascospores (Jeffers 1940; Hawksworth 1982; AbdelWahab and Jones 2000; Raja and Shearer 2008; Zhao and
Zhao 2012; Ariyawansa et al. 2015). Caryospora submersa
does not fit the generic concept of Caryospora except for the
large, superficial, carbonaceous ascomata, which are similar to those in other genera within the family. However, C.
submersa cannot be placed in other genera due to its distinct
morphology and phylogenetically nests in Caryospora with
strong bootstrap support (Fig. 2). Thus, Caryospora submersa sp. nov. is introduced.
Key to freshwater Caryospora species
1.
1.
2.
2.
3.
3.
4.
4.
Ascospores rough-walled…………………C. obclavata
Ascospores smooth-walled……………………………2
Ascospores thin-walled……………………C. submersa
Ascospores thick-walled………………………………3
Ascospores pale brown when mature………C. minima
Ascospores dark brown to black when mature………4
Ascospores 35–48 × 18–28 μm……………C. aquatica
Ascospores 47–60 × 19–33 μm……………C. quercus
Pithomyces Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74):
100 (1873) [1875]
Saprobic in terrestrial or freshwater habitats. Sexual
morph: Ascomata dark opaque, solitary to gregarious,
erumpent to superficial, mammiform to conical, with ruptured, reflexed, stellate, host remnants around the base, uniloculate, glabrous, brittle, carbonaceous, with an indistinct
ostiole. Peridium unequal thickness, poorly developed at the
base, composed of thick, dark brown to black cells. Pseudoparaphyses dense, trabeculate, filiform, hyaline, sparsely
septate. Asci 8-spored, bitunicate, fissitunicate, cylindrical,
with furcate pedicel, apically rounded. Ascospores overlapping biseriate, pale brown to dark brown, fusiform with acute
ends, septate, smooth-walled (Pratibha and Prabhugaonkar
2015; Wanasinghe et al. 2018a). Asexual morph: Hyphomycetous. Conidiophores mononematous, micronematous,
mostly intercalary, sometimes denticulate, aseptate. Conidiogenous cells mono or polyblastic, hyaline to light brown,
smooth. Conidia solitary, brown to black, obovate to oblong,
verruculose to spinulose, transversely septate (Pratibha and
Prabhugaonkar 2015; Wanasinghe et al. 2018a).
Type species: Pithomyces flavus Berk. & Broome
Notes: Pithomyces is similar to Astrosphaeriellaceae
members in having dark, carbonaceous ascomata with a
poorly developed base and fusiform ascospores (Liu et al.
389
2011; Phookamsak et al. 2015b; Wanasinghe et al. 2018a;
Jayasiri et al. 2019). Pithomyces clustered in Astrosphaeriellaceae with low bootstrap support (Jayasiri et al.
2019; this study, Fig. 2), but morphologically belongs to
Astrosphaeriellaceae.
Over 40 epithets are listed in Index Fungorum (2020),
but only three have sequence data, viz. Pithomyces caryotae
Wanas. et al., P. flavus and P. licualae Wanas. et al. Pithomyces flavus was found on the natural substratum associated with its sexual morph Astrosphaeriella vesuvius, which
was confirmed in the phylogenetic analysis (Pratibha and
Prabhugaonkar 2015). Wanasinghe et al. (2018a) found the
asexual morphs of P. caryotae and P. licualae from culture, respectively. The asexual morphs of three species share
similar morpholigical characters in having obovate, septate,
pigmented and rough-walled conidia (Pratibha and Prabhugaonkar 2015; Wanasinghe et al. 2018a). Pithomyces flavus
was recorded from freshwater habitats (Cai et al. 2002a; Hu
et al. 2013), but not confirmed by molecular data.
List of freshwater Pithomyces species
*Pithomyces flavus Berk. & Broome, J. Linn. Soc., Bot.
14(no. 74): 100 (1873) [1875]
Basionym: Sphaeria vesuvius Berk. & Broome, J. Linn.
Soc., Bot. 14(no. 74): 127 (1873) [1875]
Synonymy: Astrosphaeriella vesuvius (Berk. & Broome)
D. Hawksw. & Boise, Sydowia 38: 122 (1986) [1985]
Freshwater distribution: China (Cai et al. 2002a; Hu et al.
2013)
Xenoastrosphaeriella Jayasiri, E.B.G. Jones & K.D. Hyde,
Mycosphere 10(1): 36 (2019)
Saprobic on bamboo and palms in freshwater or terrestrial habitats. Sexual morph: Ascostromata dark opaque,
gregarious, erumpent to superficial, mammiform to conical
with ruptured, reflexed, stellate, host remnants around the
base, uniloculate, glabrous, brittle, carbonaceous, central
ostiolate, with pore-like opening. Peridium unequal thickness, poorly developed at the base, thick at the sides towards
the apex, composed of thick, opaque and melanized cells.
Pseudoparaphyses dense, trabeculate, hyaline, distinctly
septate, anastomosing between the asci, embedded in a
hyaline gelatinous matrix. Asci 8-spored, bitunicate, cylindrical, short pedicellate, apically rounded with an ocular
chamber. Ascospores overlapping uni- to bi-seriate, brown
to reddish brown, with paler end cells, fusiform with acute
ends, septate, smooth-walled (Hawksworth and Boise 1985;
Phookamsak et al. 2015b; Jayasiri et al. 2019). Asexual
morph: Undetermined.
Type species: Xenoastrosphaeriella tornata (Cooke) Jayasiri & K.D. Hyde
Notes: Xenoastrosphaeriella was established by Jayasiri et al. (2019) to accommodate Astrosphaeriella tornata
13
390
(Cooke) D. Hawksw. & Boise which formed a weakly supported clade with other genera in Astrosphaeriellaceae
(Phookamsak et al. 2015b; this study, Fig. 2). Xenoastrosphaeriella is distinct in the family in having paler end cells
of ascospores (Hawksworth and Boise 1985; Phookamsak et al. 2015b; Jayasiri et al. 2019). The second species
Xenoastrosphaeriella trochus (D. Hawksw.) Phookamsak
et al. was recently introduced based on morphology and phylogeny (Hongsanan et al. 2020a). Both species X. tornata
and X. trochus were ever collected from freshwater habitats
(see list below), but not confirmed by molecular data.
List of freshwater Xenoastrosphaeriella species
*Xenoastrosphaeriella tornata (Cooke) Jayasiri & K.D.
Hyde, Mycosphere 10(1): 36 (2019)
Basionym: Trematosphaeria tornata Cooke, Grevillea
16(no. 79): 91 (1888)
Synonymy: Sphaeria tornata Berk. & M.A. Curtis, J.
Acad. nat. Sci. Philad., N.S. 2(6): 290 (1854)
Astrosphaeriella tornata (Cooke) D. Hawksw. & Boise,
Sydowia 38: 119 (1986) [1985]
Freshwater distribution: Philippines (Cai et al. 2003a)
*Xenoastrosphaeriella trochus (D. Hawksw.) Phookamsak,
H.B. Jiang & K.D. Hyde, Mycosphere 11(1): 1762 (2020)
Basionym: Melanomma trochus Penz. & Sacc., Malpighia
11(9–10): 401 (1897)
Synonymy: Leptosphaeria trochus (Penz. & Sacc.) Höhn.,
Sber. Akad. Wiss.Wien, Math.-naturw. Kl., Abt. 1 118: 328
[54 repr.] (1909)
Asterella trochus (Penz. & Sacc.) Hara, Nippon-gaikingaku: 187 (1936)
Asterotheca trochus (Penz. & Sacc.) I. Hino, Bull.
Miyazaki Coll. Agric. Forest.10: 57 (1938)
Trematosphaeria trochus (Penz. & Sacc.) Teng, Sinensia,
Shanghai 9: 257 (1938)
Astrosphaeriella trochus (Penz. & Sacc.) D. Hawksw., J.
Linn. Soc., Bot. 82: 46(1981)
Freshwater distribution: China (Tsui et al. 2000; Cai et al.
2002a, 2006b; Hu etal. 2013), Australia (Vijaykrishna and Hyde
2006), South Africa (Hyde and Frohlich1998; Hyde et al. 1998)
Key to freshwater Xenoastrosphaeriella species
1. Ascospores 3-septate ······································ X. tornata
1. Ascospores mostly 5-septate ·························· X. trochus
Bambusicolaceae D.Q. Dai & K.D. Hyde, Fungal Diversity
63: 49 (2013)
Bambusicola D.Q. Dai & K.D. Hyde, Cryptog. Mycol.
33(3): 367 (2012)
13
Fungal Diversity (2020) 105:319–575
Saprobic on decaying bamboo or parasitic on living
plant materials in terrestrial or freshwater habitats. Sexual
morph: Ascostromata solitary to gregarious, immersed or
semi-immersed and becoming erumpent, variable in shape,
conical, convex or dome-shaped to ampulliform, globose to
subglobose, with centrally located ostiole, with or without red
pigment in the ostiole, uni- to multi- loculate, coriaceous, visible as raised, dark spots or elongate structures on host surface. Peridium composed of dark cells of textura angularis
or textura epidermoidea or pseudoparenchymatous cells, normally with the flattened base comprising of thinner, hyaline,
smaller cells or intermingling with host cells. Pseudoparaphyses dense, trabeculate, filiform, hyaline, indistinctly septate,
anastomosing and branching above the asci. Asci 8-spored,
bitunicate, cylindrical or cylindric-clavate, short pedicellate,
with a shallow or well-developed chamber. Ascospores unito tri-seriate, fusiform, septate, mostly hyaline, rerely pale
brown, mostly surrounded by a gelatinous sheath. Asexual
morph: Coelomycetous. Conidiomata pycnothyrial, solitary
to gregarious, scattered, immersed to semi-immersed, acerose
or subglobose, pyriform or irregular, uni- to multi-loculate.
Pycnidial wall composed of several layers, thin- to thickwalled, hyaline to dark brown cells of textura globulosa or
textura angularis. Conidiophores indistinct. Conidiogenous
cells holoblastic, annellidic, discrete, cylindrical, smooth.
Conidia sometimes two types, macro and microconidia. Macroconidia cylindrical to ellipsoidal, pale brown to brown,
septate, smooth, guttulate. Microconidia globose, oblong to
ellipsoidal, hyaline to pale brown, aseptate, smooth, guttulate.
Type species: Bambusicola massarinia D.Q. Dai & K.D.
Hyde
Notes: Bambusicola is well-studied because all species have
sequence data in GenBank. Bambusicola is distinct in Bambusicolaceae in having fusiform, hyaline, mostly smooth-walled
ascospores and cylindrical to ellipsoidal or globose conidia
(Dai et al. 2012). The asexual morphs can be found in natural
substrate (Bambusicola splendida D.Q. Dai & K.D. Hyde and
B. irregulispora D.Q. Dai & K.D. Hyde) or in culture (B. massarinia D.Q. Dai & K.D. Hyde) (Dai et al. 2012). Bambusicola
sometimes produces two types of conidia, such as B. dimorpha
Thambugala et al. producing macro and micro conidia (Thambugala et al. 2017). Other Bambusicola species only formed
macroconidia characterized as cylindrical to ellipsoidal, pale
brown to brown, septate conidia. Bambusicola species are saprobic on decaying bamboo from Thailand or parasitic on living
leaves or branches of Phyllostachys heteroclada Oliv. from
China (Dai et al. 2012, 2015, 2017; Thambugala et al. 2017;
Yang et al. 2019a). Herein, we report a new Bambusicola species isolated from submerged wood in China.
List of freshwater Bambusicola species
*Bambusicola aquatica W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Fungal Diversity (2020) 105:319–575
391
Fig. 36 Bambusicola aquatica
(MFLU 18-1164, holotype).
a, b Appearance of black
ascomata on host. c, d Vertical
sections of ascomata. e Structure of peridium. f Pseudoparaphyses. g–i Bitunicate asci. j–l
Ascospores. m Germinated
ascospore. n Colony on PDA
(from front). Scale bars: c, d =
100 μm, e = 50 μm, f, j–m = 10
μm, g–i = 20 μm
Index Fungorum number: IF557906; Facesoffungi number: FoF09249; Fig. 36
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-1164
Saprobic on decaying bamboo submerged in freshwater.
Sexual morph: Ascomata 180–210 μm high, 250–370 μm
diam., black, gregarious, densely clustered in a large group,
semi-immersed to superficial, conical or dome-shaped in
section. Peridium comprising host and fungal tissues, laterally 85–95 μm thick, composed of dark brown to black,
thick-walled cells of textura angularis intermingled with host
cells, with thinner basal part 10–35 μm thick, composed of
subhyaline, rim-like cells. Pseudoparaphyses 1.4–1.8 μm
diam., numerous, trabeculate, filiform, hyaline, indistinctly
13
392
septate. Asci 80–126.5(–202) × 7–9.5(–10.5) μm (x̄ = 99.5
× 9 μm, n = 15), 8-spored, bitunicate, cylindrical, apically
rounded, with a shallow chamber and a short, twisted or
straight pedicel, easily and rapidly elongate when mounted
in water, up to 202 μm long. Ascospores 18–22.5 × 4–5 μm
( x̄ = 20.5 × 4.5 μm, n = 40), overlapping biseriate or partially uniseriate, narrowly fusiform, the upper cell slightly
broader than the lower one, sharply narrowed towards to ends
which are narrowly rounded, mostly slightly curved, rarely
straight, hyaline, 1-septate, deeply constricted at the septum,
slightly constricted at a quarter, guttulate, the guttula in upper
cell larger than the lower cell, thin-walled, smooth, with a
thin, hyaline, inconspicuous, mucilaginous sheath which is
2 μm thick when mounted in water, but 5–10 μm thick when
mounted in Indian Ink. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony irregular, reaching 30 mm diam. in 30 days at 25 °C, brown with pale green
margin from above, dark brown from below, surface rough,
with dense mycelium, dry, mucoid after old, edge undulate.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H29A-1 (MFLU 18-1164, holotype), ex-type living
culture MFLUCC 18-1031; ibid., H29A-2 (HKAS 101730,
isotype), ex-type living culture KUMCC 18-0080.
Notes: Bambusicola aquatica is most similar to B. loculata D.Q. Dai & K.D. Hyde in having narrowly fusiform,
1-septate, hyaline ascospores with narrowly rounded ends
and an inconspicuous mucilaginous sheath (Dai et al. 2015).
However, they can be distinguished by their ascus characteristics (cylindric-clavate, bi- to tri-seriate, 80–105 × 8–13
μm in B. loculata vs. cylindrical, overlapping biseriate or
partially uniseriate, 80–126.5(–202) × 7–9.5(–10.5) μm in B.
aquatica). Additionally, the ascomata of B. loculata are solitary or clustered in a small group, while those of B. aquatica
are gregarious and densely clustered in a large group.
Bambusicola aquatica forms a strongly supported clade
with B. irregulispora (Fig. 2). Bambusicola aquatica is an
ascomycetous species, while B. irregulispora is a coelomycetous species. There are respectively 5, 6, 30 and 14 nucleotide differences in the LSU, ITS, TEF and RPB2 sequence
data between B. aquatica (MFLUCC 18-1031) and B. irregulispora (MFLUCC 11-0437), which supports them to be
different species (Jeewon and Hyde 2016).
Nigrogranaceae Jaklitsch & Voglmayr, Stud. Mycol. 85:
54 (2016)
Nigrograna Gruyter, Verkley & Crous, Stud. Mycol. 75: 31
(2012) [2013]
Saprobic on submerged wood, decaying twigs of shrubs
and trees, old fructifications of pyrenomycetes, sometimes as
human pathogenic. Sexual morph: Ascomata solitary to gregarious, immersed to superficial, somewhat globose, brown to
13
Fungal Diversity (2020) 105:319–575
black, ostiolate. Peridium comprising several layers, angular
cells. Pseudoparaphyses numerous, trabeculate, filiform, septate, branching and anastomosing. Asci 8-spored, bitunicate,
cylindrical to clavate, or broadly filiform, with short pedicellate
and knob-like base. Ascospores overlapping, broadly fusiform
to narrowly ellipsoid, inequilateral, septate, hyaline to brown,
smooth or faintly verruculose (Jaklitsch and Voglmayr 2016;
Hyde et al. 2017; Tibpromma et al. 2017). Asexual morph:
Coelomycetous. Pycnidia similar to ascomata, solitary or
rarely grerarious, superficial or submerged in agar, globose to
subglobose or pyriform, olivaceous to olivaceous-black, with
dark brown, septate mycelial outgrowths, with ostiolate papilla.
Peridium comprising several layers of pseudoparenchymatous
cells. Conidiophores filiform, simple to sparsely branched,
with pegs along one or two sides and solitary terminal phialides, reduced in culture. Conidiogenous cells phialidic, discrete,
ampulliform, lageniform or subcylindrical, hyaline. Conidia
oblong, cylindrical or allantoid, sometimes ellipsoid, hyaline or
subhyaline, brown in mass, aseptate, smooth (de Gruyter et al.
2013; Jaklitsch and Voglmayr 2016). Hyphomycetous. Synnemata superficial, effuse, gregarious, black. Mycelium mostly
immersed, composed of septate, pale brown, smooth hyphae.
Conidiophores macronematous, synnematous on substratum,
septate, unbranched to branched, smooth, brown, compacted
below half and flared at the tip. Conidiogenous cells polyblastic,
terminal or intercalary, sympodial, smooth, swollen subcylindrical, ampulliform, denticulate, covered with several conidiogenous loci, hyaline to subhyaline. Conidia solitary, acrogenous,
simple, dry, ellipsoidal, smooth, thin-walled, aseptate, hyaline.
Type species: Nigrograna mackinnonii (Borelli) Gruyter,
Verkley & Crous
Notes: Most Nigrograna species were collected from terrestrial habitats and are morphologically very similar. Sexual
and asexual morphs have been reported from natural substrates
and characterized as ascomycetous and coelomycetous species (Jaklitsch and Voglmayr 2016; Tibpromma et al. 2017).
Only Nigrograna cangshanensis was reported from freshwater
habitats (China), and it differs from other speices by thicker
peridium and molecular characters (Tibpromma et al. 2017).
We collected a hyphomycetous species from submerged wood
in Thailand, which expands the known morph of this genus.
List of freshwater Nigrograna species
*Nigrograna aquatica W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557907; Facesoffungi number: FoF09250; Fig. 37
Etymology: in reference to aquatic habitat of this fungus
Holotype: MFLU 17-1661
Saprobic on submerged wood in freshwater. Sexual morph:
Undetermined. Asexual morph: Synnemata 110–125 ×
8.5–14.5 μm (x̄ = 118 × 11.8 μm, n = 5), superficial, effuse,
gregarious, black, upper part covered with a white or pale
Fungal Diversity (2020) 105:319–575
393
Fig. 37 Nigrograna aquatica
(MFLU 17-1661, holotype). a,
b Colonies on submerged wood.
c–f Conidiophores with attached
conidia. g, h Conidiogenous
cells. i Conidia. j, k Colony on
PDA (up-front, down-reverse).
Scale bars: c, d = 20 μm, e =
40 μm, f, g = 10 μm, h = 3 μm,
i = 5 μm
grey powdery mass of conidia, composed of paralleled conidiophores. Mycelium mostly immersed, composed of septate,
pale brown, smooth hyphae. Conidiophores 100–120 × 1.5–2
μm (x̄ = 100.5 × 1.7 μm, n = 15), macronematous, synnematous on substratum, not compacted, become divergent after
mounted in water, unbranched, septate, not constricted at the
septa, smooth, dark brown at the base, gradually paler towards
the apex, flared in upper part. Conidiogenous cells 4–7.5 ×
1.7–2.5 μm (x̄ = 5.6 × 2.2 μm, n = 10), polyblastic, terminal
or intercalary, integrated when terminal and discrete when
intercalary, sympodial, smooth, swollen subcylindrical, flaskshaped, ampulliform, curved or straight, denticulate, covered
with several conidiogenous loci at the upper part, hyaline to
subhyaline. Conidia 6–7 × 2–2.5 μm (x̄ = 6.6 × 2.2 μm, n =
15), solitary, acrogenous, simple, dry, ellipsoidal or cylindrical,
smooth, thin-walled, aseptate, hyaline.
Culture characteristics: On PDA, colony circular, reaching 25 mm diam. in 30 days at 25 °C, brown to grey from
above, dark brown from below, surface rough, with dense
mycelium, dry, raised, edge entire.
Material examined: THAILAND, Phayao Province, on
submerged wood in a stream, 4 August 2017, G.N. Wang,
G11 (MFLU 17-1661, holotype), ex-type living culture
MFLUCC 17-2318.
Notes: Nigrograna aquatica is morphologically similar to
Phaeoisaria species which are characterized by synnematous conidiophores with polyblastic, sympodial conidiogenous cells (Cai et al. 2006a). However, they cannot be congeneric because N. aquatica differs in having not compacted
synnemata which easily become divergent after mounted in
water. In contrast, the conidiophores of Phaeoisaria are
compacted and parallel adpressed, and never become divergent in water (Liu et al. 2015; Hyde et al. 2018a; Luo et al.
2018). The ellipsoidal or cylindrical, hyaline, aseptate, small
conidia of N. aquatica are similar to those coelomycetous
species in Nigrograna (de Gruyter et al. 2013; Jaklitsch and
Voglmayr 2016). However, phylogenetic analyses showed
13
394
them as distinct species (Fig. 2). The phylogenetically closest species N. locuta-pollinis F. Liu & L. Cai was isolated
from hive-stored pollen collected in Italian honey bee colonies, while N. aquatica was collected from submerged wood
in freshwater habitats in Thailand. Morphology of N. locutapollinis was not given in protologue (Zhao et al. 2018). A
comparison of their nucleotides shows that there are six and
eight (including two gaps) nucleotide differences in LSU and
ITS sequence data, respectively. We introduce N. aquatica
sp. nov. based on the guideline of Jeewon and Hyde (2016)
and this is the first hyphomycetous species in Nigrograna.
*Nigrograna cangshanensis Z.L. Luo, H.Y. Su & K.D.
Hyde, Fungal Diversity 83: 52 (2017)
Freshwater distribution: China (Tibpromma et al. 2017)
Key to freshwater Nigrograna species
1. Sexual morph………………………N. cangshanensis
2. Asexual morph……………………………N. aquatica
Corynesporascaceae Sivan., Mycol. Res. 100(7): 786
(1996)
Corynespora Güssow, Z. PflKrankh. 16: 13 (1906)
Pathogenic and saprobic on leaves or submerged wood. Sexual
morph: Corynesporasca. Ascomata solitary to aggregated, superficial or immersed, uniloculate, globose, dark brown to black, lacking ostioles. Peridium comprising several layers of thin-walled,
brown cells of textura globosa or angularis. Pseudoparaphyses
cylindrical, branched, septate, apically free, disintegrate when
asci mature. Asci 8-spored, bitunicate, obovoid, deliquescent.
Ascospores oblong, pale to dark brown, usually asymmetric,
1-euseptate near the center, with or without indistinctly 1-distoseptate in the upper and lower cell, constricted at the central euseptum
(Sivanesan 1996; Hyde et al. 2013). Asexual morph: Hyphomycetous. Colonies on the natural substrate effuse, hairy, amphigenous, brown. Mycelium superficial to immersed in the substrate,
composed of branched, septate, thin-walled, smooth, subhyaline to
brown hyphae. Conidiophores macronematous, mononematous,
erect to procumbent, simple, none to few septate, smooth. Conidiogenous cells monotretic, integrated, terminal, determinate or
percurrent proliferation. Conidia acrogenous, solitary, obclavate,
wide at the truncate base, tapering towards the apex, pale brown to
pale olivaceous brown, multi-distoseptate, smooth-walled, usually
with a slightly melanized basal scar (Hyde et al. 2013).
Type species: Corynespora cassiicola (Berk. & M.A.
Curtis) C.T. Wei
Notes: Corynespora is characterized by obclavate, brown,
multi-distoseptate conidia with slightly melanized basal scar
(Hyde et al. 2013). Sivanesan (1996) linked C. caryotae
Sivan. as the sexual morph of Corynespora based on cultural
method. Based on the extensive use of the name Corynespora
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Fungal Diversity (2020) 105:319–575
for plant pathogenic fungi and its priority, Rossman et al.
(2015) formally synonymized Corynesporasca Sivan. under
Corynespora. Corynespora is therefore the only genus in
Corynesporascaceae (Sivanesan 1996; Wijayawardene et al.
2018). Corynespora was revealed as polyphyletic (Schoch
et al. 2009; Voglmayr and Jaklitsch 2017). Two freshwater
species C. lignicola Z.L. Luo et al. and C. submersa Z.L. Luo
et al. formed a well-supported clade with the type species C.
cassiicola, which was represented by a strain CBS 100822
(Hyde et al. 2020a). Corynespora aquatica R.F. Castañeda
et al. was reported from freshwater habitats, but not confirmed
by molecular data (Castañeda-Ruiz et al. 2004).
List of freshwater Corynespora species
Corynespora aquatica R.F. Castañeda, Heredia & R.M.
Arias, Mycotaxon 89: 298 (2004)
Freshwater distribution: Mexico (Castañeda-Ruiz et al.
2004)
*Corynespora lignicola Z.L. Luo, H.Y. Su & K.D. Hyde,
Asian Journal of Mycology 3: 65 (2020)
Freshwater distribution: China (Hyde et al. 2020a)
*Corynespora submersa Z.L. Luo, H.Y. Su & K.D. Hyde,
Asian Journal of Mycology 3: 63 (2020)
Freshwater distribution: China (Hyde et al. 2020a)
Key to freshwater Corynespora species
1.
1.
2.
2.
Conidia cylindrical…………………………C. lignicola
Conidia obclavate………………………………………2
Conidia 100–150 μm long…………………C. submersa
Conidia 34–46 μm long……………………C. aquatica
Delitschiaceae M.E. Barr, Mycotaxon 76: 109 (2000)
Delitschia Auersw., Hedwigia 5: 49 (1866)
Saprobic on old herbivore dung or submerged wood.
Sexual morph: Ascomata solitary or scattered, immersed to
semi-immersed, globose to subglobose or subpyriform, often
covered with dense, hyphoid hairs, semitransparent to brown
or black, uniloculate, membranous to coriaceous, with a neck.
Neck blackish, central, usually coriaceous, with a wide opening
and a smooth or tuberculate to hairy surface. Peridium thick
at the apex, thinner at the base. Pseudoparaphyses numerous,
trabeculate, filiform, long, hyaline, septate, anastomosing and
branching. Asci 4- to poly-spored, bitunicate, cylindrical to
cylindro-clavate, short or long pedicellate. Ascospores partially
overlapping, obliquely uniseriate, ellipsoid, reddish brown,
1-septate, cells often easily separable from each other, each
cell with a full length germ slit (Luck-Allen and Cain 1975;
Hyde et al. 2013). Asexual morph: Undetermined.
Fungal Diversity (2020) 105:319–575
Type species: Delitschia didyma Auersw.
Notes: Delitschia is characterized by usually fimicolous habit, with bitunicate asci, and pigmented, 1-septate
ascospores, with an elongate germ slit in each cell (LuckAllen and Cain 1975). Most taxa of this genus are coprophilous (Luck-Allen and Cain 1975), some are wood-inhabiting
(Eaton and Jones 1970; Hyde and Steinke 1996) and three species were reported from freshwater habitats (see list below).
It is reasonable to infer that the lignicolous freshwater taxa
might phylogenetically separate from the coprophilous taxa.
This cannot be confirmed until sequences of three freshwater species are available. Delitschia fasciatispora K.D. Hyde
and D. palmietensis K.D. Hyde & Steinke have 8-spored asci,
while they are 2-spored in D. bispora Eaton & E.B.G. Jones.
List of freshwater Delitschia species
Delitschia bispora Eaton & E.B.G. Jones, Nova Hedwigia
19(3–4): 781 (1971) [1970]
Freshwater distribution: England (Eaton and Jones 1970;
Eaton and Jones 1971; Eaton 1972)
Delitschia fasciatispora K.D. Hyde, Mycoscience 37(1):
100 (1996)
Freshwater distribution: Mauritius (Hyde and Steinke
1996)
Delitschia palmietensis K.D. Hyde & Steinke, Mycoscience
37(1): 101 (1996)
Freshwater distribution: Mauritius (Hyde and Steinke
1996)
Key to freshwater Delitschia species
1. Asci 2-spored………………………………D. bispora
1. Asci 8-spored…………………………………………2
2. Ascospores with a wide dark brown central band………
…………………………………………D. fasciatispora
2. Ascospores lack above characters………D. palmietensis
Dictyosporiaceae Boonmee & K.D. Hyde, Fungal Diversity
80: 462 (2016)
Key to sexual genera of freshwater Dictyosporiaceae
species
1. Ascomata somewhat soft, collapsing when dry, with
hyphomycetous asexual morph………Dictyosporium
1. Ascomata not as above, with coelomycetous asexual
morph………………………………Pseudocoleophoma
395
Key to asexual genera of freshwater Dictyosporiaceae
species
1.
1.
2.
2.
3.
3.
4.
Coelomycetous…………………Pseudocoleophoma
Hyphomycetous………………………………………2
Conidiomata synnemata…………Aquaticheirospora
Conidiomata sporodochia……………………………3
Conidia non-cheiroid…………………Dendryphiella
Conidia cheiroid………………………………………4
Conidiophores form distinct sterile branches…………
………………………………………Cheirosporium
4. Conidiophores with no sterile branches………………5
5. Conidia with divergent arms…………………………6
5. Conidia with compacted arms………………………7
6. Conidiophores composed of moniliform hyphae………
………………………………………Digitodesmium
6. Conidiophores not as above…………………Jalapriya
7. Conidia complanate……………………Dictyosporium
7. Conidia non-complanate……………………………8
8. Conidia with basal hyaline cell……Aquadictyospora
8. Conidia not as above…………………………………9
9. Conidial arms mostly curved towards tip………………
……………………………………Dictyocheirospora
9. Conidial arms mostly parallel………………………10
10. Conidia elongated subcylindrical……………Vikalpa
10. Conidia mostly ellipsoidal……Pseudodictyosporium
Aquadictyospora Z.L. Luo, K.D. Hyde & H.Y. Su, Mycosphere 8(10): 1590 (2017)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Conidiomata on natural substratum
sporodochial, superficial, compact, scattered, circular or subglobose, dark brown to black, velvety. Mycelium immersed,
consisted of septate, hyaline hyphae. Conidiophores reduced
to conidiogenous cells. Conidiogenous cells holoblastic,
monoblastic. Conidia ellipsoidal to broadly clavate, composed of medium brown, appressed cells, rounded and
brown at the apex, oval to subglobose and hyaline at the
base, muriform, cheiroid, non-complanate, without appendages (Li et al. 2017a).
Type species: Aquadictyospora lignicola Z.L. Luo, W.L.
Li, K.D. Hyde & H.Y. Su
Notes: The monotypic genus Aquadictyospora was introduced to accommodate a freshwater species A. lignicola collected from China (Li et al. 2017b; Bao et al. 2018). Aquadictyospora differs from other genera in Dictyosporiaceae
by its basal hyaline cell and conidia without appendages
(Li et al. 2017b; Bao et al. 2018). Aquadictyospora nested
in Dictyosporiaceae with high bootstrap support (Li et al.
2017b; this study, Fig. 47).
List of freshwater Aquadictyospora species
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Fungal Diversity (2020) 105:319–575
Fig. 38 Aquadictyospora
lignicola (Material examined:
CHINA, Yunnan Province, Dali,
on submerged wood in a stream
in Cangshan Mountain, July
2016, H.Y. Su, 4XP H 2–9–3,
MFLU 17–1422, holotype). a–e
Conidia. Scale bars: a = 5 μm,
b–e = 10 μm
*Aquadictyospora lignicola Z.L. Luo, W.L. Li, K.D. Hyde
& H.Y. Su, Mycosphere 8(10): 1591 (2017); Fig. 38
Freshwater distribution: China (Li et al. 2017)
*Aquaticheirospora lignicola Kodsueb & W. H. Ho, J. Linn.
Soc., Bot. 155: 286 (2007); Fig. 39
Freshwater distribution: Thailand (Kodsueb et al. 2007)
Aquaticheirospora Kodsueb & W. H. Ho, J. Linn. Soc., Bot.
155: 286 (2007)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Conidiomata
synnema, scattered or gregarious, erect, brown. Mycelium
immersed in the substratum, hyaline to pale brown. Conidiogenous cells holoblastic, monoblastic, integrated, terminal,
determinate, oblong, hyaline to pale brown. Conidia acrogenous,
solitary, non-complanate, hyaline to brown, cheiroid, euseptate,
arms vertically inserted in different planes, on a basal cell; basal
cells pale brown, cuneiform, smooth, thin-walled; arms discrete,
mostly divergent, cylindrical (Kodsueb et al. 2007).
Type species: Aquaticheirospora lignicola Kodsueb &
W.H. Ho
Notes: The monotypic genus Aquaticheirospora is distinguished from other cheirosporous genera by synnematous
conidiomata and acrogenous, euseptate, cheiroid conidia
with divergent arms (Kodsueb et al. 2007). The synnematous
conidiomata were thought to be not important to delimit a
separated genus because the type species A. lignicola nested
amongst Dictyosporium based on ITS sequence data (Cai
et al. 2008). However, Boonmee et al. (2016) demonstrated
that Aquaticheirospora was a morphologically and phylogenetically well-separated genus based on multigene phylogenetic analyses.
Aquaticheirospora lignicola has 3(–5) tightly packed
arms on the substrate, but has (3–)5(–10) divergent arms
with a brown vacuole in culture (Kodsueb et al. 2007).
Cheirosporium L. Cai & K.D. Hyde, Persoonia 20: 55
(2008)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Conidiomata sporodochial, scattered, punctiform, brown to black. Mycelium immersed or
partly superficial, consisted of branched, septate, subhyaline
to pale brown hyphae. Conidiophores semi-macronematous to
macronematous, mononematous, flexuous, septate, unevenly
branched, branches sterile or fertile; branches with an elongate,
relatively enlarged, obclavate cell. Conidiogenous cells holoblastic, monoblastic, terminal, determinate. Conidia acrogenous, cheiroid, olivaceous to brown, solitary, dry, smoothwalled (Cai et al. 2008).
Type species: Cheirosporium triseriale L. Cai & K.D.
Hyde
Notes: Cheirosporium differs from other genera in Dictyosporiaceae by its semi-macronematous to macronematous,
branched conidiophores with sterile or fertile branches, on
which grow an elongate, relatively enlarged, obclavate cell
(Cai et al. 2008). Two species are listed in Index Fungorum (2020) and both are from freshwater habitats (Cai
et al. 2008). Cheirosporium vesiculare Abdel-Aziz, with
only LSU sequence deposited in GenBank, clustered with
the type species C. triseriale with low bootstrap support in
Abdel-Aziz (2016b) and they are unrelated in our phylogenetic analysis (data not shown). Cheirosporium triseriale has punctiform sporodochia, complanate conidia with
1–2 cells per row, without appendages, while C. vesiculare
has compact sporodochia, non-complanate conidia with
9–12 cells per row and with 2 to 5 large, globose to subglobose appendages (Cai et al. 2008; Abdel-Aziz 2016b).
List of freshwater Aquaticheirospora species
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397
1. Conidia 15–25 × 11–15 µm………………C. triseriale
1. Conidia 42–65 × 19–22 μm………………C. vesiculare
Fig. 39 Aquaticheirospora lignicola (redrawn from Kodsueb et al.
(2007), HKU(M) 17493, holotype). a Conidia produced from synnemata. b Conidium. Scale bars: a = 30 μm, b = 20 μm
The conidial length are also considerably different (see key
below). The phylogenetic relationships of two species need
further study.
List of freshwater Cheirosporium species
*Cheirosporium triseriale L. Cai & K.D. Hyde, Persoonia
20: 56 (2008)
Freshwater distribution: Yunnan, China (Cai et al. 2008)
*Cheirosporium vesiculare Abdel-Aziz, Mycosphere 7(4):
450 (2016)
Freshwater distribution: Egypt (Abdel-Aziz 2016b)
Key to freshwater Cheirosporium species
Dendryphiella Bubák & Ranoj., Annls mycol. 12(4): 417
(1914)
Saprobic on dead branch, living leaves, leaf litter, dead
or old herbaceous stems in terrestrial or submerged wood in
freshwater habitats. Sexual morph: Undetermined. Asexual
morph: Colonies effuse, rubiginous, brown or black, hairy
or velvety. Mycelium mostly immersed. Conidiophores
macronematous, mononematous, evenly or irregularly
branched, with terminal or intercalary nodose swellings, mid
to dark brown or reddish brown, smooth or verruculose. Conidiogenous cells polytretic, integrated, sympodial, reddish
brown to pale brown, subspherical or clavate to subcylindrical, cicatrized, covered with some conidiogenous loci. Conidia
solitary or catenate, acropleurogenous, simple, cylindrical or
oblong, pale brown to mid brown, burnt sienna or olivaceous
brown, septate, smooth or verruculose (Ellis 1971).
Type species: Dendryphiella vinosa (Berk. & M.A. Curtis) Reisinger
Notes: Dendryphiella was introduced by Ranojevic
(1914) with the type species D. interseminata (Berk. &
Ravenel) Bubák which was later synonymized as D. vinosum (Matsushima 1971). Fifteen species are accepted in
the genus and six of which have sequence data (Crous et al.
2014, 2016; Liu et al. 2017c; Hyde et al. 2018a; IturrietaGonzález et al. 2018). Taxonomic notes and a key to Dendryphiella species were provided by Liu et al. (2017c). Dendryphiella has non-cheiroida conidia which are similar to
Neodendryphiella Iturrieta-González et al., but differ in having up to ten conidia in a chain. These two genera separated
well in a phylogenetic tree (Iturrieta-González et al. 2018).
All Dendryphiella species were collected from terrestrial
habitats, except our new collection D. vinosa is collected
from freshwater habitats.
List of freshwater Dendryphiella species
*Dendryphiella vinosa (Berk. & M.A. Curtis) Reisinger,
Bull. trimest. Soc. mycol. Fr. 84(1): 27 (1968)
Facesoffungi number: FoF08673; Fig. 40
Basionym: Helminthosporium vinosum Berk. & M.A.
Curtis [as ‘Helmisporium’], in Berkeley, J. Linn. Soc., Bot.
10(no. 46): 361 (1868) [1869]
Synonymy: Dendryphiella phitsanulokensis N.G. Liu &
K.D. Hyde, Mycosphere 9(2): 287 (2018)
Freshwater Distribution: Thailand (this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies on natural substrate
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Fig. 40 Dendryphiella vinosa (MFLU 17-1666, new habitat
record). a Colonies on submerged wood. b–d Conidiophores with
attached conidia. e Conidiogenous cell and conidium. f–h Conidia
i Two overlapping conidia. j Germinated conidium. k, l Colony on
PDA (left-front, right-reverse). Scale bars: b, d = 50 μm, c, e, i, j =
20 μm, f–h = 10 μm
superficial, effuse, greyish-brown to black. Mycelium partly
immersed, partly superficial, composed of septate, pale
brown, smooth hyphae. Conidiophores uneven in width,
160–300 × 4–6 µm ( x̄ = 235 × 5 µm, n = 10), macronematous, mononematous, occasionally fasciculate, branched at
upper part, slightly wider at nodes, 8–10 µm diam., septate,
slightly constricted at the septa, brown to reddish brown,
sometimes gradually paler and tapering at tip, smooth or
delicately verruculose. Conidiogenous cells 35–40 × 3–5 µm
( x̄ = 37 × 4 µm, n = 5), polytretic, terminal or intercalary,
integrated, proliferating symmetrically or asymmetrically,
smooth, subcylindrical, covered with 1–2 conidiogenous
loci, brown to reddish brown. Conidia 23–34 × 5–8 µm ( x̄ =
27 × 6.5 µm, n = 15), solitary, acrogenous or pleurogenous,
oblong with obtuse ends, 3-septate, slightly constricted at
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the septa, brown to reddish brown, darker at the hilum, simple, dry, smooth or delicately verruculose, thin-walled.
Culture characteristics: On PDA, colony circular, reaching 40 mm in 15 days at 25 °C, pale yellow-brown to white
from above, yellow-brown from below, surface rough, with
dense mycelium in the middle, sparse mycelium in outer
layer, dry, raised, edge entire.
Material examined: THAILAND, Phayao Province, on
submerged wood in a stream, 4 August 2017, G.N. Wang,
G19 (MFLU 17-1666), ex-type living culture MFLUCC
17-2321.
Notes: Dendryphiella vinosa was initially collected from
dead stems of Congo Bean in Cuba (Berkeley and Curtis
1869). Ellis (1971) redescribed and illustrated D. vinosa with
reddish brown, verruculose conidiophores and 3-septate, verruculose, pale brown to reddish brown conidia. Dendryphiella vinosa only has LSU (EU848590) and ITS (DQ307316)
sequence data in GenBank, which have been proved in following phylogenetic analyses (Crous et al. 2016, 2019; Liu
et al. 2017c; Hyde et al. 2018a). Hyde et al. (2018a) introduced a new species D. phitsanulokensis N.G. Liu & K.D.
Hyde, which differs from D. vinosa in having shorter and
unbranched conidiophores. However, they have identical LSU
sequence data and there are only one nucleotide differences in
ITS sequence data between D. phitsanulokensis (MFLUCC
17-2513) and D. vinosa (NBRC 32669). Further morphological comparison shows that D. phitsanulokensis has identical
characters with D. vinosa in all structures (Ellis 1971; Hyde
et al. 2018a), and we consider that the conidiophore length
and branch of D. vinosa can be variable in different living
habitats. Based on single gene comparison, phylogenetic analysis (Fig. 47) and morphological comparison, we synonymize
D. phitsanulokensis with D. vinosa.
Our new collection MFLUCC 17-2321 clusters with D.
vinosa in our multigene phylogenetic analysis (Fig. 47).
They have identical LSU sequence data and there are only
one nucleotide differences in ITS sequence data between
MFLUCC 17-2321 and D. vinosa MFLUCC 17-2513. Our
collection MFLUCC 17-2321 is very similar to D. vinosa
MFLUCC 17-2513 in all morphological characters and
they have overlapping size of conidiophores (160–300 µm
long vs. 130–260 µm long) and conidia (23–34 × 5–8 µm
vs. 16–26 × 2.5–4.4 µm) (Hyde et al. 2018a). We observed
branched conidiophores in our collection, which are present
in Ellis (1971) but absent in Hyde et al. (2018a). We identify
MFLUCC 17-2321 as D. vinosa based on phylogenetic analysis (Fig. 47) and their very similar morphology. This is a
new habitat record of D. vinosa from freshwater in Thailand.
Dictyocheirospora M.J. D’souza, Boonmee & K.D. Hyde,
Fungal Diversity 80: 465 (2016)
399
Saprobic on decaying wood in freshwater and terrestrial
habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Conidiomata on natural substrate sporodochial, punctiform, dark brown. Mycelium immersed,
consisting of branched, septate, subhyaline to pale brown
hyphae. Conidiophores micronematous or semi-macronematous, septate, hyaline to pale brown, smooth. Conidiogenous
cells holoblastic, integrated, terminal, determinate, doliiform
to cylindrical. Conidia acrogenous, solitary, dry, cheiroid,
pale brown, complanate or non-complanate, euseptate or
distoseptate, smooth-walled (Boonmee et al. 2016).
Type species: Dictyocheirospora rotunda M.J. D’souza,
Bhat & K.D. Hyde
Notes: Dictyocheirospora was introduced for the species
producing dark sporodochial colonies with aeroaquatic cheiroid dictyospores (Boonmee et al. 2016). Many new species
and combinations were reported in the genus (Yang et al.
2018), and 21 species are listed in Index Fungorum (2020).
Nine of them are from freshwater habitats, mostly in China
and Thailand (see list below).
List of freshwater Dictyocheirospora species
*Dictyocheirospora aquadulcis Sorvongxay, S. Boonmee
& K.D Hyde, Fungal Diversity 96: 23 (2019)
Freshwater distribution: Thailand (Hyde et al. 2019)
*Dictyocheirospora aquatica Z.L. Luo, Bhat & K.D. Hyde,
Mycosphere 7(9): 1361 (2017)
Freshwater distribution: China (Wang et al. 2016)
*Dictyocheirospora garethjonesii Z.L. Luo, H.Y. Su & K.D.
Hyde, Mycosphere 7(9): 1361 (2017)
Freshwater distribution: China (Wang et al. 2016)
*Dictyocheirospora gigantica (Goh & K.D. Hyde) M.J.
D’souza, Boonmee & K.D. Hyde, Fungal Diversity 80: 469
(2016)
Freshwater distribution: KwaZulu-Natal (Goh et al.
1999)
*Dictyocheirospora indica (I.B. Prasher & R.K. Verma) J.
Yang & K.D. Hyde, MycoKeys 36: 90 (2018)
Freshwater distribution: Thailand (Yang et al. 2018)
*Dictyocheirospora rotunda M.J. D’souza, Bhat & K.D.
Hyde, Fungal Diversity 80: 465 (2016); Fig. 41
Freshwater distribution: China (Wang et al. 2016; this
study), Thailand (Boonmee et al. 2016)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Conidiomata sporodochial, punctiform, velvety, black. Mycelium mostly immersed.
Conidiophores micronematous, reduced. Conidiogenous cells
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Fig. 41 Dictyocheirospora rotunda a–g HKAS 102146. h–k HKAS 102143. a–c Colonies on submerged wood. d–k Conidia. Scale bars: d–k =
20 μm
holoblastic, monoblastic, integrated, terminal, determinate, pale
brown, smooth-walled. Conidia 50–62 × 19–25 µm (x̄ = 56
× 22 µm, n = 15), solitary, acrogenous, cheiroid, pale brown
to dark brown, consisting of 4–6 rows of cells, rows digitate,
cylindrical to narrowly clavate, inwardly curved at the apex,
with a subglobose, cuneiform basal cell, each arm composed
of 10–12 cells, septate, constricted at septa.
Culture characteristics: On PDA, colony circular, reaching 50 mm in 25 days at 25 °C, grey to brown from above,
black from below, surface rough, with dense mycelium,
produced conidia in culture after 10 days, dry, raised, edge
entire.
Material examined: CHINA, Yunnan Province, Kunming University of Science & Technology, on submerged
wood in a stream, 10 May 2017, C.X. Liu, L13 (HKAS
13
102143), living culture KUMCC 19-0103; ibid., Qujing, on
submerged wood in a stream, 10 May 2017, W. Dong, L40
(HKAS 102146), living culture KUMCC 19-0105.
Notes: Our two new collections KUMCC 19-0103 and
KUMCC 19-0105 have slightly thinner conidia (50–62 ×
19–25 µm vs. 42–58 × 19–38 μm) than the holotype in Thailand (Boonmee et al. 2016). Two collections cluster with D.
rotunda with high bootstrap support (Fig. 47).
*Dictyocheirospora tetraploides (L. Cai & K.D. Hyde) J.
Yang & K.D. Hyde, MycoKeys 36: 93 (2018)
Freshwater distribution: China (Cai et al. 2003b)
*Dictyocheirospora thailandica X.D. Yu, W. Dong & H.
Zhang, sp. nov.
Fungal Diversity (2020) 105:319–575
401
Fig. 42 Dictyocheirospora thailandica (MFLU 18-0999, holotype). a Colonies on submerged
wood. b–k Conidia. i Conidiogenous cell with conidium. Scale
bars: b–k = 20 μm
Index Fungorum number: IF557908; Facesoffungi number: FoF09251; Fig. 42
Etymology: referring to Thailand, where the holotype was
collected
Holotype: MFLU 18-0999
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Conidiomata sporodochial, punctiform, black. Mycelium mostly
immersed, composed of pale brown, thin-walled hyphae.
Conidiophores micronematous, reduced. Conidiogenous
cells holoblastic, monoblastic, integrated, terminal, determinate, hyaline to subhyaline, smooth-walled. Conidia
42–65 × 20–45 µm ( x̄ = 51 × 27 µm, n = 15), solitary,
acrogenous, cheiroid, non-complanate, pale brown to dark
brown, consisting of 6–7 rows of cells, rows compacted,
inwardly curved at the apex, easily become divergent and
digitate when in water, broadly clavate, with a subglobose
or cuneiform basal cell, each arm subcylindrical to narrowly
clavate, 7–7.5 µm wide, composed of 9–12 cells, 8–11-septate, constricted at the septa, without appendages or sheaths.
13
402
Fungal Diversity (2020) 105:319–575
Culture characteristics: On PDA, colony circular, reaching 40 mm in 45 days at 25 °C, pale yellow to grey from
above, dark brown from below, surface rough, with dense
mycelium, dry, raised, edge undulate to filiform.
Material examined: THAILAND, Phayao Province, on
submerged wood in a stream, 23 February 2018, X.D. Yu,
Y21 (MFLU 18-0999, holotype), ex-type living culture
MFLUCC 18-0987.
Notes: Dictyocheirospora thailandica clusters with D.
taiwanense Tennakoon et al. with low bootstrap support
(Fig. 47). Morphologically, they are entirely different species. Dictyocheirospora thailandica has broadly clavate
conidia which easily become divergent and digitate when in
water, while D. taiwanense has narrowly cylindrical conidia
comprising of tightly compacted arms (Hyde et al. 2019).
In addition, D. thailandica has shorter and wider conidia
(42–65 × 20–45 µm vs. (72–)74–84(–86) × 16–20(–24) µm)
with a higher number of conidial rows (6–7 vs. 5) than D.
taiwanense (Hyde et al. 2019). Unfortunately, D. taiwanense
only has LSU and ITS sequence data in GenBank, which
show two and three nucleotide differences between D. thailandica (MFLUCC 18-0987) and D. taiwanense (MFLUCC
17-2654), respectively. No other genes can be compared.
Based on the very distinct morphology, we introduce D.
thailandica as a new species in Dictyocheirospora. We
believe that D. thailandica and D. taiwanense will be separated clearly in the phylogenetic tree with protein coding
genes added in future.
*Dictyocheirospora vinaya M.J. D’souza, Bhat & K.D.
Hyde, Fungal Diversity 80: 467 (2016)
Freshwater distribution: Thailand (Boonmee et al. 2016)
Key to freshwater Dictyocheirospora species
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
8.
8.
Conidia with appendage………………………………2
Conidia without appendages……………………………3
Conidial appendage 5–13 × 5–7 μm…………D. indica
Conidial appendage 10–25 × 5–10 μm…………………
…………………………………………D. tetraploides
Conidial rows closely compacted, become divergent when
squashed………………………………………………4
Conidial rows not as above……………………………5
Conidia 58–67 × 15.5–26.5 μm………………D. vinaya
Conidia 42–65 × 20–45 µm……………D. thailandica
Conidial rows composed of > 20 cells……D. gigantica
Conidial rows composed of < 20 cells………………6
Conidia > 60 μm long……………………D. aquadulcis
Conidia < 60 μm long…………………………………7
Conidial arm composed of up to 12 cells……D. rotunda
Conidial arm composed of 7–10 cells…………………8
Conidia 34–42 × 12.5–19.5 μm……………D. aquatica
Conidia 45.5–54.5 × 15.5–24.5 μm……D. garethjonesii
13
Dictyosporium Corda, Weitenweber’s Beitr. Nat.: 87 (1836)
Saprobic on decaying wood and plant debris in terrestrial
and freshwater habitats. Sexual morph: Ascomata perithecial, superficial, solitary or scattered, globose to subglobose,
dark brown to black, collapsed when dry, ostiolate. Peridium
membranaceous, composed of several layers of dark brown,
small cells of textura angularis or textura epidermoidea.
Pseudoparaphyses numerous, cellular, subcylindrical,
hyaline, septate. Asci bitunicate, fissitunicate, clavate to
cylindrical or saccate, pedicellate, apically rounded with an
ocular chamber. Ascospores biseriate, fusiform, elongatedellipsoid, septate, hyaline, with or without mucilaginous
sheath (Boonmee et al. 2016). Asexual morph: Hyphomycetous. Colonies punctiform, sporodochial, effuse or
compact, olive, brown to black, glistening, with immersed
mycelium. Conidiophores micronematous, mononematous,
flexuous, irregularly branched, hyaline to brown, smooth,
sometimes reduced to conidiogenous cells. Conidiogenous
cells holoblastic, monoblastic, integrated, terminal or sometimes intercalary, determinate, cylindrical, doliiform, spherical or subspherical, hyaline to pale brown. Conidia solitary,
dry, acrogenous or sometimes pleurogenous, cheiroid, complanate, olive to brown, smooth, multiseptate (Boonmee
et al. 2016).
Type species: Dictyosporium elegans Corda
Notes: Dictyosporium was introduced for asexual species
producing punctiform, sporodochial colonies and cheiroid,
complanate conidia. The sexual morph D. meiosporum
Boonmee & K.D. Hyde and D. sexualis Boonmee & K.D.
Hyde were reported based on phylogenetic analysis (Liu
et al. 2015). Nineteen freshwater Dictyosporium species are
accepted in the genus, and 11 species are confirmed with
molecular data. Dictyosporium species can be distinguished
by the conidial size, appendage, sheath and rows (see key
below).
List of freshwater Dictyosporium species
*Dictyosporium aquaticum Abdel-Aziz, Fungal Diversity
72: 72 (2015)
Freshwater distribution: Egypt (Liu et al. 2015)
Dictyosporium biseriale D.M. Hu, L. Cai & K.D. Hyde,
Sydowia 62(2): 197 (2010)
Freshwater distribution: Yunnan, China (Hu et al. 2010a)
Dictyosporium canisporum L. Cai & K.D. Hyde, Sydowia
55(2): 130 (2003)
Freshwater distribution: Yunnan, China (Cai et al. 2003b)
*Dictyosporium digitatum J.L. Chen, C.H. Hwang & Tzean,
Mycol. Res. 95(9): 1145 (1991)
Fungal Diversity (2020) 105:319–575
403
Freshwater distribution: Australia (Goh et al. 1999), Brunei (Goh et al. 1999), China (Goh et al. 1999; Tsui et al.
2000, 2001b), Thailand (Sivichai et al. 2002)
Dictyosporium triramosum Aramb., Cabello & Cazau,
Mycotaxon 78: 185 (2001)
Freshwater distribution: Buenos Aires (Arambarri 2001)
Dictyosporium lakefuxianense L. Cai, K.D. Hyde &
McKenzie [as ‘lakefuxianensis’], Cryptog. Mycol. 24(1):
5 (2003)
Freshwater distribution: Yunnan, China (Cai et al. 2003c)
*Dictyosporium tubulatum J. Yang, K.D. Hyde & Z.Y. Liu,
MycoKeys 36: 94 (2018)
Freshwater distribution: Thailand (Yang et al. 2018)
*Dictyosporium nigroapice Goh, W.H. Ho & K.D. Hyde,
in Goh, Hyde & Yanna, Fungal Diversity 103(1): 83 (1999)
Freshwater distribution: Hong Kong, China (Goh et al.
1999; Sivichai et al. 2002; Yang et al. 2018)
*Dictyosporium olivaceosporum Kaz. Tanaka, K. Hiray.,
Boonmee & K.D. Hyde, Fungal Diversity 80: 474 (2016)
Freshwater distribution: Japan (Boonmee et al. 2016)
*Dictyosporium palmae Abdel-Aziz, Mycosphere 7(4):
[453] (2016)
Freshwater distribution: Egypt (Abdel-Aziz 2016b)
Dictyosporium pelagicum (Linder) G.C. Hughes ex E.B.G.
Jones [as ‘pelagica’], Trans. Br. mycol. Soc. 46(1): 137
(1963)
Freshwater distribution: USA (Shearer 1972)
Dictyosporium polystichum (Höhn.) Damon, Lloydia 15:
118 (1952)
Freshwater distribution: Yunnan, China (Luo et al. 2004)
*Dictyosporium stellatum G.P. White & Seifert, Persoonia
26: 156 (2011)
Freshwater distribution: Ontario (Crous et al. 2011)
Dictyosporium tetraseriale Goh, Yanna & K.D. Hyde, Fungal Diversity 103(1): 87 (1999)
Freshwater distribution: Brunei (Goh et al. 1999), Hong
Kong, China (Goh et al. 1999)
*Dictyosporium tetrasporum L. Cai & K.D. Hyde, Mycoscience 48(5): 290 (2007)
Freshwater distribution: Yunnan, China (Cai and Hyde
2007a)
*Dictyosporium thailandicum M.J. D’souza, Bhat & K.D.
Hyde, Fungal Diversity 72: 78 (2015)
Freshwater distribution: Thailand (Liu et al. 2015)
*Dictyosporium tratense J. Yang & K.D. Hyde, MycoKeys
36: 96 (2018)
Freshwater distribution: Thailand (Yang et al. 2018)
Dictyosporium yunnanense L. Cai, K.D. Hyde & McKenzie
[as ‘yunnanensis’], Cryptog. Mycol. 24(1): 7 (2003)
Freshwater distribution: Yunnan, China (Cai et al. 2003c)
*Dictyosporium zhejiangense Wongs., H.K. Wang, K.D.
Hyde & F.C. Lin, Cryptog. Mycol. 30(4): 358 (2009)
Freshwater distribution: Zhejiang, China (Wongsawas
et al. 2009)
Key to freshwater Dictyosporium species (only species
confirmed with molecular data)
1.
1.
2.
Conidia without appendages…………………………2
Conidia with appendage………………………………3
Conidia (50–)95–140(–175) μm × (27.5–)30–40(–
52.5) μm…………………………………D. stellatum
2. Conidia 23.5–40 μm × 16–21.5 μm……………………
………………………………………D. tetrasporum
3. Conidia with sheath…………………………………4
3. Conidia without sheath………………………………5
4. Conidia 60–85 μm × 20–30 μm………D. aquaticum
4. Conidia (40–)43–54(–57) μm × (20–)23–32(–36)
μm…………………………………………D. tratense
5. Conidia with two types of appendages………………
……………………………………………D. palmae
5. Conidia with one type of appendage…………………6
6. Conidia > 45 μm long……………………D. digitatum
6. Conidia < 45 μm long…………………………………7
7. Conidia not complanate…………D. olivaceosporum
7. Conidia complanate…………………………………8
8. Conidia consist of five rows of cells…………………9
8. Conidia mostly consist of four rows of cells………10
9. Appendages variable in shape………D. zhejiangense
9. Appendages tubular…………………D. thailandicum
10. Conidial appendage 22–34 × 4–5 μm…D. nigroapice
10. Conidial appendage 19–24 × 3.5–7 μm………………
…………………………………………D. tubulatum
Digitodesmium P. M. Kirk, Trans. Br. mycol. Soc. 77: 284
(1981)
Saprobic on submerged bamboo, wood in freshwater or
decaying wood, soil in terrestrial habitats. Sexual morph:
Undetermined. Asexual morph: Hyphomycetous. Conidiomata sporodochial, punctiform, pulvinate, scattered, pale
13
404
Fungal Diversity (2020) 105:319–575
brown to brown. Mycelium mostly immersed, composed of
branched, septate, thin-walled, smooth, pale brown hyphae.
Conidiophores semi-macronematous, mononematous, fasciculate, composed of moniliform, pale brown, septate, smooth
hyphae. Conidiogenous cells holoblastic, monoblastic, integrated, terminal, determinate, globose to doliiform, minutely
cicatrized. Conidia acrogenous, solitary, dry, digitate, cheiroid, slightly divergent arms, euseptate, some with a hyaline
gelatinous cap at the apex (Kirk 1981; Boonmee et al. 2016).
Type species: Digitodesmium elegans P.M. Kirk
Notes: Digitodesmium is unique by its digitate conidia
and moniliform conidiophores (Kirk 1981; Boonmee et al.
2016). Tsui et al. (2006a) indicated that Digitodesmium
might be a synonym of Dictyosporium because the phylogenetic position of Digitodesmium was within Dictyosporium,
but with low bootstrap support. A similar result was shown
in Cai et al. (2008). Digitodesmium was distinguished from
Dictyosporium by its divergent conidial arms which were
considered as an artificial feature by Tsui et al. (2006a) and
Cai et al. (2008). Boonmee et al. (2016) introduced family Dictyosporiaceae to accommodate most cheirosporous
hyphomycete genera and accepted Digitodesmium as a distinct genus based on multigene phylogenetic analyses. We
think that the moniliform conidiophores of Digitodesmium
are also important to distinguish this genus.
Three freshwater Digitodesmium species morphologically fit well with the generic concept of Digitodesmium in
producing divergent conidial arms (Ho et al. 1999; Cai et al.
2002b, 2003b). The placement of D. heptasporum L. Cai
& K. D. Hyde and D. recurvum W. H. Ho et al. needs to be
confirmed with molecular data.
List of freshwater Digitodesmium species
*Digitodesmium bambusicola L. Cai, K. Q. Zhang, McKenzie,
W. H. Ho & K. D. Hyde, Nova Hedwigia 75: 528 (2002); Fig. 43d
Freshwater distribution: Philippines (Cai et al. 2002b)
Digitodesmium heptasporum L. Cai & K. D. Hyde,
Sydowia 55: 133 (2003)
Freshwater distribution: Yunnan, China (Cai et al. 2003b)
Digitodesmium recurvum W. H. Ho, K. D. Hyde & Hodgkiss, Mycologia 91: 900 (1999); Fig. 43a–c
Freshwater distribution: China (Ho et al. 1999)
Key to freshwater Digitodesmium species
1.
1.
2.
2.
Conidia with appendage………………D. bambusicola
Conidia without appendages…………………………2
Conidia 30–45 × 12–21 μm………………D. recurvum
Conidia 50–75 × 32.5–70 μm…………D. heptasporum
13
Fig. 43 Digitodesmium spp. a–c D. recurvum (redrawn from Ho
et al. (1999), HKU(M) 4552, holotype). a Conidiophore bearing
conidium. b Conidium. c Conidium at the basal portion. d D. bambusicola (redrawn from Cai et al. (2002b), PDD 74494, holotype).
d Conidium with apical or subapical appendages. Scale bars: a–d =
10 μm
Pseudocoleophoma Kaz. Tanaka & K. Hiray., Stud. Mycol.
82: 89 (2015)
Saprobic on decaying pod, leaves, stems and submerged
wood. Sexual morph: Ascomata scattered or gregarious,
immersed to erumpent, globose to subglobose, ostiolate.
Ostiolar neck central, composed of subglobose, dark brown
cells. Peridium composed of several layers of polygonal
to rectangular cells. Pseudoparaphyses numerous, cellular, hypha-like, septate, branched, anastomosing. Asci
8-spored, bitunicate, fissitunicate, cylindrical to clavate,
short pedicellate. Ascospores fusiform, septate, smooth, with
a conspicuous, mucilaginous sheath (Tanaka et al. 2015).
Asexual morph: Coelomycetous. Conidiomata immersed
to erumpent, subglobose. Ostiolar neck well-developed,
cylindrical, central. Peridium composed of several layers
of slightly thick-walled, pale brown, polygonal to subglobose cells. Conidiophores absent. Conidiogenous cells
Fungal Diversity (2020) 105:319–575
enteroblastic, phialidic, doliiform to lageniform. Conidia
cylindrical, hyaline, aseptate or septate, smooth (Tanaka
et al. 2015).
Type species: Pseudocoleophoma calamagrostidis Kaz.
Tanaka & K. Hiray.
Notes: Pseudocoleophoma is characterized by fusiform, septate ascospores with a conspicuous sheath and
coleophoma-like conidia (Tanaka et al. 2015). Pseudocoleophoma was placed in Dictyosporiaceae based on molecular data (Tanaka et al. 2015; Boonmee et al. 2016; Yang et al.
2018). Four species are listed in Index Fungorum (2020).
Pseudocoleophoma typhicola Kamolhan et al. (Hyde et al.
2016b) is known from only an asexual morph, while the
other species have both sexual and asexual morphs. Pseudocoleophoma typhicola is the only species producing septate
conidia (Tanaka et al. 2015; Hyde et al. 2016b; Jayasiri et al.
2019).
List of freshwater Pseudocoleophoma species
*Pseudocoleophoma typhicola Kamolhan, Banmai, Boonmee, E.B.G. Jones & K.D. Hyde, Fungal Diversity 80: 34
(2016)
Freshwater distribution: UK (Hyde et al. 2016b)
Jalapriya M.J. D’souza, H.Y. Su, Z.L. Luo & K.D. Hyde,
Fungal Diversity 80: 476 (2016)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Colonies effuse, dark brown to black.
Mycelium immersed or partly superficial, consisting of
branched, septate, smooth, subhyaline to pale brown hyphae.
Conidiophores micronematous, unbranched, thin-walled,
smooth, cylindrical. Conidiogenous cells holoblastic, integrated, determinate, terminal. Conidia acrogenous, solitary,
cheiroid, complanate or non-complanate, smooth-walled,
euseptate, with 5–7 rows of cells, rows converging or divergent at the apex, apical cells with or without appendages
(Boonmee et al. 2016).
Type species: Jalapriya pulchra M.J. D’souza, H.Y. Su,
Z.L. Luo & K.D. Hyde
Notes: Jalapriya was introduced to accommodate a new
freshwater species J. pulchra and two new combinations of
species from terrestrial habitats (Boonmee et al. 2016). The
cheirosporous genera are morphologically very similar in
the family and Jalapriya was established based on multigene phylogenetic analyses (Boonmee et al. 2016). Jalapriya
pulchra is distinctive based on its complanate conidia with
hyaline appendages on the apical cells.
List of freshwater Jalapriya species
*Jalapriya pulchra M.J. D’souza, H.Y. Su, Z.L. Luo & K.D.
Hyde, Fungal Diversity 80: 477 (2016); Fig. 44
405
Freshwater distribution: China (Boonmee et al. 2016)
Pseudodictyosporium Matsush., Bull. natn. Sci. Mus.,
Tokyo, N.S. 14: 473 (1971)
Saprobic on rotten leaves or submerged wood. Sexual
morph: Undetermined. Asexual morph: Hyphomycetous.
Conidiomata on natural substratum sporodochial, superficial, punctiform to effuse, velvety, scattered to gregarious,
pale brown to dark brown, olive. Conidiophores micronematous, aseptate, simple, hyaline to pale brown, smooth. Conidiogenous cells holoblastic, monoblastic, integrated, terminal,
determinate, doliiform to cylindrical. Conidia acrogenous,
solitary, dry, cheiroid, pale brown, smooth-walled, septate,
composed of three rows of cells arising parallelly from a
truncate basal cell, compactly, non-complanate, with or
without appendages (Matsushima 1971; Li et al. 2017).
Type species: Pseudodictyosporium wauense Matsush.
Notes: Pseudodictyosporium wauense is a terrestrial species collected from New Guinea. It was re-collected from
submerged wood in China and described as cheiroid conidia
containing three parallel rows of cells arising from a basal
cell (Li et al. 2017). The parallel rows of cells of Pseudodictyosporium are similar to Vikalpa, however, phylogeny
separates them as distinct genera (Boonmee et al. 2016; Li
et al. 2017; Yang et al. 2018).
List of freshwater Pseudodictyosporium species
*Pseudodictyosporium wauense Matsush., Bull. natn. Sci.
Mus., Tokyo 14(3): 473 (1971); Fig. 45
Freshwater distribution: China (Li et al. 2017)
Vikalpa M.J. D’souza, Boonmee, Bhat & K.D. Hyde, Fungal
Diversity 80: 479 (2016)
Saprobic on submerged wood in freshwater and decaying
wood, leaves in terrestrial habitats. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Conidiomata
sporodochial, punctiform or effuse, scattered to gregarious,
superficial, pale brown to dark brown, with or without a
mucilage, inconspicuous covering. Mycelium immersed,
composed of branched, septate, subhyaline to pale brown,
smooth-walled hyphae. Conidiophores micronematous,
aseptate, hyaline to pale brown. Conidiogenous cells holoblastic, integrated, terminal, determinate, cylindrical to
doliiform. Conidia acrogenous, solitary, dry, cheiroid, pale
brown, smooth-walled, euseptate or distoseptate, consisting of three rows of cells arised parallel on a truncate basal
cell, closely compacted, non-complanate, with or without
appendages (Boonmee et al. 2016).
Type species: Vikalpa australiensis (B. Sutton) M.J.
D’souza, Boonmee & K.D. Hyde
Notes: Vikalpa was introduced for one new species and
three new combinations transferred from Dictyosporium
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406
Fungal Diversity (2020) 105:319–575
Fig. 44 Jalapriya pulchra
(Material examined: CHINA,
Yunnan Province, Dali,
Cangshan Mountain, Lingquan
stream, on submerged wood,
March 2014, Z. Li, LQXM
47–1(S-013), HKAS 83979,
holotype). a Colonies on
submerged wood. b–f Conidia.
Scale bars: b = 10 μm, c–f =
20 μm
(Boonmee et al. 2016). Vikalpa has non-complanate conidia
consisting of three rows of cells arising parallel on a truncate
basal cell, while Dictyosporium has complanate conidia with
several unparallel rows of cells (Boonmee et al. 2016). The
placement of Vikalpa was unstable, as it clustered basal to
Dictyosporium in Boonmee et al. (2016), but distant from
Dictyosporium (Yang et al. 2018). The freshwater species
V. lignicola M.J. D’souza et al. was introduced based on its
3 rows and non-complanate conidia, but without molecular
evidence (Boonmee et al. 2016).
List of freshwater Vikalpa species
Vikalpa lignicola M.J. D’souza, Bhat, H.Y. Su & K.D.
Hyde, Fungal Diversity 80: 479 (2016); Fig. 46
Freshwater distribution: China (Boonmee et al. 2016)
Didymosphaeriaceae Munk, Dansk bot. Ark. 15(no. 2): 128
(1953)
Key to freshwater genera of Didymosphaeriaceae
1. Ascomata with a pore-like opening…Didymosphaeria
1. Ascomata without above character……………………2
2. Asci cylindrical, ascospores broadly ellipsoidal, yellowish brown…………………………Paraphaeosphaeria
13
2. Asci elongate-clavate to sub-cylindrical, ascospores
fusoid, hyaline or pale brown……………Pseudotrichia
Didymosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 140
(1870) [1869-70]
Saprobic on woody branches and herbaceous stems and
leaves in freshwater or terrestrial habitats. Sexual morph:
Ascomata solitary, scattered, or in small groups, immersed
to erumpent, globose to ovoid, with ostiolate papilla. Ostiole
with a pore-like opening, periphysate. Peridium one-layered,
thin, composed of brown pseudoparenchymatous cells of
textura intricata. Pseudoparaphyses dense, trabeculate,
filiform, hyaline, indistinctly septate, anastomosing, mostly
above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical, with a furcated pedicel, apically rounded with an
indistinct ocular chamber. Ascospores uniseriate, ellipsoidal, brown, 1-septate (Zhang et al. 2012b; Ariyawansa et al.
2014a). Asexual morph: Undetermined.
Type species: Didymosphaeria futilis (Berk. & Broome)
Rehm
Notes: Didymosphaeria comprises over 520 epithets in
Index Fungorum (2020). The type species D. futilis was
examined by Ariyawansa et al. (2014a) who also introduced a new species D. rubi-ulmifolii Ariyaw. et al. based
on immersed ascomata under clypeus with a single layered
peridium and brown, 1-septate ascospores. Didymosphaeria
represented by D. rubi-ulmifolii clustered with members of
Fungal Diversity (2020) 105:319–575
407
Fig. 45 Pseudodictyosporium
wauense (Material examined:
CHINA, Yunnan Province,
Dali, saprobic on decaying
wood submerged in a stream in
Cangshan Mountain, June 2016,
S.M. Tang, 3XP H 3–9–1, DLU
0801). a–f Conidia. Scale bars:
a–f = 10 μm
Didymosphaeriaceae (Liu et al. 2017a; this study, Fig. 2).
The placement of Didymosphaeria futilis is questionable as
two strains clustered in two different families (HKUCC 5834
in Cucurbitariaceae and CMW22186 in Didymellaceae)
(Zhang et al. 2012b; Ariyawansa et al. 2014a). The freshwater species D. pittospora Udaiyan & Hosag. was proposed
by Udaiyan and Hosagoudar (1991) but is an invalid name
(Index Fungorum 2020) because the holotype was not indicated when introduced (Fig. 47).
List of freshwater Didymosphaeria species
Didymosphaeria pittospora Udaiyan & Hosag., J. Econ.
Taxon. Bot. 15(3): 658 (1992) [1991]
Freshwater distribution: Tamil Nadu, India (Udaiyan and
Hosagoudar 1991)
Paraphaeosphaeria O.E. Erikss., Ark. Bot., Ser. 2 6: 405
(1967)
Saprobic on various hosts in terrestrial and submerged wood
in freshwater habitats. Sexual morph: Ascomata immersed
to semi-immersed, depressed globose, with ostiolate papilla.
Ostiole black, with a short neck, without periphyses. Peridium
composed of several layers cells of textura prismatica. Pseudoparaphyses dense, trabeculate, filamentous, hyaline, septate.
Asci 8-spored, bitunicate, fissitunicate, cylindrical with a short,
broad pedicel. Ascospores uniseritate or partially overlapping,
broadly ellipsoidal, yellowish-brown, septate, smooth, with a
thin sheath (Ariyawansa et al. 2014b). Asexual morph: Coelomycetous. Conidiomata eustromatic or pycnidial. Conidiogenous cells discrete or integrated, phialidic or annellidic, with
percurrent proliferations. Conidia variable in shape, subglobose, ellipsoid to obovoid-pyriform, or more cylindrical, aseptate or 1-septate, smooth to verrucose (Verkley et al. 2014).
Type species: Paraphaeosphaeria michotii (Westend.)
O.E. Erikss.
Notes: Paraphaeosphaeria was confirmed in Didymosphaeriaceae based on molecular data derived from epitype of
P. michotii (Ariyawansa et al. 2014b). The asexual morph of
Paraphaeosphaeria was reported as coelomycetous species,
e.g. P. angularis Verkley & Aa, P. arecacearum Verkley
et al. and P. neglecta Verkley et al. (Verkley et al. 2014).
Paraphaeosphaeria is quite comparable to Phaeosphaeria I.
Miyake, but phylogeny places them in Didymosphaeriaceae
and Phaeosphaeriaceae, respectively (Liu et al. 2017a).
Two freshwater species, Paraphaeosphaeria michotii and
P. schoenoplecti M.K.M. Wong et al., are similar in having
2-septate, yellowish brown ascospores, but can be distinguished by ascospores shape (cylindrical vs. long fusiform)
and size (see key below) (Wong et al. 2000).
List of freshwater Paraphaeosphaeria species
13
408
Fungal Diversity (2020) 105:319–575
Fig. 46 Vikalpa lignicola (Material examined: CHINA, Yunnan
Province, Dali, Cangshan Mountain, Xue Shan Stream, on decaying
wood submerged in the stream, 11 August 2014, Z.L. Luo, XS-69-2,
MFLU 15-1506, holotype). a, b Colonies on submerged wood. c–g
Conidia. Scale bars: c, g = 15 μm, d–f = 10 μm
Paraphaeosphaeria michotii (Westend.) O.E. Erikss., Arch.
Botan. 6: 405 (1967)
Basionym: Sphaeria michotii Westend., Bull. Acad. R.
Sci. Belg., Cl. Sci., sér. 2 7(5): 87 (1859)
Synonymy: see Index Fungorum (2020)
Freshwater distribution: England (Pugh and Mulder
1971)
Key to freshwater Paraphaeosphaeria species
Paraphaeosphaeria schoenoplecti M.K.M. Wong, Goh &
K.D. Hyde, Fungal Diversity 4: 172 (2000)
Freshwater distribution: Hong Kong, China (Wong et al.
2000)
13
1. Ascospores (12–)16–24 × 4–5(–6) μm………P. michotii
1. Ascospores 27–31 × 3.8–4 μm………P. schoenoplecti
Pseudotrichia Kirschst., Annls mycol. 37(1/2): 125 (1939)
Saprobic in terrestrial or rarely aquatic habitats. Sexual
morph: Ascomata solitary or scattered, immersed to nearly
superficial, globose to pyriform, carbonaceous or coriaceous, with ostiolate papilla. Peridium two-layered, comprising thick-walled cells of textura angularis, outer layer
heavily pigmented, comprising small, dark brown to black
cells, inner layer comprising lightly pigmented or hyaline
Fungal Diversity (2020) 105:319–575
Fig. 47 Phylogram generated
from maximum likelihood analysis of combined ITS, LSU and
TEF sequence data for species
of Dictyosporiaceae. Bootstrap
values for maximum likelihood
equal to or greater than 75% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree is
rooted to Lentithecium clioninum KT 1220 (Lentitheciaceae)
409
Dictyocheirospora indica MFLUCC 15-0056 F
99/1.00
Dictyocheirospora subramanianii BCC 3503
Dictyocheirospora xishuangbannaensis MFLUCC 17-2267
Dictyocheirospora aquadulcis MFLUCC 17-2571 F
89/0.99
Dictyocheirospora heptaspora CBS 396.59
Dictyocheirospora lithocarpi MFLUCC 17-2537
100/1.00
Dictyocheirospora bannica KH 332
Dictyocheirospora bannica MFLUCC 16-0874
Dictyocheirospora pseudomusae KH 412
95/1.00
Dictyocheirospora pandanicola MFLUCC 16-0365
100/1.00
Dictyocheirospora vinaya MFLUCC 14-0294 F
90/0.99
Dictyocheirospora pseudomusae yone 234
Dictyocheirospora gigantica BCC 11346
100/1.00
Dictyocheirospora metroxylonis MFLUCC 15-0282a
100/1.00
Dictyocheirospora metroxylonis MFLUCC 15-0282b
Dictyocheirospora
Dictyocheirospora thailandica MFLUCC 18-0987 F
Dictyocheirospora taiwanense MFLUCC 17-2654
88/0.97
Dictyocheirospora rotunda KUMCC 19-0103 F
Dictyocheirospora rotunda MFLUCC 17-1313 F
Dictyocheirospora rotunda KUMCC 19-0105 F
99/1.00
Dictyocheirospora rotunda MFLUCC 14-0222 F
92/1.00
Dictyocheirospora rotunda MFLUCC 14-0293 F
98/1.00
Dictyocheirospora rotunda DLUCC 0576 F
Dictyocheirospora cheirospora MFLUCC 17-0888
77/1.00
Dictyocheirospora garethjonesii MFLUCC 16-0909 F
92/1.00
--/0.99
100/1.00
Dictyocheirospora garethjonesii DLUCC 0848 F
Dictyocheirospora garethjonesii KUMCC 15-0396 F
100/1.00
Dictyocheirospora nabanheensis MFLUCC 17-0562
Dictyocheirospora aquatica KUMCC 15-0305 F
Digitodesmium bambusicola CBS 110279 F
Aquaticheirospora lignicola HKUCC 10304 F
89/0.95
100/1.00
93/1.00
Digitodesmium
Aquaticheirospora
Jalapriya pulchra MFLUCC 15-0348 F
Jalapriya pulchra MFLUCC 17-1683 F
Jalapriya inflata NTOU 3855
Jalapriya
Jalapriya toruloides CBS 209.65
Vikalpa australiensis HKUCC 8797
Aquadictyospora lignicola MFLUCC 17-1318 F
cells. Pseudoparaphyses numerous, cellular, narrow, hyaline, septate, unbranched. Asci 8-spored, bitunicate, fissitunicate, elongate clavate to sub-cylindrical, with narrow, long,
furcate pedicellate, apically rounded with a minute ocular
chamber. Ascospores uni- or bi-seriate, fusoid, straight to
curved, hyaline or pale brown, 1- to multiseptate, smoothwalled, thick-walled (Thambugala et al. 2014; Tian et al.
2015). Asexual morph: Undetermined.
Type species: Pseudotrichia stromatophila Kirschst.
Notes: Pseudotrichia was placed in Didymosphaeriaceae
by Thambugala et al. (2014) based on re-examination of the
holotype of P. stromatophila. Pseudotrichia was polyphyletic
as shown in Tian et al. (2015) as one strain of P. guatopoensis
Vikalpa
Aquadictyospora
Huhndorf formed an independent lineage in Pleosporales and
two strains of P. mutabilis (Pers.) Wehm. clustered in Pleomassariaceae. The sequence data of the type species P. stromatophila and other Pseudotrichia species are needed to clarify
the familial and generic status (Thambugala et al. 2014). Pseudotrichia allequashensis Fallah & Shearer differs from other
species in the genus in having a narrow cylindric neck with a
clypeus and hyaline ascospores with three eusepta and two distosepta (Huhndorf 1994; Fallah and Shearer 2001). However,
the placement of P. allequashensis in Didymosphaeriaceae
needs confirmation with molecular data.
List of freshwater Pseudotrichia species
13
410
Fungal Diversity (2020) 105:319–575
Fig. 47 (continued)
Dictyosporium thailandicum MFLUCC 13-0773 F
Dictyosporium alatum ATCC 34953
Dictyosporium strelitziae CBS 123359
100/1.00
Dictyosporium bulbosum yone 221
93/1.00
Dictyosporium elegans NBRC 32502
Dictyosporium hughesii KT 1847
Dictyosporium zhejiangensis MW 2009a F
95/0.99
95/1.00
Dictyosporium sp. MFLUCC 15-0629 F
Dictyosporium strelitziae CCFC 241241
Dictyosporium aquaticum MF1318 F
93/0.97
Dictyosporium digitatum yone 280
Dictyosporium
Dictyosporium digitatum KT 2660
Dictyosporium wuyiense CGMCC 3.18703
Dictyosporium tratense MFLUCC 17-2052 F
100/1.00
Dictyosporium tubulatum MFLUCC 17-2056 F
100/1.00
Dictyosporium tubulatum MFLUCC 15-0631 F
100/0.99
Dictyosporium nigroapice MFLUCC 17-2053 F
92/1.00
Dictyosporium nigroapice BCC 3555
99/--
Dictyosporium meiosporum MFLUCC 10-0131
Dictyosporium tetrasporum KT 2865
Dictyosporium olivaceosporum KH 375 F
Dictyosporium sexualis MFLUCC 10-0127
97/0.99
Pseudodictyosporium wauense DLUCC 0801 F
79/--
Pseudodictyosporium wauense KRP88 6
94/-100/1.00
Pseudodictyosporium wauense NBRC 30078
Pseudodictyosporium indicum CBS 471.95
100/1.00
Pseudodictyosporium
Pseudodictyosporium elegans CBS 688.93
Pseudodictyosporium thailandica MFLUCC 16-0029
Cheirosporium triseriale HMAS 180703 F
96/1.00
Cheirosporium
Dendryphiella paravinosa CPC 26176
Dendryphiella paravinosa CBS 141286
100/0.99
--/0.98
Dendryphiella paravinosa CBS 121797
Dendryphiella paravinosa CBS 118716
Dendryphiella fasciculata MFLUCC 17-1074
Dendryphiella variabilis CBS 584.96
Dendryphiella
Dendryphiella stromaticola LAMIC 90/16
100/1.00
Dendryphiella vinosa
Dendryphiella vinosa MFLUCC 17-2321 F
Dendryphiella vinosa MFLUCC 17-2513
95/1.00
96/0.99
Dendryphiella eucalyptorum CBS 137987
100/1.00
100/1.00
--/0.99
100/1.00
Neodendryphiella mali FMR 17003
Neodendryphiella mali CBS 139.95
Neodendryphiella michoacanensis FMR 16098
Neodendryphiella
Neodendryphiella tarraconensis FMR 16234
Gregarithecium curvisporum KT 922
85/0.99
87/0.99
Gregarithecium
Pseudocoleophoma calamagrostidis KT 3284
Pseudocoleophoma polygonicola KT 731
Pseudocoleophoma
Pseudocoleophoma flavescens CBS 178.93
77/0.99
Pseudocoleophoma typhicola MFLUCC 16-0123 F
Lentithecium clioninum KT 1220
0.03
13
Outgroup
Fungal Diversity (2020) 105:319–575
Pseudotrichia allequashensis Fallah & Shearer, Mycologia
93(3): 597 (2001)
Freshwater distribution: USA (Fallah and Shearer 2001)
Key to freshwater Didymosphaeriaceae asexual genera
1. Conidiogenous cells phialidic or annellidic, with percurrent proliferations…………………Paraphaeosphaeria
1. Conidiogenous cells phialidic, sometimes percurrent…………………………………Paraconiothyrium
Paraconiothyrium Verkley, Stud. Mycol. 50(2): 327 (2004)
Saprobic on submerged wood in freshwater or plant materials in terrestrial habitats. Sexual morph: Undetermined.
Asexual morph: Conidiomata eustromatic, simple or complex, rarely pycnidial. Conidiogenous cells discrete or integrated, phialidic, sometimes percurrent. Conidia narrowly
ellipsoidal or short-cylindrical, aseptate or sometimes 1-septate, thin-walled, smooth-walled or minutely warted, hyaline
when young, brown when mature (Verkley et al. 2004).
Type species: Paraconiothyrium estuarinum Verkley &
Manuela Silva
Notes: Paraconiothyrium was introduced to accommodate the mycoparasite P. minitans (W.A. Campb.) Verkley
and the other four species (Verkley et al. 2004). Paraconiothyrium species are coelomycetous and regarded as asexual
morph of Paraphaeosphaeria because they clustered in a
well-supported clade (Verkley et al. 2004). However, they
clustered in two different clades in the later study of Wanasinghe et al. (2018b) who mentioned that the sexual morph of
Paraconiothyrium is undetermined, which was in agreement
with Wijayawardene et al. (2017). Paraconiothyrium fuckelii
(Sacc.) Verkley & Gruyter was found from freshwater habitats in USA (Shearer 1972), but not confirmed by molecular
data. The name P. fuckelii is illegitimate (Index Fungorum
2020) according to code, Rejection of names article 52. 1.
List of freshwater Paraconiothyrium species
Paraconiothyrium fuckelii (Sacc.) Verkley & Gruyter, Stud.
Mycol. 75: 25 (2012) [2013]
Basionym: Coniothyrium fuckelii Sacc., Fungi venet. nov.
vel. Crit., Sér. 5: 200 (1878)
Synonymy: see Index Fungorum (2020)
Freshwater distribution: USA (Shearer 1972)
Latoruaceae Crous, IMA Fungus 6(1): 176 (2015)
Pseudoasteromassaria M. Matsum. & Kaz. Tanaka, Fungal
Diversity 75: 77 (2015)
Saprobic on submerged wood or parasitic on twigs.
Sexual morph: Ascomata scattered, immersed, subglobose, brown to black, ostiolate. Peridium two-layered.
411
Pseudoparaphyses numerous, cellular, hyaline, septate,
branched. Asci 8-spored, bitunicate, cylindrical to clavate,
pedicellate. Ascospores bi- to tri-seriate, fusiform, brown,
septate (Ariyawansa et al. 2015). Asexual morph: Coelomycetous. Conidiomata pycnidial, solitary or scattered,
immersed or superficial, globose, ellipsoidal, conical, dark
brown to black, ostiolate. Peridium two-layered. Conidiophores absent. Conidiogenous cells enteroblastic, phialidic,
cylindrical or pyramidal, doliiform to ampulliform. Conidia
variable in shape, truncate at the base, aseptate or septate,
hyaline (Ariyawansa et al. 2015; Tibpromma et al. 2017).
Type species: Pseudoasteromassaria fagi M. Matsum. &
Kaz. Tanaka
Notes: The type species Pseudoasteromassaria fagi was
found on twigs with both sexual and asexual morphs, and
it produces cylindrical, hyaline, septate conidia in culture
(Ariyawansa et al. 2015). The second species P. spadicea
W. Dong et al. was described from submerged wood in
freshwater habitats. Pseudoasteromassaria spadicea differs
from the asexual morph of P. fagi by its subglobose, aseptate
conidia (Tibpromma et al. 2017). We report another species
P. aquatica from Thailand.
List of freshwater Pseudoasteromassaria species
*Pseudoasteromassaria aquatica W. Dong, H. Zhang &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557909; Facesoffungi number: FoF09252; Fig. 48
Etymology: in reference to aquatic habitat of the fungus
Holotype: MFLU 18-1519
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Undetermined. Asexual morph: Pycnidia
140–170 μm high, 210–250 μm diam., dark brown, scattered,
immersed, conical, coriaceous, ostiolate. Peridium 15–20 μm
thick, comprising 4–5 layers of brown to dark brown, irregular
cells arranged in a textura angularis. Conidiophores reduced.
Conidiogenous cells 5–8 × 2–4 μm (x̄ = 6.5 × 3 μm, n = 5),
phialidic, determinate, discrete, cylindrical, hyaline, thinwalled. Conidia 12–18 × 8–13 μm (x̄ = 15.5 × 10.5 μm, n =
20), ellipsoidal, obovoid or irregular, aseptate, guttulate, hyaline, smooth, thin-walled, straight, lacking mucilaginous sheath.
Culture characteristics: On PDA, colony circular, reaching
15 mm in 50 days at 25 °C, white to grey from above, brown
from below, surface rough, dry, raised, edge entire or undulate.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat829-1 (MFLU 18-1519, holotype), ex-type living culture MFLUCC 18-1397; ibid., hat829-2 (HKAS 105044,
isotype), living culture KUMCC 19-0063.
Notes: Pseudoasteromassaria aquatica is morphologically similar to P. spadicea W. Dong et al. in having immersed
pycnidia, phialidic conidiogenous cells, and hyaline, aseptate
13
412
Fig. 48 Pseudoasteromassaria aquatica (MFLU 18-1519, holotype).
a Conidiomata immersed in host. b Vertical section of pycnidium.
c Structure of peridium. d, e Conidiogenous cells with conidia. f–j
13
Fungal Diversity (2020) 105:319–575
Conidia. k Germinated conidium. l, m Colony on PDA (up-front,
down-reverse). Scale bars: b = 50 μm, c–e, j = 10 μm, f, k = 20 μm,
g–i = 5 μm
Fungal Diversity (2020) 105:319–575
conidia (Tibpromma et al. 2017). However, P. aquatica has
smaller pycnidia (140–170 × 210–250 μm vs. 290–320 ×
460–480 μm) and slightly larger conidia (12–18 × 8–13 μm
vs. 11–15 × 7–10 μm). Moreover, P. aquatica has ellipsoidal to
obovoid conidia, while P. spadicea has subglobose conidia. Phylogenetic analysis supports them to be different species (Fig. 2).
*Pseudoasteromassaria spadicea W. Dong, H. Zhang &
K.D. Hyde, Fungal Diversity 83: 59 (2017)
Freshwater distribution: Thailand (Tibpromma et al.
2017)
Key to freshwater Pseudoasteromassaria species
1. Pycnidia 140–170 μm high, 210–250 μm diam., conidia
ellipsoidal to obovoid, 12–18 × 8–13 μm………………
………………………………………………P. aquatica
1. Pycnidia 290–320 μm high, 460–480 μm diam., conidia
mostly subglobose, 11–15 × 7–10 μm………P. spadicea
Lentitheciaceae Y. Zhang ter, C.L. Schoch, J. Fourn., Crous
& K.D. Hyde, Stud. Mycol. 64: 93 (2009)
Key to freshwater sexual genera of Lentitheciaceae
1. Ascomata with ostiolate papilla covered by short dark
setae………………………………………Keissleriella
1. Ascomata without setae………………………………2
2. Ascospores filiform………………………Poaceascoma
2. Ascosproes fusiform to cylindrical……………………3
3. Ascospores surrounded by a large expanding sheath…
……………………………………………Tingoldiago
3. Ascospores without expanding sheath…………………4
4. Ascospores fusiform ……………………Lentithecium
4 . Ascospores cylindrical……………………Setoseptoria
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 128: 582 (1919)
Saprobic on submerged wood in freshwater or dead wood
in terrestrial habitats. Sexual morph: Ascomata immersed,
solitary to gregarious, erumpent to nearly superficial, uniloculate, globose, dark brown to black, with ostiolate papilla,
covered by dark setae or small blackened cells. Peridium
thick, composed of pseudoparenchymatous cells and pale
cells. Pseudoparaphyses dense, trabeculate, filiform, long,
rarely septate, anastomosing and branching. Asci 4- or
8-spored, bitunicate, fissitunicate, cylindro-clavate, with
a furcate pedicel and a small ocular chamber. Ascospores
medium-sized, uni- to bi-seriate, ellipsoidal to fusiform,
or clavate, cylindrical, straight to slightly curved, hyaline
to pale brown, or yellow, septate, sometimes with a vertical septum in central cells, constricted at the septa (Zhang
et al. 2012b). Asexual morph: Coelomycetes, developed
413
in culture. Pycnidia erumpent becoming superficial, uniloculate, subglobose to hemispherical, glabrous, dark
brown to black, with ostiolate papilla. Mycelium immersed.
Peridium thin, composed of thin-walled, flattened cells.
Conidiophore reduced. Conidiogenous cells holoblastic,
determinate, lageniform, cylindrical to doliiform, hyaline,
smooth, formed from the inner most layer of pycnidium wall.
Conidia medium-sized, cylindrical to bone-shaped, straight
to curved, hyaline to brown at aged, septate, smooth, thinwalled, without sheath (Tanaka et al. 2015).
Type species: Keissleriella aesculi (Höhn.) Höhn.
Notes: Keissleriella is an old genus comprising over
40 epithets in Index Fungorum (2020) with ca. 18 species
confirmed with molecular data (Phookamsak et al. 2019).
Keissleriella is characterized by globose ascomata, with a
small black papilla and short black external setae (Zhang
et al. 2012b). Keissleriella was shown to belong in Lentitheciaceae based on a few species (e.g. K. cladophila (Niessl)
Corbaz and K. linearis E. Müll. ex Dennis) (Schoch et al.
2009; Zhang et al. 2012b), which has been accepted by other
studies (Wijayawardene et al. 2017, 2018, 2020; Phookamsak et al. 2019). Keissleriella was regarded as congeneric
with Trichometasphaeria because their species (K. cladophila (Niessl) Corbaz and T. gloeospora (Berk. & Curr.) L.
Holm) clustered in one clade (Tanaka et al. 2015). However,
Wijayawardene et al. (2017, 2020) treated them as distinct
genera. Sequence data of their type species are needed to
solve their phylogenetic relationships. All Keissleriella species clustered in a weakly supported clade, which probably
indicated a polyphyletic nature.
Keissleriella linearis is the only species reported from
freshwater habitats (Zhang et al. 2009d). It was isolated from
dry culms of Phragmites communis in UK (Dennis 1964)
and later collected from an artificial lake in France (Zhang
et al. 2009d). Keissleriella linearis was transferred to Lentithecium based on phylogenetic analyses and examination
of a fresh specimen (IFRD 2008) (Zhang et al. 2009d). With
more sequences added, K. linearis was shown to be a member of Keissleriella, rather than Lentithecium (Tanaka et al.
2015; this study, Fig. 53). This result was consistant with
Singtripop et al. (2015), who re-examined the specimen of
K. linearis (IFRD 2008) and described the brown to dark
brown setae covering ostiole which is the typical character
of Keissleriella. In the phylogenetic analysis of Phookamsak
et al. (2019), K. linearis grouped in Lentithecium but with
low bootstrap support. We follow the results of Singtripop
et al. (2015) and Tanaka et al. (2015) until more sequences
are added to the family. Keissleriella linearis is characterized
by immersed ascomata covered with a clypeus and numerous, dark brown, cylindrical bristles surrounding the ostiole,
clavate asci and fusiform, hyaline, 3-septate ascospores with
a thick, mucous sheath (Dennis 1964).
13
414
List of freshwater Keissleriella species
*Keissleriella linearis E. Müll. ex Dennis, Kew Bull. 19(1):
120 (1964)
Synonymy: Lentithecium lineare (E. Müll. ex Dennis)
K.D. Hyde, J. Fourn. & Ying Zhang, Fungal Diversity 38:
236 (2009)
Freshwater distribution: France (Zhang et al. 2009d)
Lentithecium K.D. Hyde, J. Fourn. & Ying Zhang, Fungal
Diversity 38: 234 (2009)
Saprobic on submerged woody plant, grass in freshwater or dead leaf sheaths, stems, culms of grass in terrestrial
habitats. Sexual morph Ascomata scattered to grouped,
immersed to erumpent, uniloculate, globose to subglobose,
glabrous, dark brown to black, with ostiolate papilla. Peridium composed of several layers of thin-walled cells. Pseudoparaphyses numerous, cellular, hyaline, septate, branched,
persistent. Asci 8-spored, bitunicate, fissitunicate, clavatecylindrical to oblong-cylindrical, broadly rounded at the
apex, tapering to a short pedicel. Ascospores bi- or tri-seriate, fusiform, straight to slightly curved, hyaline to brown,
septate, usually asymmetrical, smooth to roughed, with or
without gelatinous sheath. Asexual morph Conidiomata
pycnidial, solitary or aggregated, unilocular, immersed,
erumpent to superficial, subglobose, ellipsoidal, dark brown
to black, glabrous or surrounded by brown, septate, thickwalled hyphae, with ostiolate papilla. Pycnidial wall composed of several layers of dark cells of textura angularis,
paler towards the inner layer. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells holoblastic, determinate, smooth-walled, hyaline, subglobose to pearshaped.
Conidia variable in shape, hyaline, aseptate, smooth (Ryckegem 2001; Zhang et al. 2009d; Hyde et al. 2016b).
Type species: Lentithecium fluviatile (Aptroot & Van
Ryck.) K.D. Hyde, J. Fourn. & Ying Zhang
Notes: Lentithecium was established to accommodate L.
aquaticum Ying Zhang et al. and three new combinations L.
arundinaceum (Sowerby) K.D. Hyde et al., L. fluviatile (type
species) and L. lineare (E. Müll. ex Dennis) K.D. Hyde et al.
(Zhang et al. 2009d). However, L. arundinaceum was later
transferred to Setoseptoria based on phylogenetic analyses,
although no morphological connection between L. arundinaceum (ascomycetes) and S. phragmitis Quaedvl. et al. (type species, coelomycetes) was shown (Tanaka et al. 2015). Lentithecium was previously characterized by the lenticular ascomata
(Zhang et al. 2009d) and later emended to globose after reexamination of the holotype of L. fluviatile (Hyde et al. 2013).
The freshwater species Lentithecium clioninum (Kaz.
Tanaka et al.) Kaz. Tanaka & K. Hiray. and L. pseudoclioninum Kaz. Tanaka & K. Hiray. formed a well-supported clade
with the type species, L. fluviatile (Fig. 53). Lentithecium
cangshanense Z.L. Luo et al., L. carbonneanum J. Fourn.
13
Fungal Diversity (2020) 105:319–575
et al., L. unicellulare Abdel-Aziz and a new species L. kunmingense clustered in a separated clade with low bootstrap
support (Fig. 53), and close to the genus Halobyssothecium
Dayarathne et al. Lentithecium cangshanense, collected from
submerged wood, is probably conspecific with the marine
species L. voraginesporum Abdel-Wahab et al., because of
their nearly identical sequences (one nucleotide difference in
LSU and SSU sequence data, respectively). Unfortunately,
their ITS sequence data are lacking. Hyde et al. (2016b)
distinguished them based on the size of ascomata, asci and
ascospores which are considered as tiny differences in this
study. More sequenced collections are needed in future.
Lentithecium aquaticum Ying Zhang et al., which was
collected from submerged wood in France, clustered in
Lentithecium (Zhang et al. 2009d). With more sequenced
species added in the family, the placement of L. aquaticum
appeared to be unstable and had a weak relationship with
Lentithecium clade (Tanaka et al. 2015; Hyde et al. 2016b;
Su et al. 2016a). The most recent publications showed it was
phylogenetically distant from Lentithecium and clustered
with the members of Setoseptoria, but with low bootstrap
support (Wanasinghe et al. 2018b; Phookamsak et al. 2019).
Our phylogenetic analysis show L. aquaticum forms a basal
clade to Darksidea, Halobyssothecium and Lentithecium
(Fig. 53). We exclude L. aquaticum from Lentithecium but
more phylogenetic evidence is required.
Lentithecium unicellulare Abdel-Aziz, which was collected from submerged wood in the River Nile in Egypt, is
the first asexual species in the genus (Hyde et al. 2016b).
We report the second asexual species L. kunmingense, which
was collected from China.
List of freshwater Lentithecium species
*Lentithecium cangshanense Z.L. Luo, X.J. Su & K.D.
Hyde, Phytotaxa 267: 65 (2016)
Freshwater distribution: China (Su et al. 2016a)
*Lentithecium carbonneanum J. Fourn., Raja & Oberlies,
Persoonia 40: 295 (2018)
Freshwater distribution: France (Crous et al. 2018b)
*Lentithecium clioninum (Kaz. Tanaka et al.) Kaz. Tanaka
& K. Hiray., Stud. Mycol. 82: 99 (2015); Fig. 49a–f
Basionym: Massarina clionina Kaz. Tanaka et al., Mycoscience 46(5): 288 (2005)
Freshwater distribution: Japan (Tanaka et al. 2005b)
*Lentithecium kunmingense W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557910; Facesoffungi number: FoF09253; Fig. 50
Fungal Diversity (2020) 105:319–575
415
Fig. 49 Lentithecium spp. (Material examined: JAPAN, Hokkaido,
Akkeshi, Bekanbeushi-river, on submerged twigs of woody plant,
2 June 2003, K. Tanaka & S. Hatakeyama, KT 1149A, holotype;
Hokkaido, Akkeshi, Toraibetsu-river, on submerged twigs of woody
plant, 3 June 2003, K. Tanaka & S. Hatakeyama, KT 1220, paratype;
Aomori, Hirosaki, Aoki, Mohei pond, on submerged twigs of woody
plant, 3 May 2003, K. Tanaka & N. Asama, KT 1113, holotype).
a–f L. clioninum (a–d from KT 1149A, holotype; e from KT 1220,
paratype; f from culture of KT 1149A). a Ascomata on the host surface. b Section of ascoma. c Peridium. d Pseudoparaphyses. e Ascus.
f Ascospore. g Ascospore of L. pseudoclioninum (KT 1113, holotype). Scale bars: a = 500 μm, b = 100 μm, c–g = 10 μm
Etymology: referring to Kunming, where the holotype
was collected
Holotype: HKAS 102150
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Undetermined. Asexual morph: Pycnidia
210–250 μm high, 320–350 μm diam., scattered or solitary,
unilocular, immersed to erumpent, ellipsoidal, dark brown to
black, glabrous, with ostiolate papilla. Peridium 60–80 μm
thick, composed of several layers of brown to black brown
cells of textura angularis, hyaline to pale brown towards
the inner layer which bearing conidiogenous cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
5–19 × 2–5 μm (x̄ = 11.5 × 3.5 μm, n = 10), phialidic, determinate, smooth-walled, hyaline, subglobose to ellipsoidal.
Conidia 8–14 × 5–8 μm (x̄ = 10.5 × 7 μm, n = 55), subglobose, ovate, ellipsoid, clavate, pyriform or irregular, with
rounded or truncate base, hyaline, aseptate, several small to
one big guttulate, smooth-walled.
Culture characteristics: On PDA, colony filamentous,
reaching 45 mm in 25 days at 25 °C, black from above and
below, surface dry, with sparse aerial mycelium, mostly
immersed in culture, with a filiform edge.
Material examined: CHINA, Yunnan Province, Kunming
University of Science and Technology, on submerged wood
in a stream, 10 May 2017, C.X. Liu, L54 (HKAS 102150,
holotype), ex-type living culture KUMCC 19-0101.
Notes: Lentithecium kunmingense clusters with L. voraginesporum without bootstrap support (Fig. 53). Lentithecium
kunmingense is a coelomycetous species, while L. voraginesporum is an ascomycetous species (Hyde et al. 2016b). Lentithecium voraginesporum only has LSU and SSU sequence
data in GenBank, and there are thirteen and four nucleotide
differences in LSU and SSU sequence data between L. voraginesporum (CBS H-22560) and L. kunmingense (KUMCC
19-0101), respectively, which indicate them to be different
species (Jeewon and Hyde 2016).
Only one asexual species Lentithecium unicellulare has
been reported in Lentithecium (Hyde et al. 2016b). Lentithecium kunmingense is similar to L. unicellulare in having
dark pycnidia, determinate, hyaline conidiogenous cells and
variable, hyaline, aseptate conidia. However, L. kunmingense
differs from L. unicellulare in having glabrous pycnidia and
larger conidia (8–14 × 5–8 μm vs. 6–9 × 4–5 μm) (Hyde
et al. 2016b). Phylogenetic analysis supports them to be different species (Fig. 53).
*Lentithecium pseudoclioninum Kaz. Tanaka & K. Hiray.,
Stud. Mycol. 82: 99 (2015); Fig. 49g
13
416
Fungal Diversity (2020) 105:319–575
Fig. 50 Lentithecium kunmingense (HKAS 102150,
holotype). a Conidiomata
immersed in host. b Vertical
section of pycnidium. c Structure of peridium. d–g Conidiogenous cells with conidia.
h–m Conidia. n Germinated
conidium. o, p Colony on PDA
(left-front, right-reverse). Scale
bars: b = 50 μm, c = 20 μm,
d–n = 5 μm
Freshwater distribution: Japan (Tanaka et al. 2015)
*Lentithecium unicellulare Abdel-Aziz, Fungal Diversity
80: 53 (2016)
Freshwater distribution: Egypt (Hyde et al. 2016b)
Key to freshwater Lentithecium species
1. Asexual morph…………………………………………2
1. Sexual morph…………………………………………3
13
2.
2.
3.
3.
4.
Conidia 6–9 × 4–5 μm…………………L. unicellulare
Conidia 8–14 × 5–8 μm………………L. kunmingense
Ascospores hyaline……………………………………4
Ascospores pigmented…………………………………5
Ascospores surrounded by a wing-like sheath…………
……………………………………………L. clioninum
4. Ascospores lacking wing-like sheath…………………
……………………………………L. pseudoclioninum
5. Asci > 100 µm long, ascospores rough-walled…………
………………………………………L. carbonneanum
Fungal Diversity (2020) 105:319–575
417
5. Asci < 100 µm long, ascospores smooth-walled………
………………………………………L. cangshanense
*Poaceascoma aquaticum Z.L. Luo & K.D. Hyde, Phytotaxa 253: 75 (2016)
Freshwater distribution: Thailand (Luo et al. 2016a)
Poaceascoma Phookamsak & K.D. Hyde, Cryptogamie,
Mycologie 36: 231 (2015)
Saprobic on submerged bamboo, wood in freshwater or intertidal, dead stem in terrestrial habitats. Sexual
morph: Ascomata solitary to gregarious, semi-immersed to
erumpent, uniloculate, globose to subglobose, with or without setose, black, with a central ostiole, with short to long
papilla. Peridium thick walled, of equal thickness, composed
of several layers of dark brown to black, pseudoparenchymatous cells. Pseudoparaphyses dense, cellular, hyaline, septate, persistent, anastomosing above the asci, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
elongate-cylindrical, short pedicellate, apically rounded
with an ocular chamber. Ascospores fasciculate, spirally
arranged within the ascus, filiform, curved, hyaline, multiseptate, smooth, thin-walled, without sheath or appendage,
ascospores often longer than asci (Phookamsak et al. 2015a).
Asexual morph: Undetermined.
Type species: Poaceascoma helicoides Phook. & K.D.
Hyde
Notes: Poaceascoma was introduced to accommodate
ophiosphaerella-like species associated with Poaceae, which
form semi-immersed to erumpent, setose ascomata with short
to long necks and filiform ascospores spirally arranged in asci
(Phookamsak et al. 2015a). Poaceascoma is a well-studied
genus with sequence data available for all species in GenBank; they formed a monophyletic clade in the family with
high bootstrap support (Phookamsak et al. 2015a; Luo et al.
2016a; Hyde et al. 2017, 2018a). Some genera also form filiform ascospores in Dothideomycetes, such as Acanthophiobolus, Leptospora, Ophiobolus and Ophiosphaerella (Shoemaker
1976; Boonmee et al. 2011; Zhang et al. 2012b; Boonmee et al.
2014). Acanthophiobolus differs from these genera in having
rather smaller, superficial, apapillate ascomata (Boonmee et al.
2011, 2014). Leptospora is distinct in having possesses glabrous ascomata which often stain host tissues red or purple
(Hyde et al. 2016b). Ophiobolus and Ophiosphaerella nested
in Phaeosphaeriaceae (Phookamsak et al. 2014), which was
phylogenetically distant from Poaceascoma (Lentitheciaceae).
Poaceascoma aquaticum Z.L. Luo & K.D. Hyde was
collected from submerged bamboo in Thailand (Luo et al.
2016a). It was described as globose to subglobose ascomata
without setae, cylindrical to cylindric-clavate asci and pale
brown to brown ascospores (Luo et al. 2016a). However,
the ascospores were obviously hyaline in the illustration of
Luo et al. (2016b).
Setoseptoria Quaedvl., Verkley & Crous, Stud. Mycol. 75:
382 (2013)
Saprobic on decaying leaves or wood/branch in terrestrial
or submerged wood in fershwater habitats. Sexual morph:
Ascomata erumpent to superficial, scattered to gregarious,
sometimes scattered beneath the host periderm or on decorticated wood, subglobose to hemispherical with flattened
base, glabrous, sometimes with sparse brown hyphae at
sides, dark brown to black, with a central ostiole, with or
without short papilla. Peridium thin, composed of several
layers of thin-walled cells. Pseudoparaphyses numerous,
cellular, hyaline, septate, branched, persistent, anastomosing between and above the asci. Asci 8-spored, bitunicate,
fissitunicate, clavate to cylindric-clavate, or narrowly ovoid,
pedicellate, rounded at the apex, with an indistinct or minute
ocular chamber. Ascospores uni- to tri-seriate, cylindrical to
narrowly fusiform, mostly hyaline, sometimes hyaline to pale
olivaceous, septate, deeply constricted at the primary septum, mostly asymmetrical, smooth, thin-walled, surrounded
by a gelatinous sheath (Tanaka et al. 2004; Hyde et al. 2017;
Wanasinghe et al. 2018b). Asexual morph: Coelomycetous.
Pycnidia immersed, uniloculate, globose, with brown, verruculose to warty setae, brown, ostiolate, somewhat papillate. Peridium composed of several layers of brown, angular
cells which inwardly become hyaline. Conidiophores reduced
to conidiogenous cells or with one supporting cell. Conidiogenous cells subcylindrical to doliiform, hyaline, smooth,
apical region with several inconspicuous percurrent proliferations, or with periclinal thickening; collarette inconspicuous, or prominent, flared. Conidia subcylindrical, straight
to somewhat curved, hyaline, smooth, transversely septate,
thin-walled, without appendages or sheath (Quaedvlieg et al.
2013).
Type species: Setoseptoria phragmitis Quaedvl., Verkley
& Crous
Notes: Setoseptoria was introduced for a coelomycetous species S. phragmitis which forms setose pycnidia and
subcylindrical, (1–)3-septate, hyaline conidia that become
olivaceous and verruculose in older cultures (Quaedvlieg
et al. 2013). Another six ascomycetous species were added
to the genus, but the asexual morphs were not produced in
culture. These species forms a well-supported clade in our
phylogenetic tree (Fig. 53), but lack strong bootstrap support
between each other (Hyde et al. 2017; Wanasinghe et al.
2018b). Tanaka et al. (2015) mentioned that the species
producing both sexual and asexual morphs were needed to
confirm the validity of this generic treatment.
Setoseptoria arundinacea (Sowerby) Kaz. Tanaka & K.
Hiray., a common species occurring on Phragmites culms,
List of freshwater Poaceascoma species
13
418
Fungal Diversity (2020) 105:319–575
Fig. 51 Setoseptoria spp. (Material examined: JAPAN, Aomori,
Hirosaki, Kadoke, Oowasawa-river, on dead culms of Phragmites
australis, 29 July 2001, K. Tanaka, KT 600 = HHUF 27544; Hokkaido, Akkeshi, Ariake, small stream, on submerged stems of herbaceous plant, 3 June 2003, K. Tanaka & S. Hatakeyama, KT 1174
= HHUF 28293, holotype). a–e S. arundinacea (KT 600 = HHUF
27544). a Ascomata on the host surface. b, c Section of ascomata. d
Ascus. e Ascospore. f, g Ascospores of S. magniarundinacea (culture of KT 1174 = CBS 139702). Scale bars: a = 1 mm, b = 500 μm,
c = 100 µm, d–g = 10 μm
was also reported from freshwater habitats (Aptroot 1998;
Tanaka and Harada 2003c; Luo et al. 2004; Zhang et al.
2009d). Setoseptoria arundinacea may consist of several
cryptic species because two strains, KT 552 and KT 600
(Tanaka et al. 2015), did not form a clade with another two
strains CBS 619.86 (Schoch et al. 2009) and CBS 123131
(Zhang et al. 2009c), which showed in Tanaka et al. (2015),
Hyde et al. (2017), Wanasinghe et al. (2018b) and our study
(Fig. 53). The epitype of this species is expected to solve
this S. arundinacea complex. The second freshwater species S. magniarundinacea (Kaz. Tanaka & Y. Harada) Kaz.
Tanaka & K. Hiray. has larger ascospores than S. arundinacea (Tanaka et al. 2004) (see key below).
Basionym: Massarina magniarundinacea Kaz. Tanaka &
Y. Harada, Mycotaxon 90(2): 349 (2004)
Freshwater distribution: Japan (Tanaka and Harada 2004)
List of freshwater Setoseptoria species
*Setoseptoria arundinacea (Sowerby) Kaz. Tanaka & K.
Hiray., Stud. Mycol. 82: 101 (2015); Fig. 51a–e
Basionym: Sphaeria arundinacea Sowerby, Col. fig. Engl.
Fung. Mushr. (London) 3: tab. 336 (1803)
Synonymy: Possible synonyms see Index Fungorum (2020)
Freshwater distribution: China (Luo et al. 2004), Denmark (Zhang et al. 2009d)
*Setoseptoria magniarundinacea (Kaz. Tanaka & Y.
Harada) Kaz. Tanaka & K. Hiray., Stud. Mycol. 82: 102
(2015); Fig. 51f, g
13
Key to freshwater Setoseptoria species
1. Ascospores 23–40 × 3.5–6 µm………S. arundinacea
1. Ascospores 67–82 × 6.5–9 µm…S. magniarundinacea
Tingoldiago K. Hiray. & Kaz. Tanaka, Mycologia 102(3):
740 (2010)
Saprobic on submerged culms. Sexual morph: Ascomata
scattered, immersed to erumpent, uniloculate, depressed globose to conical, with a flattened base, glabrous, black, with
a central, rounded ostiole. Peridium composed of hyaline to
brown, small cells. Pseudoparaphyses numerous, cellular,
hyaline, septate, immersed in gel matrix. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate, rounded at the apex,
with or without a shallow chamber. Ascospores bi- to triseriate, clavate, straight, asymmetrical, the upper cell often
broader than the lower one, smooth, thin-walled, usually surrounded by a large, expanding gelatinous sheath (Hirayama
et al. 2010; Xu et al. 2020). Asexual morph: Undetermined.
Type species: Tingoldiago graminicola K. Hiray. & Kaz.
Tanaka
Notes: Tingoldiago resembles Lindgomyces in ascospore
characters and aquatic habitats (Hirayama et al. 2010).
Fungal Diversity (2020) 105:319–575
419
Fig. 52 Tingoldiago graminicola (Material examined: JAPAN,
Aomori, Hirosaki, Kadoke, Oowasawa River, on submerged culms of
Phragmites japonica, 28 September 2002, KT 891, paratype). a, b,
d, e, g from KT 891, paratype. c, f from culture of KT 891. a Asco-
mata on the host surface. b Section of ascoma. c Pseudoparaphyses. d
Ascus. e Ascospore. f Ascospore with an elongate gelatinous sheath
in Indian Ink. Scale bars: a = 500 μm, b = 100 μm, c–e = 10 μm, f =
50 μm
However, Tingoldiago differs by its depressed globose to
mammiform ascomata with a wedge of palisade-like cells
at the rim and graminicolous substrate. Tingoldiago was
treated as a synonym of Lentithecium due to the lenticular, immersed to erumpent ascomata, cylindro-clavate asci
and hyaline, 1-septate ascospores with sheath (Zhang et al.
2012b). Later, its generic placement was supported by
molecular studies (Tanaka et al. 2015; Hyde et al. 2016b;
Wanasinghe et al. 2018b).
Freshwater species Tingoldiago graminicola (type) has
clavate, hyaline, 1-septate ascospores when young, becoming pale brown, 3-septate with age and with a fusiform
gelatinous sheath which expands to form a long appendage
in water (Hirayama et al. 2010). Other two freshwater species, T. clavata D.F. Bao et al. and T. hydei D.F. Bao et al.,
have several long or short, equatorial appendages, but lack
the expanding sheath (Xu et al. 2020).
Key to freshwater Tingoldiago species
List of freshwater Tingoldiago species
*Tingoldiago clavata D.F. Bao, L. Xu & H.Y. Su, MycoKeys 65: 128 (2020)
Freshwater distribution: Thailand (Xu et al. 2020)
*Tingoldiago hydei D.F. Bao, Z.L. Luo & H.Y. Su, MycoKeys 65: 126 (2020)
Freshwater distribution: Thailand (Xu et al. 2020)
1. Ascospores without appendages at the septum…………
…………………………………………T. graminicola
1. Ascospores with appendages at the septum……………2
2. Ascospores 48–51 × 7.5–9 μm………………T. clavata
2. Ascospores 37.5–42 × 7.5–9 μm………………T. hydei
Lindgomycetaceae K. Hiray., Kaz. Tanaka & Shearer, Mycologia 102(3): 733 (2010)
Key to freshwater sexual genera of Lindgomycetaceae
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
*Tingoldiago graminicola K. Hiray. & Kaz. Tanaka, Mycologia 102(3): 740 (2010); Fig. 52
Freshwater distribution: Japan (Hirayama et al. 2010)
Ascomata with a clypeus………………Neolindgomyces
Ascomata without clypeus……………………………2
Ascospores < 3-septate…………………………………3
Ascospores > 13-septate………………………………4
Ascospores hyaline to pale brown, 1(–2)-septate, or only
brown and > 3-septate in old ascospores…Lindgomyces
Ascospores hyaline, 1-septate………Hongkongmyces
Ascospores hyaline to golden-pale brown………
……………………………………………Arundellina
Ascospores brown to dark brown………………………5
Ascospores fusiform to clavate, transversely septate,
occasionally with one longitudinal or oblique septa, asci
4- to 8-spored……………………………………Lolia
Ascospores fusiform or vermiform, transversely
septate, without longitudinal or oblique septa, asci
8-spored………………………Aquimassariosphaeria
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420
Fungal Diversity (2020) 105:319–575
Fig. 53 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS and TEF sequence
data for species of Lentitheciaceae. Bootstrap values for
maximum likelihood equal to or
greater than 75% and Bayesian
posterior probabilities equal to
or greater than 0.95 are placed
near the branches as ML/BYPP.
Newly generated sequences are
in red and ex-type strains are
in bold. The new species introduced in this study are indicated
with underline. Freshwater
strains are indicated with a red
letter “F”. The tree is rooted to
Massarina cisti CBS 266.62,
M. eburnea CBS 473.64 and
M. eburnea H 3953 (Massarinaceae)
75/1.00 Keissleriella breviasca KT 649
100/1.00
Keissleriella breviasca KT 581
Keissleriella breviasca KT 540
Keissleriella poagena CBS 136767
100/1.00
Keissleriella culmifida KT 2308
76/--
Keissleriella culmifida KT 2642
Keissleriella trichophoricola CBS 136770
99/1.00
Keissleriella sp. KT 895
Keissleriella quadriseptata KT 2292
--/0.97
Keissleriella gloeospora KT 829
Keissleriella taminensis KT 678
92/1.00
100/1.00
99/1.00
Keissleriella taminensis KT 594
Keissleriella taminensis KT 571
Keissleriella rosae MFLUCC 15-0180
Keissleriella cirsii MFLUCC 16-0454
Keissleriella dactylidicola MFLUCC 13-0866
Keissleriella lineare IFRD 2008 F
99/1.00
Keissleriella phragmiticola MFLUCC 17-0779
99/--
Pleurophoma ossicola CPC 24985
Keissleriella
Pleurophoma ossicola CBS 139905
Pleurophoma pleurospora CBS 130329
Keissleriella genistae CBS 113798
Pleurophoma italica MFLUCC 15-0061
Keissleriella rosacearum MFLUCC 15-0045
Keissleriella rosarum MFLUCC 15-0089
Keissleriella sparticola MFLUCC 14-0196
Keissleriella cladophila CBS 104.55
Keissleriella tamaricicola MFLUCC 14-0168
Keissleriella pleurospora CBS 116668
Phragmocamarosporium hederae MFLUCC 13-0552
Phragmocamarosporium platani MFLUCC 14-1191
Phragmocamarosporium rosae MFLUCC 17-0797
Murilentithecium clematidis MFLUCC 14-0561
Murilentithecium rosae MFLUCC 15-0044
100/1.00
Keissleriella caraganae KUMCC 18-0163
Keissleriella caraganae KUMCC 18-0164
81/0.98
Keissleriella yonaguniensis KT 2604
Arundellina Wanas., E.B.G. Jones & K.D. Hyde, Fungal
Diversity 80: 59 (2016)
Saprobic on dead submerged stem. Sexual morph:
Ascomata solitary to scattered, immersed, uniloculate,
globose, glabrous, dark brown to black, coriaceous, with
ostiolate papilla. Peridium 4–5 layered, outer layer heavily
pigmented, comprising reddish to dark brown, thick-walled
cells of textura angularis, inner layer comprising hyaline,
13
thin-walled cells of textura angularis. Pseudoparaphyses
numerous, cellular, septate, branched. Asci 8-spored, bitunicate, fissitunicate, cylindrical to cylindric-clavate, pedicellate, with an ocular chamber. Ascospores overlapping
uni- to bi-seriate, fusiform, hyaline to golden-pale brown,
transversely septate, with pointed ends (Hyde et al. 2016b).
Asexual morph: Undetermined.
Fungal Diversity (2020) 105:319–575
421
Fig. 53 (continued)
Halobyssothecium obiones 08AV2569
100/1.00
90/1.00
Halobyssothecium obiones 13AV2143
96/1.00
Halobyssothecium obiones 07AV2562
99/1.00
Halobyssothecium obiones 27AV2385
Halobyssothecium
Halobyssothecium obiones 20AV2566
Halobyssothecium obiones MFLUCC 15-0381
Lentithecium carbonneanum CBS 144076 F
Lentithecium unicellulare MD 6004 F
Lentithecium kunmingense KUMCC 19-0101 F
--/0.98
Lentithecium voraginesporum CBS H-22560
Lentithecium cangshanense DLUCC 0143 F
91/1.00
100/0.99 Lentithecium pseudoclioninum CBS 139695 F
Lentithecium pseudoclioninum KT 1111 F
98/1.00
Lentithecium
Lentithecium fluviatile CBS 123090
98/1.00
100/1.00 Lentithecium fluviatile CBS 122367
Lentithecium clioninum CBS 139694 F
79/0.99
100/1.00 Lentithecium clioninum MAFF 243839 F
97/0.97 Darksidea epsilon CBS 135658
Darksidea delta CBS 135638
76/0.95
100/1.00
Darksidea gamma CBS 135634
--/0.99
Darksidea
Darksidea beta CBS 135637
100/1.00
Darksidea alpha CBS 135650
100/1.00
Darksidea zeta CBS 135640
Lentithecium aquaticum CBS 123099 F
Setoseptoria arundinacea KT 600
95/--
90/0.98 Setoseptoria arundinacea KT 552
Setoseptoria arundelensis MFLUCC 17-0759
Setoseptoria englandensis MFLUCC 17-0778
100/0.99 Setoseptoria phragmitis CBS 114802
85/--
Setoseptoria phragmitis CBS 114966
Setoseptoria arundinacea CBS 123131
98/1.00
Setoseptoria
Setoseptoria arundinaceum CBS 619.86
Stagonospora macropycnidia CBS 114202
98/1.00
Setoseptoria magniarundinacea KT 1174 F
Setoseptoria lulworthcovensis MFLU 18-0110
99/1.00
Poaceascoma helicoides MFLUCC 11-0136
Poaceascoma taiwanense MFLU 18-0083
100/1.00
Poaceascoma aquaticum MFLUCC 14-0048 F
Poaceascoma
Poaceascoma halophilum MFLUCC 15-0949
87/-- Tingoldiago clavata MFLUCC 19-0495 F
98/1.00
99/1.00
Tingoldiago clavata MFLUCC 19-0496 F
Tingoldiago clavata MFLUCC 19-0498 F
Tingoldiago hydei MFLUCC 19-0499 F
96/1.00
Tingoldiago
Tingoldiago graminicola KH 155 F
Tingoldiago graminicola KT 891 F
100/1.00
Tingoldiago graminicola KH 68 F
Towyspora aestuari MFLUCC 15-1274
Neoophiosphaerella sasicola KT 1706
--/1.00
Katumotoa bambusicola KT 1517a
100/1.00
100/1.00
Towyspora
Neoophiosphaerella
Katumotoa
Massarina eburnea CBS 473.64
Massarina eburnea H 3953
Outgroup
Massarina cisti CBS 266.62
0.02
Type species: Arundellina typhae Wanas., E.B.G. Jones
& K.D. Hyde
Notes: Arundellina is a monotypic genus introduced
for A. typhae, which was isolated from submerged stems
of Typha sp. (Typhaceae) in Arun River in UK (Hyde
et al. 2016b). The combined LSU and SSU sequence data
placed it in Lindgomycetaceae (Hyde et al. 2016b), which
is confirmed in this study (Fig. 2). The globose ascomata,
13
422
Fungal Diversity (2020) 105:319–575
Fig. 54 Arundellina typhae
(Material examined: UK,
England, Arun River, on dead
submerged stem of Typha sp.
(Typhaceae), 6 April 2015,
E.B.G. Jones, GJ122, MFLU
16-1276, holotype). a Appearance of immersed ascomata
on host substrate (arrows). b
Section of ascoma. c Peridium.
d Pseudoparaphyses. e, f Asci.
g–i Ascospores. Scale bars: b =
100 µm, c, e, f = 20 µm, d = 5
µm, g–i = 10 µm
cylindric-clavate asci with thick-walled apex, fusiform, 3–4
transversely septate, golden-pale brown ascospores with
pointed ends of A. typhae are unique in Lindgomycetaceae
(Fig. 53).
List of freshwater Arundellina species
*Arundellina typhae Wanas., E.B.G. Jones & K.D. Hyde,
Fungal Diversity 80: 61 (2016); Fig. 54
Freshwater distribution: UK (Hyde et al. 2016b)
Hongkongmyces C.C.C. Tsang et al., Medical Mycol. 52(7):
740 (2014)
Parasitic on biopsy tissues of an infected foot of a
patient or saprobic on submerged wood. Sexual morph:
Ascomata immersed to semi-immersed, scattered or gregarious, subglobose, coriaceous, dark brown to black,
with papilla. Peridium composed of several layers of dark
brown cells of textura epidermoidea or textura angularis.
13
Pseudoparaphyses numerous, trabeculate, filiform, hyaline, septate, branched. Asci 8-spored, bitunicate, fisitunicate, cylindric-clavate, pedicellate. Ascospores overlapping
biseriate, broad-fusiform, sometimes tapering towards the
ends, hyaline, septate, surrounded with mucilaginous sheath
(Hyde et al. 2017). Asexual morph: Coelomycetous. Conidiomata pycnidial, globose to ampulliform or ellipsoidal, dark
brown to black, with a central ostiole to multiple ostioles.
Peridium two-layered, or sometimes poorly developed and
thinner at the base. Conidiophores reduced. Conidiogenous
cells phialidic, sometimes with sympodial proliferations,
discrete, subulate to ampulliform or subcylindrical, hyaline,
smooth. Conidia white in mass, hyaline, solitary, ellipsoid to
obovoid, globose, subglobose, lacking mucilaginous sheath
(Crous et al. 2018b).
Type species: Hongkongmyces pedis C.C.C. Tsang et al.
Notes: Hongkongmyces pedis was isolated from biopsy
tissues of an infected foot of a patient and it did not produce
Fungal Diversity (2020) 105:319–575
fruiting bodies or conidia in culture (Tsang et al. 2014).
Later, an asexual morph H. snookiorum Raudabaugh et al.
was isolated from submerged wood (Crous et al. 2018b). In
our phylogenetic analysis (Fig. 2), the sexual species H. thailandicus Phukhams. & K.D. Hyde and the new species H.
aquaticus forms a weakly-supported clade, separated from
the type species, H. pedis. In previous studies, H. thailandicus also had a weak relationship with H. pedis (Hyde et al.
2017; Crous et al. 2018b). This group needs further study
with more collections.
423
List of freshwater Hongkongmyces species
*Hongkongmyces aquaticus W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557911; Facesoffungi number: FoF09254; Fig. 55
Etymology: in reference to aquatic habitat of this fungus
Holotype: MFLU 18-1516
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Undetermined. Asexual morph: Pycnidia
270–320 μm high, 300–350 μm diam., dark brown to black,
Fig. 55 Hongkongmyces
aquaticus (MFLU 18-1516,
holotype). a, b Appearance of
black conidiomata on host. c,
d Vertical section of pycnidia.
e–g Conidiogenous cells with
conidia. h–k Conidia. l Germinated conidium. m, n Colony on
PDA (up-front, down-reverse).
Scale bars: c, d = 50 μm, e–k =
10 μm, l = 20 μm
13
424
scattered, semi-immersed or erumpent to host surface, globose
or ellipsoidal, coriaceous, ostiolate. Peridium 50–70 μm thick
at the sides, poorly developed and thinner at the base, 30–40
μm thick, composed of dark brown, large, compressed cells
of textura angularis. Conidiophores reduced. Conidiogenous
cells 9–12 × 3–7 μm ( x̄ = 10.5 × 5 μm, n = 5), phialidic,
determinate, cylindrical to subcylindrical, hyaline, thin-walled.
Conidia 14–19 × 11–15 μm ( x̄ = 16.5 × 12.5 μm, n = 20),
globose, subglobose or obovoid, aseptate, guttulate, hyaline,
smooth, thin-walled, straight, lacking mucilaginous sheath.
Culture characteristics: On PDA, colony circular, reaching
15 mm in 10 days at 25 °C, grey to brown from above, dark
brown from below, surface rough, dry, raised, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
20180525-1 (MFLU 18-1516, holotype), ex-type living culture MFLUCC 18-1150; ibid., 20180525-2 (HKAS 105003,
isotype), ex-type living culture KUMCC 19-0019.
Notes: Hongkongmyces aquaticus resembles H. snookiorum in having phialidic conidiogenous cells and hyaline,
variable shaped conidia, but it lacks sympodial proliferations (Crous et al. 2018b). Additionally, H. aquaticus has
larger conidia (14–19 × 11–15 μm vs. 4.5–5.5 × 3.5–4 μm).
Both species were collected from freshwater habitats, but
from different substrates (submerged wood vs. submerged
detritus). Phylogenetic analysis supports them to be different
species (Fig. 2).
*Hongkongmyces snookiorum Raudabaugh, Iturr. & A.N.
Mill., Persoonia 40: 289 (2018)
Freshwater distribution: USA (Crous et al. 2018b)
Key to freshwater Hongkongmyces species
1. Conidia 4.5–5.5 × 3.5–4 μm……………H. snookiorum
1. Conidia 14–19 × 11–15 μm………………H. aquaticus
Aquimassariosphaeria W. Dong & Doilom, gen. nov.
Index Fungorum number: IF557825; Facesoffungi number: FoF08733
Etymology: referring to type species was collected
from aquatic habitat and its morphological similarity to
Massariosphaeria
Saprobic on submerged wood or dry branches. Sexual
morph: Ascomata scattered or solitary, immersed, erumpent
to superficial, subglobose to ellipsoidal, black, coriaceous,
with ostiolate papilla, sometimes stain substrate purple.
Peridium two-layered, outer layer comprising several layers
of dark brown to black, thin-walled, compressed cells, inner
layer comprising several layers of pale brown to hyaline,
thin-walled, large cells of textura angularis. Pseudoparaphyses numerous, cellular, hyaline, septate, embedded in a
13
Fungal Diversity (2020) 105:319–575
matrix. Asci 8-spored, bitunicate, narrowly clavate, short
pedicellate, apically rounded. Ascospores bi- to tri-seriate,
narrowly fusiform or vermiform, straight or curved, brown,
transversely septate, thin-walled, with or without a sheath.
Asexual morph: Undetermined.
Type species: Aquimassariosphaeria kunmingensis W.
Dong, Doilom & K.D. Hyde
Notes: Aquimassariosphaeria is characterized by its
immersed, erumpent to superficial ascomata, sometimes
staining substrate purple, and narrowly fusiform or vermiform, brown, transversely septate ascospores. Morphologically, Aquimassariosphaeria is similar to Lolia in having
fusiform to clavate, multiple transversely septate, brown
ascospores, but the conidia of the latter equipped with apical,
sub-apical and basal appendages, which are never formed
in Aquimassariosphaeria. Phylogenetic analyses also support them to be different genera (Figs. 2, 61). Additionally,
the type species of Massariosphaeria, M. phaeospora (E.
Müll.) Crivelli, clusters in Cyclothyriellaceae (Jaklitsch and
Voglmayr 2016), while M. typhicola (CBS 609.86) clusters
in Lindgomycetaceae (Fig. 2). Although Ariyawansa et al.
(2015) transferred Neomassariosphaeria Y. Zhang ter et al.
from Amniculicolaceae to Lindgomycetaceae, to accommodate Massariosphaeria typhicola, we do not accept in this
study (see notes under Neomassariosphaeria, Amniculicolaceae). A new genus Aquimassariosphaeria is, therefore,
established to accommodate Massariosphaeria typhicola
and the new species A. kunmingensis.
List of freshwater Aquimassariosphaeria species
*Aquimassariosphaeria kunmingensis W. Dong, Doilom
& K.D. Hyde, sp. nov.
Index Fungorum number: IF557912; Facesoffungi number: FoF08734; Fig. 56
Etymology: referring to Kunming, where the holotype
was collected
Holotype: HKAS 102148
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 300–350 × 370–400 μm, scattered
or solitary, erumpent, mostly superficial, subglobose to ellipsoidal, black, coriaceous, with ostiolate papilla. Peridium
30–40 μm thick, two-layered, outer layer comprising 4–6
layers of dark brown or black, thin-walled, compressed cells
of textura angularis or irregular cells, inner layer comprising
3–5 layers of pale brown to hyaline, thin-walled, large cells
of textura angularis. Pseudoparaphyses 2 μm diam., numerous, cellular, hyaline, unbranched, septate, embedded in a
matrix. Asci 160–200 × 15–18 μm ( x̄ = 180 × 17 μm, n =
5), 8-spored, bitunicate, narrowly clavate, short pedicellate,
apically rounded. Ascospores 35–46(–55) × 6.5–9.5 μm (x̄ =
40 × 7.8 μm, n = 25), bi- to tri-seriate in the upper part, uniseriate in the lower part, narrowly fusiform or vermiform,
Fungal Diversity (2020) 105:319–575
425
Fig. 56 Aquimassariosphaeria
kunmingensis (HKAS 102148).
a, b Appearance of ascomata
on host substrate. c Vertical
section of ascoma. d Structure
of peridium. e, f Bitunicate
asci. g Pseudoparaphyses. h–m
Ascospores. n Germinated
ascospore. o, p Colony on PDA
(left-front, right-reverse). Scale
bars: c = 50 μm, d–n = 20 μm
occasionally narrowly clavate with elongate basal cells,
with rounded ends, brown, 6–7-septate, occasionally 5- or
8-septate, constricted at the septa, asymmetric, obviously
enlarged at the third or fourth cell from the apex, straight
or curved, smooth, thin-walled, without a sheath. Asexual
morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 15 days at 25 °C, black from above, dark
brown from below, surface rough, with sparse mycelium,
mostly immersed in culture, dry, raised, edge entire.
Material examined: CHINA, Yunnan Province, Kunming
University of Science & Technology, on submerged wood
13
426
in a stream, 10 May 2017, C.X. Liu, L47 (HKAS 102148),
living culture KUMCC 18-1019.
Notes: Aquimassariosphaeria kunmingensis clusters with
A. typhicola with moderate bootstrap support (Figs. 2, 61).
Aquimassariosphaeria kunmingensis has superficial ascomata and thin-walled, smooth ascospores without a sheath,
while the latter has completely immersed ascomata that
staining substrate purple and thick-walled, heavily grained
sculpted ascospores with a well-defined, mucilaginous
sheath (Leuchtmann 1984). Comparison of single genes
between A. kunmingensis (KUMCC 18-1019) and A. typhicola (CBS 609.86) shows that there are 4, 1 and 16 nucleotide differences in LSU, SSU and RPB2 sequence data,
respectively, which supports them to be different species.
*Aquimassariosphaeria typhicola (P. Karst.) W. Dong &
Doilom, comb. nov.
Index Fungorum number: IF557913; Facesoffungi number: FoF09255
Basionym: Leptosphaeria typhicola P. Karst., Bidr. Känn.
Finl. Nat. Folk 23: 100 (1873)
Synonymy: Massariosphaeria typhicola (P. Karst.)
Leuchtm., Sydowia 37: 168 (1984)
Other synonyms see Index Fungorum (2020), but Neomassariosphaeria typhicola (P. Karst.) Y. Zhang ter, J.
Fourn. & K.D. Hyde is not accepted as a synonymy in this
study
Possible freshwater distribution: USA (Fallah and
Shearer 2001)
Notes: Massariosphaeria typhicola was transferred
from Leptosphaeria based on some collections ZT 9428,
ZT 9430, ZT 9431 and ZT 9435, and the description and
drawing of Berlese (1894), which were designed from the
original material by Karsten. The holotype of Leptosphaeria
typhicola was not indicated when published and few information of L. typhicola can be obtained in Karsten (1873)
and Berlese (1894). Leuchtmann (1984) mentioned that M.
typhicola showed variability in the shape, size and septation of ascospores. Some authors named several strains CBS
123126, CBS 609.86, KT 667 and KT 797 as M. typhicola
(Leuchtmann 1984; Tanaka and Harada 2004; Zhang et al.
2009c). However, phylogenetic analyses placed CBS 123126
in Amniculicolaceae (Zhang et al. 2009c, 2012b) and KT
667, KT 797, CBS 609.86 in Lindgomycetaceae (Hyde
et al. 2016b; this study, Fig. 2). It is, therefore, considerably important to identify M. typhicola and its relatives with
DNA sequence data. To solve the taxonomic confusion of
M. typhicola, we select strain CBS 609.86 (specimen ZT
9428) designated by Leuchtmann (1984) as its representative materials. The ascospores from a drawing of M. typhicola based on four specimens ZT 9428, ZT 9430, ZT 9431
and ZT 9435, share similar morphology to the new species
13
Fungal Diversity (2020) 105:319–575
Aquimassariosphaeria kunmingensis (Leuchtmann 1984).
We, therefore, place two species in the same genus Aquimassariosphaeria. The examination and description for the
specimen ZT 9428 is expected.
Our phylogenetic analyses show that KT 667 and KT 797
represent a different species with CBS 609.86 (Figs. 2, 61).
Since the very less morphological information of KT 667
and KT 797 in Tanaka and Harada (2004), the real identity
of these two strains are uncertain, only if their specimens
are re-examined in the future. KT 667 and KT 797 probably
represent another new genus, but the establishment needs
evidence from their specimens or new collections with sufficient DNA sequence data. We name KT 667 and KT 797 as
Massariosphaeria sp. in this study. The strain CBS 123126
has been treated as a different species Neomassariosphaeria
typhicola based on morphology and phylogeny in this study
(see notes under Neomassariosphaeria, Amniculicolaceae).
Aquimassariosphaeria typhicola was reported from freshwater habitats in USA with detailed description and illustration (Fallah and Shearer 2001). This USA specimen has
shorter and thinner asci ((93–)113–133 × 11.5–18 μm vs.
100–160 × 15–25 μm) and shorter ascospores (32–36 × 6–8
μm vs. 26–52 × 6–11 μm) than collections of Leuchtmann
(1984). Sequence data of the USA specimen are needed to
confirm if A. typhicola occurred in freshwater environment
in USA.
Key to freshwater Aquimassariosphaeria species
1. Ascospores smooth………A. kunmingensis
1. Ascospores heavily grained sculpted………A. typhicola
Lindgomyces K. Hiray., Kaz. Tanaka & Shearer, Mycologia
102(3): 733 (2010)
Saprobic on submerged woody plant. Sexual morph:
Ascomata scattered to clustered, semi-immersed to nearly
superficial, ellipsoidal or lenticular, subglobose to globose,
uniloculate, glabrous, dark brown to black, with ostiolate
papilla. Peridium composed of hyaline to brown, small, thinwalled cells, sometimes poorly developed at the base. Pseudoparaphyses numerous, cellular, hyaline, septate, branched,
anastomosing. Asci 8-spored bitunicate, fissitunicate, cylindrical to clavate, sessile to minutely pedicellate, rounded at
the apex, with an ocular chamber. Ascospores bi- to tri-seriate, fusiform to cylindrical, straight to slightly curved, hyaline to pale brown, with a median primary septum, smooth
or finely verruculose, thin-walled, usually covered with a
sheath or bearing bipolar mucilaginous appendages. Senescent ascospores brown, > 3-septate (Hirayama et al. 2010).
Asexual morph: Undetermined.
Type species: Lindgomyces ingoldianus (Shearer & K.D.
Hyde) K. Hiray., Kaz. Tanaka & Shearer
Fungal Diversity (2020) 105:319–575
Notes: All Lindgomyces species have been reported from
freshwater habitats (see list below). Lindgomyces was typified by Massarina ingoldiana Shearer & K.D. Hyde, which
is a species complex with some strains assigned to Lindgomyces, and others to Tingoldiago (Hirayama et al. 2010).
The fusiform to cylindrical ascospores with a sheath or bearing bipolar mucilaginous appendages are main characters of
Lindgomyces. Lindgomyces aquaticus sp. nov. is introduced
in this study.
Fig. 57 Lindgomyces spp. (Material examined: USA, Wisconsin,
Adams County, Lemonweir River, on submerged decorticated woody
debris, 31 July 1992, CAS and JLC, A-39-1, ILLS 52289, holotype;
JAPAN, Okinawa, Iriomote, Oomijya River, on submerged decorticated woody debris, 28 September 2007, KH 100; ibid., Aomori,
Hirosaki, Aoki, Mohei-pond, on submerged twigs of woody plant,
3 May 2003, KT 1108, HHUF 28988, holotype; ibid., Hokkaido,
Akkeshi, Ootakita, Sattebetsu River, on submerged wood, 7 September 2003, KT 1399, HHUF 28194, holotype; ibid., Okinawa,
Kunigami, Aha, Tanagakumui, small river, on submerged dead twigs
of woody plant, 19 May 2015, K. Tanaka et al., KT 3531, HHUF
30498, holotype; ibid., Aomori, Nishimeya, Seisyu trail, Ooshirosawa River, on submerged dead twigs of woody plant, 28 August
427
List of freshwater Lindgomyces species
*Lindgomyces angustiascus Raja, A.N. Mill. & Shearer,
Mycoscience 54(5): 357 (2013)
Freshwater distribution: USA (Raja et al. 2013a)
*Lindgomyces apiculatus K. Hiray. & Kaz. Tanaka, Mycologia 103(6): 1424 (2011); Fig. 57g
Freshwater distribution: Japan (Raja et al. 2011)
2010, K. Tanaka et al., KT 2742, HHUF 30513, holotype; ibid.,
Hirosaki, Aoki, Mohei Pond, on submerged woody debris, 7 December 2002, KT 966, HHUF 27883, holotype). a–f L. ingoldianus (a,
c, d from ILLS52289, holotype; b from KH 100; e, f from culture
of ATCC 200398). a Ascomata on the host surface. b Section of
ascoma. c Pseudoparaphyses. d Ascus. e Ascospore with an elongate
sheath. f Ascospore. g Ascospore of L. apiculatus (KT 1108, holotype). h Ascospore of L. breviappendiculatus (culture of KT 1399).
i Ascospore of L. okinawaensis (culture of KT 3531). j Ascospore
of L. pseudomadisonensis (culture of KT 2742). k Ascospore of L.
rotundatus (culture of KT 966). Scale bars: a = 1000 μm, b = 100
μm, c–k = 10 μm
13
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Fungal Diversity (2020) 105:319–575
Fig. 58 Lindgomyces aquaticus
(MFLU 18-1711, holotype).
a, b Appearance of ascomata
on host surface. c Vertical section of ascoma. d Structure of
peridium. e–g Bitunicate asci.
h Ascus embedded in pseudoparaphyses. i–k Ascospores.
l Ascospore in Indian Ink. m
Germinated ascospore. n, o
Colony on PDA (left-front,
right-reverse). Scale bars: c, m
= 50 μm, d–l = 20 μm
*Lindgomyces aquaticus W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557914; Facesoffungi number: FoF09256; Fig. 58
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-1711
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 160–200 μm high, 220–250 μm
wide, black, scattered, semi-immersed to nearly superficial, ellipsoidal or lenticular, coriaceous. Peridium 1.5–2.5
μm, comprising several layers of brown, thin-walled, large
13
cells of textura angularis or subglobose cells, flattened and
poorly developed at the base. Pseudoparaphyses 1.5–2.5
μm diam., sparse, cellular, hypha-like, hyaline, septate, persistent, embedded in a gelatinous matrix. Asci 150–250 ×
33–43 μm ( x̄ = 210 × 39 μm, n = 10), 8-spored, bitunicate,
mostly broadly clavate, rarely narrowly clavate, or cylindrical, apically rounded with well-developed ocular chamber.
Ascospores (59–)68–71 × 12–16 μm ( x̄ = 70 × 14 μm, n =
15), overlapping bi- to tri-seriate, straight or slightly curved,
hyaline and 1-septate when young, slightly constricted at the
septa, turn to pale brown and 5-septate when germinated,
Fungal Diversity (2020) 105:319–575
fusiform with rounded or acute ends, with minute and some
large guttulate, thin-walled, smooth, lacking a mucilaginous
sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 15 mm in 20 days at 25 °C, grey to brown from above,
black from below, surface rough, dry, raised, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat8115-1 (MFLU 18-1711, holotype), ex-type living culture MFLUCC 18-1416; ibid., hat8115-2 (HKAS 105029,
isotype), ex-type living culture KUMCC 19-0044.
Notes: Lindgomyces aquaticus clusters with an unnamed
species Lindgomyces sp. KH241 with high bootstrap support (Figs. 2, 61). Lindgomyces aquaticus differs from
KH241 by its larger asci (150–250 × 33–43 μm vs. 125–194
× 20.5–29 μm), ascospores ((59–)68–71 × 12–16 μm vs.
47–62 × 8–12 μm) and different habitats (freshwater vs.
intertidal) (Hirayama et al. 2010). A large, sticky sheath
is observed in KH241, but not in our collection MFLUCC
18-1416. Comparison of single gene between L. aquaticus
(MFLUCC 18-1416) and Lindgomyces sp. (KH241) shows
that there are one, zero and seven nucleotide differences in
LSU, SSU and ITS sequence data, respectively. Morphological characters mentioned above can easily separate them
to be different species. Lindgomyces aquaticus also differs
from L. ingoldianus (type) in having wider asci (150–250
× 33–43 μm vs. (110–)120–167.5(–182.5) × 25–32.5 μm),
larger ascospores ((59–)68–71 × 12–16 vs. (47–)50–59 ×
9–11(–12)) and without sheath (Hirayama et al. 2010). We
therefore introduce L. aquaticus sp. nov. and retain KH241
as Lindgomyces sp.
*Lindgomyces breviappendiculatus (Kaz. Tanaka, Sat.
Hatak. & Y. Harada) K. Hiray. & Kaz. Tanaka, Mycologia
102(3): 738 (2010); Fig. 57h
Basionym: Lophiostoma breviappendiculatum Kaz.
Tanaka, Sat. Hatak. & Y. Harada, Mycoscience 46(5): 288
(2005)
Freshwater distribution: Japan (Tanaka et al. 2005b;
Hirayama et al. 2010)
*Lindgomyces carolinensis Raja, J. Fourn., Paguigan &
Oberlies, Mycol. Progr. 16(5): 542 (2017)
Freshwater distribution: North Carolina (Raja et al. 2017)
*Lindgomyces cigarosporus Raja, J. Fourn., Paguigan &
Oberlies [as ‘cigarospora’], Mycol. Progr. 16(5): 543 (2017)
Freshwater distribution: North Carolina (Raja et al. 2017)
*Lindgomyces cinctosporus Raja, A.N. Mill. & Shearer [as
‘cinctosporae’], Mycologia 86102(3): 738 (2010)
Freshwater distribution: USA (Hirayama et al. 2010)
429
*Lindgomyces griseosporus Y. Zhang ter, J. Fourn. & K.D.
Hyde, Mycoscience 56(1): 43–48 (2014)
Freshwater distribution: France and Spain (Zhang et al.
2014c)
*Lindgomyces ingoldianus (Shearer & K.D. Hyde) K.
Hiray., Kaz. Tanaka & Shearer, 88Mycologia 102(3): 733
(2010); Fig. 57a–f
Basionym: Massarina ingoldiana Shearer & K.D. Hyde,
Mycologia 89(1): 114 (1997)
Freshwater distribution: Australia (Shearer and Hyde
1997), Brunei (Shearer and Hyde 1997), China (Tsui et al.
2000), Japan (Hirayama et al. 2010), Malaysia (Hyde and
Aptroot 1998b), Thailand (Sivichai et al. 2002), USA
(Shearer and Hyde 1997)
*Lindgomyces lemonweirensis Raja, A.N. Mill. & Shearer,
Mycologia 103(6): 1428 (2011)
Freshwater distribution: USA (Raja et al. 2011)
*Lindgomyces madisonensis Raja & Oberlies, Persoonia,
Mol. Phyl. Evol. Fungi 35: 307 90(2015)
Freshwater distribution: USA (Crous et al. 2015c)
*Lindgomyces okinawaensis Tak. Takah. & Kaz. Tanaka,
Fungal Diversity 78: 35 (2016); Fig. 57i
Freshwater distribution: Japan (Li et al. 2016a)
*Lindgomyces pseudomadisonensis Tak. Takah. & Kaz.
Tanaka, Fungal Diversisty 81 (2016); Fig. 57j
Freshwater distribution: Japan (Hyde et al. 2016b)
*Lindgomyces rotundatus K. Hiray. & Kaz. Tanaka, Mycologia 102(3): 733 (2010); Fig. 57k
Freshwater distribution: Japan and USA (Hirayama et al.
2010)
Key to freshwater Lindgomyces species
1.
1.
2.
2.
3.
3.
4.
4.
Ascospores hyaline, smooth…………………………2
Ascospores grey-brown, smooth to verruculose………
………………………………………L. griseosporus
Ascospores surrounded by a large, expanding gelatinous
sheath…………………………………………………3
Ascospores without expanding sheath………………4
Ascospores fusiform with acute ends…………………
…………………………………………L. ingoldianus
Ascospores cylindrical with rounded ends……………
…………………………………………L. rotundatus
Ascospores with terminal gelatinous appendages…………………………………………………5
Ascospores without terminal gelatinous appendages…………………………………………………10
13
430
5.
5.
6.
6.
7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
Fungal Diversity (2020) 105:319–575
Ascospores < 43 µm long……………………………6
Ascospores > 43 µm long……………………………7
Asci 100–157 × 14–16 μm…………L. madisonensis
Asci 85–125 × 17–25(–27) mm………L. apiculatus
Sheath evanescent……………………L. carolinensis
Sheath not evanescent………………………………8
Ascospores up to 70 μm long, with short, rounded,
terminal ephemeral appendages………L. cigarosporus
Ascospores < 60 μm long……………………………9
Ascospores 47–58 × 9–12 µm………L. angustiascus
Ascospores (40–)44–60 × (9.5–)11–17.5 µm…………
………………………………L. breviappendiculatus
Ascospores < 10 µm wide……L. pseudomadisonensis
Ascospores > 10 µm wide……………………………11
Ascospores without sheath…………………………12
Ascospores with a sheath……………………………13
Ascospores (59–)68–71 × 12–16 μm……L. aquaticus
Ascospores (38–)40–48(–51) × (10–)12–19 µm………
………………………………………L. okinawaensis
Ascospores 40–58 × 10–18 µm, with an amorphous
gelatinous sheath……………………L. cinctosporus
Ascospores 30–44 × 10–15 µm, with an oval, ephemeral sheath…………………………L. lemonweirensis
Neolindgomyces Jayasiri, E.B.G. Jones & K.D. Hyde,
Mycosphere 10(1): 79 (2019)
Saprobic on Pandanus sp or submerged wood. Sexual
morph: Ascomata scattered to gregarious, immersed, depressed
globose or conical with flattened base, dark brown to black,
coriaceous, with a small clypeus ostiole. Ostiole with a porelike opening, central, with a reduced crest, with periphyses
made up of lightly pigmented, pseudoparenchymatous cells.
Peridium composed of dark brown to black, somewhat flattened
cells of textura angularis, fusing and indistinguishable from the
host tissues, with inner stratum comprising hyaline cell layers
of textura angularis. Pseudoparaphyses numerous, trabeculate,
filamentous, hyaline, septate, branched. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate, short pedicellate, apex
rounded with a minute ocular chamber or inconspicuous apical thickening. Ascospores uni- to bi-seriate, overlapping, hyaline, cylindrical or fusiform with narrow, acute ends, septate,
smooth-walled, with a mucilaginous sheath (Hyde and Goh
1999b; Jayasiri et al. 2019). Asexual morph: Undetermined.
Type species: Neolindgomyces pandani Jayasiri, E.B.G.
Jones & K.D. Hyde
Notes: Neolindgomyces was introduced for N. pandani
and a known species Quintaria submersa K.D. Hyde & Goh
collected from freshwater habitats (Hyde and Goh 1999b;
Jayasiri et al. 2019). Neolindgomyces is distinctive in Lindgomycetaceae by its carbonaceous peridium and presence
of clypeus (Jayasiri et al. 2019). It formed a basal clade in
the family with high bootstrap support (Jayasiri et al. 2019),
13
but showed low support in Fig. 2 and moderate support in
Fig. 61.
List of freshwater Neolindgomyces species
*Neolindgomyces submersus (K.D. Hyde & Goh) Jayasiri
& K.D. Hyde [as ‘submersa’] Mycosphere 10(1): 81 (2019)
Basionym: Quintaria submersa K.D. Hyde & Goh, Nova
Hedwigia 68(1-2): 262 (1999)
Freshwater distribution: Australia (Hyde and Goh 1999b)
Key to freshwater asexual genera of Lindgomycetaceae
1. Conidia with apical, subapical and basal appendages…
……………………………………………………Lolia
1. Conidia without appendages……………………………2
2. Conidiogenous cells holoblastic………Clohesyomyces
2. Conidiogenous cells enteroblastic……Hongkongmyces
Clohesyomyces K.D. Hyde, Aust. Syst. Bot. 6(2): 170 (1993)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Conidiomata pycnidial, scattered,
subglobose to ellipsoidal, compressed lenticular, dark brown
to black, semi-immersed to erumpent, unilocular, ostiolate.
Pycnidial wall two-layered. Ostiole dark-brown, circular.
Conidiophores reduced. Conidiogenous cells holoblastic,
determinate, discrete, hyaline, smooth. Conidia variable in
shape, usually ellipsoidal, mostly 1 median-septate, guttulate, hyaline, smooth, thin-walled, straight or slightly curved,
surrounded by a mucilaginous sheath (Hyde 1993b).
Type species: Clohesyomyces aquaticus K.D. Hyde
Notes: Clohesyomyces was introduced to accommodate
a single species C. aquaticus collected from a freshwater
habitat in Australia (Hyde 1993b). Other collections were
reported from freshwater habitats in China (Cai et al. 2006b;
this study) and Thailand (Zhang et al. 2012a; this study).
The phylogenetic position of Clohesyomyces was confirmed
in Lindgomycetaceae (Zhang et al. 2012a; Raja et al. 2017)
and this study (Fig. 2). The freshwater materials of C. aquaticus collected by Zhang et al. (2012a) was re-examined and
described by Li et al. (2020) and this study (Fig. 59). We
also report two additional freshwater collections obtained
from China and Thailand (Fig. 59).
List of freshwater Clohesyomyces species
*Clohesyomyces aquaticus K.D. Hyde, Aust. Syst. Bot.
6(2): 170 (1993)
Facesoffungi number: FoF07165; Fig. 59
Freshwater distribution: Australia (Hyde 1993b), China
(Cai et al. 2006b; this study), Thailand (Zhang et al. 2012a;
this study)
Saprobic on submerged wood in freshwater habitats. Sexual morph: Undetermined. Asexual morph: Conidiomata
Fungal Diversity (2020) 105:319–575
431
Fig. 59 Clohesyomyces aquaticus (a–i from MFLU 18-1159, j–m
from MFLU 15-2699, n–o from MFLU 11-1112). a, b Appearance of
pycnidia on host surface. c Vertical section of pycnidium. d Ostiole.
e Peridium. f, g Conidiogenous cells with conidia. h, i Conidia. j–m,
o Conidia. (m, o in Indian Ink). n Conidiogenous cell with conidium.
Scale bars: c = 100 μm, d, e = 20 μm, f–o = 10 μm
pycnidial, scattered, 160–300 μm high, 180–850 μm diam.,
subglobose to ellipsoidal, compressed lenticular, dark brown
to black, semi-immersed to erumpent, unilocular, ostiolate.
Ostiole dark brown, central, circular. Peridium 25–60 μm
thick, composed of two layers, with outer wall comprising
several layers of dark brown cells, inner wall comprising 3–5
layers of brown-walled rectangle cells of textura porrecta.
Conidiophores reduced. Conidiogenous cells holoblastic, up
to 20 μm long, determinate, discrete, cylindrical to subcylindrical, hyaline, smooth, forming from the inner layer cells
of the pycnidial wall. Conidia 16–29 × 7–16 μm ( x̄ = 21.1
× 8.7 μm, n = 10), cylindrical to subcylindrical or ellipsoid,
irregular, mostly 1-septate, slightly constricted at the septum, one prominent guttulate in each cell, some with additional small guttules, hyaline, smooth, thin-walled, truncate
at the base, obtuse at the apex, straight or slightly curved,
surrounded by an irregular, thin mucilaginous sheath.
Culture characteristics: On PDA, colony circular, slow
growing, 40 mm in 30 days at 25 °C, brown to grey from
above, brown to yellow from reverse, raised, felty wooly,
fairly dense, initially aerial with an smooth edge, becoming
rough after 20 days, staining agar yellow.
Material examined: THAILAND, Prachuap Khiri Khan
Province, on submerged wood in a small river, 30 July 2015,
13
432
K.D. Hyde, 13A (MFLU 15-2699), living culture MFLUCC
15-0979; CHINA, Yunnan Province, on submerged wood
in a small river, 25 November 2017, G.N. Wang, H58A-1
(MFLU 18-1159), living culture MFLUCC 18-1037; ibid.,
H58A-2 (HKAS 101735), living culture KUMCC 18-0087.
Additional material examined: THAILAND, Chiang Mai
Province, Doi Inthanon, on submerged wood, 16 November
2010, H. Zhang, d66 (MFLU 11-1112).
Notes: Our new collections MFLUCC 15-0979 and
MFLUCC 18-1037 are identified as Clohesyomyces aquaticus based on identical LSU and ITS sequence data with
MFLUCC 11-0092, and phylogenetic analysis (Figs. 2, 61).
The conidial size of C. aquaticus varies in different collections, ranging from 15–31 × 7–16 μm. Moreover, MFLUCC
18-1037 has larger diameter of conidiomata than the holotype BRIP 20092 (260–300 μm high, 800–850 μm diam. vs.
247–390 μm high, 156–260 μm diam.) (Hyde 1993b; Zhang
et al. 2012a).
Hongkongmyces C.C.C. Tsang et al., Medical Mycol. 52(7):
740 (2014)
(see entry below freshwater sexual genera of
Lindgomycetaceae)
Lolia Abdel-Aziz & Abdel-Wahab, Mycotaxon 114: 36
(2011)
Saprobic on submerged decayed stem. Sexual morph:
Ascomata solitary or aggregated in stroma, immersed to
erumpent, uniloculate, globose to subglobose, sometimes
with flattened base, glabrous, dark brown to black, coriaceous to sub-carbonaceous, with cylindrical ostiolar canals,
with ostiolate papilla, usually stain the substrate purple.
Peridium two-layered, comprising hyaline to yellow-brown,
to dark brown, thin-walled cells of textura angularis. Pseudoparaphyses numerous, cellular, thin, hypha-like, septate,
branched, persistent, emerged into the ostiolar canal. Asci 4to 8-spored, bitunicate, fissitunicate, cylindric-clavate, pedicellate, with ocular chamber. Ascospores uni- to bi-seriate,
fusiform to clavate, with rounded ends, straight or slightly
curved, yellow- to reddish- to dark-brown, transversely septate, occasionally with one longitudinal or oblique septum,
smooth- or rough-walled, surrounded by prominent gelatinous sheath (Abdel-Aziz 2016a). Asexual morph: Coelomycetous. Conidiomata solitary or aggregated, superficial,
acervular, pearl white to dull yellow. Peridium composed
of hyaline cells of textura intricata, embedded in gel. Conidiophores lining the acervuli wall, branched and septate,
hyaline, smooth, embedded in gel. Conidiogenous cells
holoblastic, terminal, cylindrical to sub-cylindrical, hyaline, smooth, bearing a single conidium. Conidia solitary,
ellipsoidal, straight, hyaline, aseptate, smooth, thin-walled,
with apical, sub-apical and basal appendages (Abdel-Aziz
and Abdel-Wahab 2010).
Type species: Lolia aquatica Abdel-Aziz & Abdel-Wahab
13
Fungal Diversity (2020) 105:319–575
Notes: Lolia was introduced by Abdel-Aziz and AbdelWahab (2010) to accommodate the asexual species L.
aquatica collected on decayed stem of Phragmites australis from Egypt. Lolia is characterized by acervular conidiomata, clavate, ellipsoidal, cylindrical conidia with apical,
sub-apical and basal appendages (Abdel-Aziz and AbdelWahab 2010). Its sexual morph was later collected on submerged decayed stem of Phragmites australis from River
Nile (Abdel-Aziz 2016a). The authors assigned two holotypes, IMI 398675 and CBS H-22130, based on both asexual
and sexual morphs, respectively, which does not conform
to “One fungus, one name” (Abdel-Aziz and Abdel-Wahab
2010; Abdel-Aziz 2016a). We select the first holotype (IMI
398675, ex-type living culture MF644) as the exclusive
holotype.
The second species L. dictyospora Abdel-Aziz was also
collected from River Nile, but on submerged decayed rachis
of Phoenix dactylifera (Abdel-Aziz 2016a). The special
characters of L. dictyospora, such as 4- to 8-spored asci and
clavate ascospores occasionally with one longitudinal or
oblique septa, distinguish it from L. aquatica (Abdel-Aziz
2016a). Lolia aquatica has cellular, hypha-like pseudoparaphyses, while L. dictyospora has trabeculate, filiform
pseudoparaphyses. Two species cluster together, but without bootstrap support (Abdel-Aziz 2016a; this study, Figs. 2,
61).
List of freshwater Lolia species
*Lolia aquatica Abdel-Aziz & Abdel-Wahab, Mycotaxon
114: 36 (2011); Fig. 60a–j
Freshwater distribution: Egypt (Abdel-Aziz and AbdelWahab 2010; Abdel-Aziz 2016a)
*Lolia dictyospora Abdel-Aziz, Phytotaxa 267(4): 283
(2016); Fig. 60k–n
Freshwater distribution: Egypt (Abdel-Aziz 2016a)
Key to freshwater Lolia species
1. Asci 8-spored, ascospores 26–37 × 6–9 μm……………
………………………………………………L. aquatica
1. Asci 4–8-spored, ascospores 14–19 × 4–7 μm…………
…………………………………………L. dictyospora
Longipedicellataceae Phukhams., Bhat & K.D. Hyde,
Mycosphere 7(11): 1722 (2016)
Key to freshwater genera of Longipedicellataceae
1. Sexual morph…………………………Longipedicellata
1. Asexual morph…………………………………………2
Fungal Diversity (2020) 105:319–575
433
Fig. 60 Lolia spp. (Material examined: EGYPT, Sohag, El Balyana
city, on decayed stem of Phragmites australis (Cav.) Steud. at irrigation canal, March 2005, F.A. Abdel-Aziz, IMI 398675, holotype;
Sohag, River Nile, on submerged decayed stem of Phragmites australis (Poaceae), 14 August 2012, F.A. Abdel-Aziz, CBS H-22130;
Sohag, River Nile, on submerged decayed rachis of Phoenix dactylifera
(Arecaceae), 14 August 2012, F.A. Abdel-Aziz, CBS H-22131). a–e
Sexual morph of Lolia aquatica (CBS H-22130). a Vertical section
of ascoma. b, c Bitunicate asci. d Ascospore with a large gelatinous
sheath. e Ascospore in Indian Ink. f–j Asexual morph of L. aquatica
(IMI 398675, holotype). f–g Vertical section through the gelatinous
acervular (f in phase contrast). h–j Conidia. k–n L. dictyospora (CBS
H-22131, holotype). k–m Asci. n Ascospores surrounded by a large
sheath. Scale bars: a, f, g = 100 μm, b, c = 30 μm, d, e, h–n = 10 μm
1. Conidiophores semi-macronematous, slender, conidiogenous cells cuneiform, conidia variable in shape………
…………………………………………Submersispora
1. Conidiophores semi-macronematous or reduced, conidiogenous cells reduced or absent, conidia broadly
fusiform………………………………Pseudoxylomyces
Longipedicellata H. Zhang, K.D. Hyde & J.K. Liu, Phytotaxa 247(2): 102 (2016)
Saprobic on dead and submerged woody materials. Sexual morph: Ascomata scattered or clustered, immersed,
beneath a blackened pseudoclypeus, lenticular, uniloculate,
glabrous, brown to black, coriaceous, sometimes ostiolate. Ostiole relatively large, central, short, dark. Peridium
thin, comprising several layers of brown, thin-walled,
flattened cells of textura angularis, outer layer somewhat
13
434
Fig. 61 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS, TEF and RPB2
sequence data for species of
Lindgomycetaceae. Bootstrap
values for maximum likelihood equal to or greater than
70% and Bayesian posterior
probabilities equal to or greater
than 0.95 are placed near the
branches as ML/BYPP. Newly
generated sequences are in red
and ex-type strains are in bold.
The new species introduced in
this study are indicated with
underline. Freshwater strains are
indicated with a red letter “F”.
The tree is rooted to Fissuroma
maculans (MFLUCC10-0886
and MFLUCC10-0887) (Aigialaceae)
Fungal Diversity (2020) 105:319–575
99/1.00
Lolia aquatica MF 644
Lolia aquatica CBS 22130 F
Lolia
Lolia dictyospora CBS 22131 F
73/0.95
Aquimassariosphaeria kunmingensis KUMCC 18-1019 F
Aquimassariosphaeria typhicola CBS 609.86
Aquimassariosphaeria
Massariosphaeria sp. KT 667
Massariosphaeria sp. KT 797
Clohesyomyces aquaticus MFLUCC 18-1037 F
99/1.00
Clohesyomyces aquaticus MFLUCC 11-0092 F
Clohesyomyces
Clohesyomyces aquaticus MFLUCC 15-0979 F
91/0.99
96/0.99
99/1.00
Trematosphaeria hydrela CBS 880.70
Trematosphaeria hydrela HKUCC 10666
Hongkongmyces pedis HKU 35
Hongkongmyces snookiorum DAOMC 251900 F
73/1.00
100/1.00
100/1.00
Lindgomyces ingoldianus KH 100 F
Lindgomyces ingoldianus ATCC 200398 F
Lindgomyces sp. KH 241 F
73/--
100/0.99
Hongkongmyces
Lindgomyces
Lindgomyces aquaticus MFLUCC 18-1416 F
100/1.00
Neolindgomyces submersa CBS 115553
Neolindgomyces pandanae MFLUCC 18-0245
100/1.00
Arundellina typhae MFLUCC 16-0310 F
Arundellina typhae MFLUCC 16-0309
100/1.00
Neolindgomyces
Arundellina
Fissuroma maculans MFLUCC10-0886
Fissuroma maculans MFLUCC 10-0887
Outgroup
0.04
carbonaceous, easily cracked. Pseudoparaphyses absent,
few or numerous, cellular, hyphae-like, hyaline, sparsely
septate. Asci 8-spored, bitunicate, clavate, long pedicellate,
with an ocular chamber. Ascospores bi- to tri-seriate, broadly
fusiform to ellipsoidal, hyaline, 1-septate, slightly asymmetrical, the upper cell often shorter and broader than the
lower one, smooth, thin-walled, surrounded by mucilaginous
sheath (Zhang et al. 2016). Asexual morph: Hyphomycetous. Mycelium composed of hyaline to dark brown, septate,
branched, smooth hyphae. Conidiophores and conidiogenous
cells unknown. Conidia chlamydosporous monilioid, in long
acropetal, branched chains, catenate, doliiform, subglubose
to oval, brown to reddish-brown, rough-walled (Phukhamsakda et al. 2016) (Fig. 61).
Type species: Longipedicellata aptrootii (K.D. Hyde &
S.W. Wong) H. Zhang, K.D. Hyde & J.K. Liu
Notes: Longipedicellata was introduced in Bambusicolaceae to accommodate Didymella aptrootii K.D. Hyde &
S.W. Wong based on combined LSU, SSU, RPB2 and TEF
sequence data (Zhang et al. 2016). However, it clustered in
Bambusicolaceae with low bootstrap support (Zhang et al.
2016). Phukhamsakda et al. (2016) re-collected this species from Thailand and showed a close relationship between
Pseudoxylomyces elegans (Goh et al.) Kaz. Tanaka & K.
Hiray. and Longipedicellata aptrootii which represented a
new family Longipedicellataceae. They also reported the
asexual morph of L. aptrootii which formed in the culture.
We report L. aquatica sp. nov. in this study.
List of freshwater Longipedicellata species
13
*Longipedicellata aptrootii (K.D. Hyde & S.W. Wong)
H. Zhang, K.D. Hyde & J.K. Liu, Phytotaxa 247(2): 104
(2016); Fig. 62
Basionym: Didymella aptrootii K.D. Hyde & S.W. Wong,
Australas. Mycol. 18 (3): 54 (1999)
Freshwater distribution: China (Hyde and Wong 1998),
Malaysia (Hyde and Wong 1998), Philippines (Hyde and
Wong 1998; Cai et al. 2003a), Thailand (Kurniawati et al.
2010; Zhang et al. 2016; this study)
Saprobic on submerged wood in freshwater. Sexual
morph: Ascomata 150–170 μm high, 190–220 μm diam.,
immersed or semi-immersed, scattered or clustered, lenticular, covered with a blackened pseudoclypeus, black,
coriaceous, ostiolate. Peridium 10–15 μm thick, comprising several layers of brown, thin-walled, flattened cells
of textura angularis. Pseudoparaphyses 2–2.5 μm wide,
numerous, cellular, hyphae-like, hyaline, sparsely septate,
unbranched. Asci 80–115 × 19–26 μm (x̄ = 95 × 22 μm, n =
10), 8-spored, bitunicate, fissitunicate, broadly clavate, with
a long, straight or twisted pedicel and an ocular chamber.
Ascospores 17.5–22 × 8–8.5 μm (x̄ = 19 × 8 µm, n = 15),
bi- to tri–seriate, ellipsoidal to broadly fusiform, rounded
at both ends, hyaline, 1-septate, constricted at the septum,
the upper cell slightly broader and shorter than the lower
cell, guttulate, thin-walled, surrounded by a large, irregular, mucilaginous sheath, 10–15 μm wide. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 15 mm in 30 days at 25 °C, black in inner layer and white
Fungal Diversity (2020) 105:319–575
435
Fig. 62 Longipedicellata
aptrootii (a–c from MFLU
10-0162, d–k from MFLU
18-1000). a Ascomata on the
host surface. b Section of
ascoma. c Peridium. d–g Asci.
h–k Ascospores. k Ascospores
in Indian Ink. Scale bars: b =
50 μm, c, h–j = 10 μm, d–g, k
= 20 μm
in outer layer from above, black from below, surface fluffy,
with fairly dense mycelium, raised, with a smooth edge.
Material examined: THAILAND, Phayao Province, on
submerged wood in a stream, 23 February 2018, X.D. Yu,
Y22 (MFLU 18-1000), living culture MFLUCC 18-0988.
Additional material examined: THAILAND, Chiang
Rai Province, on submerged bamboo, 18 January 2010, H.
Zhang (MFLU 10-0162).
Notes: Our new collection MFLUCC 18-0988 is identified as Longipedicellata aptrootii based on their identical
LSU and ITS sequence data between MFLUCC 18-0988
and MFLUCC 10-0297, and phylogenetic analysis (Figs. 2,
68). Our collection has wider asci (80–115 × 19–26 μm vs.
70–100 × 17–20 μm) than the reference specimen in Zhang
et al. (2016). We found numerous pseudoparaphyses in our
collection, which were not mentioned in Zhang et al. (2016).
*Longipedicellata aquatica W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557915; Facesoffungi number: FoF09257; Fig. 63
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 17-1686
Saprobic on submerged wood in freshwater. Sexual
morph: Ascomata 150–200 μm high, 200–250 μm diam.,
lenticular or subglobose, immersed or semi-immersed, covered with a blackened pseudoclypeus, black, with mass of
grey-white ascospores pouring out when mature, scattered
or clustered, coriaceous, ostiolate. Peridium 5–10 μm thick,
13
436
Fungal Diversity (2020) 105:319–575
Fig. 63 Longipedicellata
aquatica (MFLU 17-1686,
holotype). a Appearance of
black ascomata on host. b
Vertical section of ascoma.
c Structure of peridium. d–g
Bitunicate asci. (g Elongate asci
with pseudoparaphyses). h–k
Ascospores. l, m Ascospores
in Indian Ink. n Geminated
ascospore. o, p Colony on PDA
(left-front, right-reverse). Scale
bars: b = 50 μm, c–g, k, l, n =
20 μm, h–j, m = 10 μm
comprising several layers of brown to black, thin-walled
cells of textura angularis. Pseudoparaphyses sparse, cellular, hyphae-like, hyaline, sparsely septate. Asci 85–120
× 22–27 μm ( x̄ = 100 × 24 μm, n = 20), 8-spored, bitunicate, fissitunicate, narrowly to broadly clavate, elongate
soon when released in water, up to 180 μm long, with a
long, straight or twisted pedicel and an ocular chamber.
Ascospores 19–22 × 8.5–10.5 μm (x̄ = 2 × 9.5 µm, n = 20),
bi- to tri-seriate, ellipsoidal to broadly fusiform, rounded
at both ends, obtuse, hyaline, 1-septate, constricted at the
13
septum, the upper cell slightly broader and shorter than
the lower cell, guttulate, with two prominent guttulate in
each cell, becoming one big guttulate when mature, thinwalled, surrounded by a large, obvious, ellipsoidal, mucilaginous sheath, 7–27 μm wide, deeply constricted near the
ascospores septa. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony filamentous,
reaching 20 mm in 15 days at 25 °C, black with brown margin from above, black from below, surface fluffy, with aerial
mycelium, raised, with a smooth edge.
Fungal Diversity (2020) 105:319–575
Material examined: THAILAND, Chiang Mai Province,
Phrao District, on submerged wood in a stream, 1 September
2017, G.N. Wang, g7 (MFLU 17-1686, holotype), ex-type
living culture MFLUCC 17-2334; Nakhon Phaanon Province, on submerged decaying wood of unidentified plants, 12
December 2018, H. Yang, T12 (MFLU 19-0530), living culture MFLUCC 19-0324; Mukdahan Province, on submerged
decaying wood of unidentified plants, 13 December 2018,
H. Yang, T45 (MFLU 19-0551), living culture MFLUCC
19-0340.
Notes: Longipedicellata aquatica clusters with L. aptrootii with high bootstrap support (Figs. 2, 68). The asci
of L. aquatica quickly become elongate when released in
water, up to 180 μm long, but not in L. aptrootii (Zhang
et al. 2016). Additionally, their ascospore sheaths are different (large, obvious, ellipsoidal, mucilaginous sheath, deeply
constricted near the ascospore septa in L. aquatica vs. large,
irregular, mucilaginous sheath in L. aptrootii). Longipedicellata aquatica is more similar to one collection of L. aptrootii
(MFLU 16–0032) in having similar ascal size, but differs in
having shorter ascospores (19–22 × 8.5–10.5 μm vs. 19–26
μm × 7–11 μm) (Phukhamsakda et al. 2016). After comparision of their single genes, there are three and 21 nucleotide differences in LSU and ITS sequence data, between
L. aquatica MFLUCC 17-2334 and L. aptrootii MFLUCC
10-0297, respectively, which indicates that they are distinct
species (Jeewon and Hyde 2016). Longipedicellata aquatica
sp. nov. is therefore introduced here.
Key to freshwater Longipedicellata species
1. Asci 70–100 × 17–20 μm, ascospores surrounded by
irregular sheath……………………………L. aptrootii
1. Asci 85–120 × 22–27 μm, ascospores surrounded by
ellipsoidal sheath, which is deeply constricted near the
ascospores septa……………………………L. aquatica
Pseudoxylomyces Kaz. Tanaka & K. Hiray., Stud. Mycol.
82: 126 (2015)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum,
scattered to gregarious, dark brown to black, glistening.
Mycelium mostly immersed, pale brown to reddish brown.
Stromata lacking. Conidiophores septate, brown, branched
or not developed. Conidiogenous cells holoblastic. Conidia
single, broadly fusiform or ellipsoidal, straight, with several
transverse septa of thick-walled, yellowish brown to dark
brown, with paler end cells, without sheath or appendages
(Tanaka et al. 2015).
Type species: Pseudoxylomyces elegans (Goh, W.H. Ho,
K.D. Hyde & C.K.M. Tsui) Kaz. Tanaka & K. Hiray.
437
Notes: Pseudoxylomyces was proposed to accommodate
Xylomyces elegans Goh et al., which was isolated from submerged wood in Australia (Goh et al. 1997; Tanaka et al.
2015). Xylomyces is characterized as lacking conidiophores
and conidiogenous cells, but has large chlamydospores
(Goos et al. 1977; Goh et al. 1997). In contrast, Pseudoxylomyces is quite distinctive in producing broadly fusiform
conidia at the tip of the conidiophores (Tanaka et al. 2015).
One new species P. aquaticus is introduced in Pseudoxylomyces in this study.
List of freshwater Pseudoxylomyces species
*Pseudoxylomyces aquaticus W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557916; Facesoffungi number: FoF09258; Fig. 64
Etymology: referring to aquatic habitat of this fungus
Holotype: HKAS 100937
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
scattered or gregarious, effuse, black. Mycelium mostly
immersed in natural substratum, hyaline to pale brown. Conidiophores reduced or absent. Conidiogenesis holoblastic.
Conidia 37–52 × 13–17 µm ( x̄ = 43 × 15 µm, n = 20),
solitary, dry, broadly fusiform, with 5 thick septa, distinctly
constricted at the septa, brown, sometimes the two central
cells swollen, guttulate when young, sometimes pale brown
at bipolar cells which are easily wizened when old, roughwalled, verruculose, straight, thick-walled.
Culture characteristics: On PDA, colony circular, reaching 20 mm in 20 days at 25 °C, pale brown to white from
above and below, surface rough, with sparse mycelium, dry,
edge entire.
Material examined: THAILAND, Chiang Mai Province,
Phrao District, on submerged wood in a stream, 1 September
2017, G.N. Wang, 4.14 (HKAS 100937, holotype), ex-type
living culture KUMCC 17-0312.
Notes: Pseudoxylomyces aquaticus shares similar characters with the type species P. elegans in having broadly fusiform conidia with thick septa (Goh et al. 1997; Tanaka et al.
2015). However, P. aquaticus has smaller conidia (37–52 ×
13–17 μm vs. 66–104 × (19–)26–40 μm), fewer and stable
number of septa (5-septate vs. 4–7-septate) and different
pigmentation (brown vs. yellowish brown to orange brown)
(Goh et al. 1997; Tanaka et al. 2015). Phylogenetic analysis
supports them to be different species (Figs. 2, 68).
*Pseudoxylomyces elegans (Goh et al.) Kaz. Tanaka & K.
Hiray., Stud. Mycol. 82: 126 (2015)
Facesoffungi number: FoF09259; Figs. 65, 66
Basionym: Xylomyces elegans Goh, W.H. Ho, K.D. Hyde
& K.M. Tsui, Mycol. Res. 101(11): 1324 (1997)
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Fig. 64 Pseudoxylomyces
aquaticus (HKAS 100937,
holotype). a Colonies on
submerged wood. b–g Conidia.
h Germinated conidium. i, j
Colony on PDA (left-front,
right-reverse). Scale bars: b–g
= 10 μm, h = 20 μm
Freshwater distribution: Australia (Goh et al. 1997),
Brazil (Barbosa and Gusmão 2011), China (Tsui and Hyde
2004), India (Patil and Borse 2015), Japan (Tanaka et al.
2015), Seychelles (Goh et al. 1997), Thailand (Sivichai et al.
2000; this study), USA (Raja et al. 2007)
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
gregarious or scattered, effuse, black, glistening. Mycelium
partly immersed in natural substratum, composed of hyaline
to pale brown, septate hyphae. Conidiophores up to 70 µm
long, semi-macronematous, mononematous, unbranched,
septate, not constricted at the septa, smooth, hyaline to
pale brown to brown, thin-walled. Conidiogenous cells
16–20 × 3.5–4.5 µm, holoblastic, monoblastic, integrated,
13
determinate, terminal, cylindrical, hyaline to pale brown,
smooth-walled. Conidia (70–)80–90(–100) × 29–37(–40.5)
µm (x̄ = 87.5 × 35 µm, n = 15), solitary, dry, mostly broadly
ellipsoidal with suddenly tapering ending cells, 5–6(–7)
thick septate, distinctly constricted and darker at the septa,
two central cells mostly swollen and greatly enlarged, which
almost occupy 1/2–3/5 of the whole conidia, reddish brown
to dark brown, guttulate when young, straight or slightly
curved, minutely rough-surfaced, thick-walled; ending cells
long or short, subcylindrical, paler, rounded and easily wizened when old.
Culture characteristics: On PDA, colony circular, reaching 30 mm in 35 days at 25 °C, reddish brown to pale grey
from above, reddish brown from below, surface rough, with
Fungal Diversity (2020) 105:319–575
439
Fig. 65 Pseudoxylomyces elegans (MFLU 17-1712). a–c Colonies on submerged wood. d Conidiophore bearing conidium. e–k Conidia. l Germinated conidium. m, n Colony on PDA (left-front, right-reverse). Scale bars: d = 20 μm, e–k = 50 μm, l = 100 μm
sparse mycelium, mostly immersed in culture, dry, edge
entire.
Material examined: THAILAND, Nan Province, on
submerged wood in a stream, 4 August 2017, Saranyaphat
Boonmee, DP8-1 (MFLU 17-1712), living culture MFLUCC
17-2350; ibid., DP8-2, living culture MFLUCC 17-2351.
Additional material examined: JAPAN, Okinawa, Isl. Iriomote, Oomijya-river, on submerged twigs of woody plant,
12 July 2011, K. Tanaka & K. Hirayama, KT 2887 = HHUF
30139, culture MAFF 243852.
Notes: Our collection MFLUCC 17-2350 clusters with
Pseudoxylomyces elegans (KT 2887) with strong bootstrap
support (Figs. 2, 68). They share similar morphology in
conidial size, pigmentation and septa (Goh et al. 1997; Tanaka et al. 2015). Comparing individual nucleotides between
MFLUCC 17-2350 and KT 2887, we noticed 100% similarities in LSU sequence data, nine and eight nucleotide
differences in ITS and TEF sequence data, respectively.
Based on their very similar morphology and no resolution
in the phylogenetic tree (Figs. 2, 68), MFLUCC 17-2350 is,
therefore, identified as P. elegans. The conidia of P. elegans
are smooth-walled in HKU (M) 3242 (Goh et al. 1997),
we observed the rough-walled conidia in our collection
MFLUCC 17-2350. The conidia in our collection mostly
have two swollen and greatly enlarged, central cells, which
are not much obvious in HKU (M) 3242 and KT 2887. The
strain SS1077 clusters basal to P. elegans with high bootstrap support (Fig. 68). SS1077 is named as P. elegans in
Prihatini et al. (2008), the real identity of SS1077 is, however, better confirmed until more evidences are obtained.
We name SS1077 as Pseudoxylomyces sp. as its morphology
is unavailable and only ITS sequence data is deposited in
GenBank.
Key to freshwater Pseudoxylomyces species
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Fig. 66 Pseudoxylomyces elegans (MAFF 243852 = KT 2887). a, b Asexual morph produced on agar medium. c Conidiophores and conidia. d,
e Conidia. Scale bars: a = 1 mm, b = 200 µm, c = 100 µm, d, e = 20 µm
1. Conidia 37–52 × 13–17 µm………………P. aquaticus
1. Conidia 66–104 × (19–)26–40 μm…………P. elegans
Submersispora W. Dong, H. Zhang & K.D. Hyde, gen. nov.
Index Fungorum number: IF557805; Facesoffungi number: FoF08719
Etymology: referring to submerged habitat of this fungus
Saprobic on decaying, submerged wood in freshwater. Sexual morph: Undetermined. Asexual morph: Colonies broadly
punctiform, gregarious or scattered or solitary, raised, black.
Mycelium mostly immersed in natural substratum, consisting of branched, septate, thin-walled, smooth, hyaline to pale
brown hyphae. Conidiophores semi-macronematous, mononematous, cylindrical, slender, unbranched, septate, smooth,
pale brown, thin-walled. Conidiogenous cells monoblastic,
holoblastic, integrated, determinate, cuneiform, pale brown,
smooth. Conidia acrogenous, solitary, dry, variable in shape,
septate, brown to dark black, smooth, thin-walled.
Type species: Submersispora variabilis W. Dong, H.
Zhang & K.D. Hyde
Notes: Two freshwater genera Longipedicellata and Pseudoxylomyces are accommodated in Longipedicellataceae
(Goh et al. 1997; Tanaka et al. 2015; Phukhamsakda et al.
2016; Zhang et al. 2016). Phylogenetic analyses show that
our collection MFLUCC 17-2360 nests between Longipedicellata and Pseudoxylomyces with strong bootstrap support
(Figs. 2, 68). Both Longipedicellata and Pseudoxylomyces
have chlamydospore-like structures or chlamydospores
(Goh et al. 1997; Phukhamsakda et al. 2016). Our collection
13
resembles Pseudoxylomyces in having holoblastic conidiogenous cells, but differs in having slender, semi-macronematous
conidiophores and variable shaped conidia. Thus, the third
freshwater genus Submersispora is introduced in this study.
List of freshwater Submersispora species
*Submersispora variabilis W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557917; Facesoffungi number: FoF08720; Fig. 67
Etymology: referring to variable shape of conidia
Holotype: MFLU 17-1719
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
broadly punctiform, gregarious or scattered or solitary, raised,
black. Mycelium mostly immersed in natural substratum,
consisting of branched, septate, thin-walled, smooth, hyaline
to pale brown hyphae. Conidiophores up to 230 µm long, 3
µm wide, semi-macronematous, mononematous, cylindrical,
slender, unbranched, septate, slightly constricted at the septa,
smooth, pale brown, thin-walled. Conidiogenous cells 5.2–6.4
× 5.5–6 µm (x̄ = 5.8 × 5.7 µm, n = 5), monoblastic, holoblastic, integrated, determinate, cuneiform, pale brown, smooth.
Conidia 26.5–185.5 × 35–90 µm (x̄ = 96.5 × 65.7 µm, n =
30), acrogenous, solitary, dry, variable in shape, mostly subglobose, ellipsoidal, oblong, irregular, septate, muriform, slightly
constricted at the septa, comprising large number of angular
cells, hyaline to brown when young, dark black and invisible
for septa when mature, smooth, thin-walled.
Fungal Diversity (2020) 105:319–575
441
Fig. 67 Submersispora variabilis (MFLU 17-1719, holotype).
a Colonies on submerged wood.
b Conidiophores with conidia.
c Conidium, conidiogenous cell
and conidiophore. d–m Conidia.
n Germinated conidium. o, p
Colony on PDA (left-front,
right-reverse). Scale bars: b, f–n
= 50 μm, c–e = 20 μm
Culture characteristics: On PDA, colony circular, reaching 30 mm in 15 days at 25 °C, grey in the middle with dark
brown margin from above, dark brown from below, surface
rough, fluffy, with dense mycelium, dry, raised, edge entire.
Material examined: THAILAND, Chiang Mai Province,
MRC, on submerged wood in a stream, 1 September 2017,
X.D. Yu, 2A (MFLU 17-1719, holotype), ex-type living
culture MFLUCC 17-2360.
Lophiostomataceae Sacc. [as ‘Lophiostomaceae’], Syll.
fung. (Abellini) 2: 672 (1883)
Key to freshwater genera of Lophiostomataceae
1. Ascospores pigmented………………………………2
1. Ascospores hyaline……………………………………4
2. Ascospores evenly coloured, yellowish brown to dark
brown…………………………………………………3
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Fig. 68 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS and TEF sequence
data for species of Longipedicellataceae. Bootstrap values for
maximum likelihood equal to or
greater than 75% and Bayesian
posterior probabilities equal to
or greater than 0.95 are placed
near the branches as ML/BYPP.
Newly generated sequences are
in red and ex-type strains are
in bold. The new species introduced in this study are indicated
with underline. Freshwater
strains are indicated with a red
letter “F”. The tree is rooted
to Bambusicola bambusae
MFLUCC11-0614 (Bambusicolaceae)
82/1.00
90/1.00
100/1.00
Longipedicellata aptrootii MFLUCC 18-0988 F
Longipedicellata aptrootii MFLUCC 16-0244
Longipedicellata aptrootii MFLUCC 10-0297 F
Longipedicellata aptrootii MFLUCC 16-0384
100/1.00
Longipedicellata
Longipedicellata aquatica MFLUCC 19-0340 F
Longipedicellata aquatica MFLUCC 17-2334 F
100/1.00
100/1.00
Longipedicellata aquatica MFLUCC 19-0324 F
Submersispora variabilis MFLUCC 17-2360 F
99/0.99
100/1.00
Submersispora
Pseudoxylomyces elegans KT 2887 F
Pseudoxylomyces elegans MFLUCC 17-2350 F
Pseudoxylomyces
Pseudoxylomyces sp. SS1077
Pseudoxylomyces aquaticus KUMCC 17-0312 F
Bambusicola bambusae MFLUCC 11-0614
Outgroup
0.009
2. Ascospores pale yellowish to olivaceous brown, but pale
at the terminal cells………………Neotrematosphaeria
3. Ascospores fusiform, often with hyaline appendages…
……………………………………Biappendiculispora
3. Ascospores cigar-shaped or ellipsoidal-fusiform, sometimes with terminal appendages……………Sigarispora
4. Ascospores with longest sheath drawn out to
form polar appendages with tips of the sheath
capped……………………………Pseudocapulatispora
4. Ascospores appendages not as above…………………5
5. Ascospores with a mucilaginous collar around its equator, surrounded by a spreading papilionaceous sheath…
…………………………………………Vaginatispora
5. Ascospores without a spreading papilionaceous
sheath…………………………………………………6
6. Ascospores with globose appendages at both ends……
………………………………………Neovaginatispora
6. Ascospores without globose appendages………………7
7. Ascospores with an internal chamber and a narrow
sheath drawn out at both ends……………Crassiclypeus
7. Ascospores without an internal chamber or a sheath…8
8. Ascomata sometimes with short papilla……Lentistoma
8. Ascomata often with crest-like apex……………………9
9. Ascospores with a narrow, bipolar sheath and coelomycetous asexual morph……………………Flabellascoma
9. Ascospores not only as above……………Lophiostoma
Saprobic on dead herbaceous stems or culms of Poaceae
or submerged wood. Sexual morph: Ascomata solitary
to gregarious, immersed, with papilla erumpent through
host surface, reddish brown to black, subglobose, coriaceous, ostiolate. Ostiole variable in shape, central, with a
pore-like opening, plugged by gelatinous tissue, made of
hyaline, pseudoparenchymatous cells. Peridium composed
of a single stratum, with several layers of cells of textura
angularis, fusing with the host tissues in outer layer. Pseudoparaphyses numerous, cellular, hypha-like, hyaline, septate,
branched. Asci 8-spored, bitunicate, fissitunicate, cylindricclavate, with short, bulbous pedicel, with an ocular chamber. Ascospores uni- to bi-seriate, fusiform, with acute ends,
septate, constricted at the septa, pale yellowish to brown,
with hyaline appendages (Thambugala et al. 2015; Bao et al.
2019c). Asexual morph: Undetermined.
Type species: Biappendiculispora japonica Thambug.,
Wanas., Kaz. Tanaka & K.D. Hyde
Notes: The monotypic genus Biappendiculispora was introduced for B. japonica (Thambugala et al. 2015) which was
initially described as Lophiostoma caulium “var. f” (Tanaka
and Harada 2003b). Three strains of B. japonica collected in
Japan clustered in a separated clade with Lophiostoma (Thambugala et al. 2015; Bao et al. 2019c; this study, Fig. 74). Bao
et al. (2019c) found this species from freshwater in China.
List of freshwater Biappendiculispora species
Biappendiculispora Thambug., Kaz. Tanaka & K.D. Hyde,
Fungal Diversity 74: 214 (2015)
13
*Biappendiculispora japonica Thambug., Wanas., Kaz.
Tanaka & K.D. Hyde, Fungal Diversity 74: 214 (2015)
Fungal Diversity (2020) 105:319–575
Freshwater distribution: China (Bao et al. 2019c)
Crassiclypeus A. Hashim., K. Hiray. & Kaz. Tanaka, Stud.
Mycol. 90: 167 (2018)
Saprobic on submerged dead twigs of woody plant. Sexual morph: Ascomata scattered to gregarious, immersed,
subglobose, dark brown to black, ostiolate. Ostiolar neck
crest-like, elongate, laterally compressed, surrounded by a
well-developed clypeus. Peridium uniform, composed of
elongate, brown cells, surrounded by brown hyphae. Pseudoparaphyses numerous, trabeculate, filiform, hyaline, septate, branching and anastomosing. Asci 8-spored, bitunicate,
fissitunicate, clavate, pedicellate, apically rounded with an
ocular chamber. Ascospores fusiform, hyaline, septate,
smooth, with an internal chamber and a narrow sheath which
drawn out at both ends (Hashimoto et al. 2018). Asexual
morph: Coelomycetous. Conidiomata pycnidial, superficial
to immersed, scattered or clustered in small groups, globose
to subglobose. Peridium composed of subglobose to rectangular, brown cells. Conidiophores reduced to conidiogenous
cells. Conidiogenous cells phialidic, ampliform, hyaline,
smooth. Conidia subglobose with rounded ends, hyaline,
aseptate, smooth (Hashimoto et al. 2018).
Type species: Crassiclypeus aquaticus A. Hashim., K.
Hiray. & Kaz. Tanaka
Notes: Crassiclypeus was introduced to accommodate
the freshwater species C. aquaticus collected in Japan
(Hashimoto et al. 2018). The asexual morph has subglobose conidia formed in culture (Hashimoto et al. 2018).
Crassiclypeus aquaticus clusters basal to Flabellascoma
with moderate bootstrap support (Fig. 74). Crassiclypeus
aquaticus shares similar morphological characters with F.
fusiforme in having fusiform, hyaline ascospores with a narrow, bipolar sheath. However, an internal chamber was not
observed in the ascospores of F. fusiforme (see description
of F. fusiforme below).
List of freshwater Crassiclypeus species
*Crassiclypeus aquaticus A. Hashim., K. Hiray. & Kaz.
Tanaka, Stud. Mycol. 90: 167 (2018)
Freshwater distribution: Japan (Hashimoto et al. 2018).
Flabellascoma A. Hashim., K. Hiray. & Kaz. Tanaka, Stud.
Mycol. 90: 167 (2018)
Saprobic on plant materials and submerged wood.
Sexual morph: Ascomata scattered, immersed with neck
erumpent through host surface, subglobose to ellipsoidal.
Ostiolar neck elongate, laterally compressed. Peridium
composed of elongate, brown to dark brown cells. Pseudoparaphyses numerous, cellular, hypha-like, hyaline,
septate, branching and anastomosing. Asci 8-spored,
bitunicate, fissitunicate, cylindrical-clavate. Ascospores
fusiform, hyaline, septate, with a narrow, bipolar sheath
443
(Hashimoto et al. 2018). Asexual morph: Coelomycetous.
Conidiomata pycnidial, globose to subglobose. Peridium
composed of subglobose to rectangular, brown cells. Conidiophores absent. Conidiogenous cells holoblastic, cylindrical or ampliform, hyaline, smooth. Conidia subglobose
with rounded ends, hyaline, smooth, aseptate (Hashimoto
et al. 2018) (Fig. 68).
Type species: Flabellascoma minimum A. Hashim., K.
Hiray. & Kaz. Tanaka
Notes: Flabellascoma was introduced to accommodate
F. cycadicola A. Hashim. et al. and F. minimum. Flabellascoma is morphologically similar to Pseudolophiostoma
Thambug. et al. in ascomata with a well-developed, crestlike ostiolar neck (Thambugala et al. 2015; Hashimoto et al.
2018), but phylogeny separates them as distinct genera
(Hashimoto et al. 2018; this study, Fig. 74). Two freshwater
species F. aquaticum D.F. Bao et al. and F. fusiforme D.F.
Bao et al. are morphologically very similar, the former has
a long, cylindrical neck, but the latter has a short, crest-like
neck (Bao et al. 2019c). They also can be distinguished by
the size of asci (see key below).
List of freshwater Flabellascoma species
*Flabellascoma aquaticum D.F. Bao, Z.L. Luo, K.D. Hyde
& H.Y. Su, Mycosphere 10, 1085 (2019)
Freshwater distribution: China (Bao et al. 2019c)
*Flabellascoma fusiforme D.F. Bao, Z.L. Luo, K.D. Hyde
& H.Y. Su, Mycosphere 10, 1089 (2019); Fig. 69
Freshwater distribution: China (Bao et al. 2019c; this
study)
Saprobic on submerged wood. Sexual morph: Ascomata 160–180 μm high, 140–150 μm diam., black, scattered, immersed with neck erumpent through host surface,
subglobose. Ostiolar neck crest-like, elongate, compressed,
130–150 μm high, 75–85 μm wide, composed of several
layers of brown to black, thick-walled cells of textura angularis or globose cells. Peridium 15–25 μm thick, composed
of brown to dark brown, thin-walled cells of textura angularis or rectangular cells. Pseudoparaphyses 2 μm wide,
numerous, trabeculate, hyaline, sparsely septate, indistinctly
branched. Asci 8-spored, bitunicate, clavate, 66–87 × 8–11.5
μm ( x̄ = 74.8 × 10 μm, n = 10), apically rounded, with an
ocular chamber and a short pedicel. Ascospores 15–22 ×
4–6 μm (x̄ = 18 × 5 μm, n = 10), fusiform, biseriate, straight
or slightly curved, hyaline, 1-septate, deeply constricted at
the septum, slightly constricted at a quarter, the upper cell
slightly broader than the lower one, sharply or gradually
narrowed towards two ends, two prominent guttulate in each
cell, thin-walled, smooth, with a narrow, hyaline, lateral narrowly pad-like sheath, 3.5–7 μm long at both ends. Asexual
morph: Undetermined.
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Fungal Diversity (2020) 105:319–575
Fig. 69 Flabellascoma fusiforme (MFLU 18-1167). a
Appearance of necks erumpent
through host substrate. b Vertical section of ascoma. c Vertical
section of neck. d Structure
of peridium. e Pseudoparaphyses. f–i Bitunicate asci. j–l
Ascospores. m Germinated
ascospore. n, o Colony on PDA
(left-front, right-reverse). Scale
bars: b = 100 μm, c = 50 μm,
d–i = 20 μm, j–l = 10 μm, m
= 40 μm
Culture characteristics: On PDA, colony circular, reaching 5 mm in 15 days at 25 °C, white from above, pale brown
from below, surface rough, with dense mycelium, dry,
raised, edge entire.
13
Material examined: CHINA, Yunnan Province, Pingbian,
on submerged wood in a stream, 20 September 2017, W.
Dong, PB30A (MFLU 18-1167), living culture MFLUCC
18-1019.
Fungal Diversity (2020) 105:319–575
Notes: Our collection MFLUCC 18-1019 has identical
LSU and ITS sequence data to F. fusiforme, which indicates
that they are conspecific. Our collection has smaller ascomata (160–180 × 140–150 μm vs. 310–420 × 320–380 μm)
and a thinner peridium (15–25 μm vs. 25–50 μm) than the
holotype (Bao et al. 2019c). Phylogenetic analysis also supports that they are the same species (Fig. 74).
Key to freshwater Flabellascoma species
1. Asci 66–80 × 10–12 μm…………………F. fusiforme
1. Asci 48–72 × 8–9 μm……………………F. aquaticum
Lentistoma A. Hashim., K. Hiray. & Kaz. Tanaka, Stud.
Mycol. 90: 169 (2018)
Saprobic on woody plant or submerged wood. Sexual
morph: Ascomata scattered, immersed to erumpent, subglobose or conical, with ostiolar, elongate, laterally compressed neck, surrounded by a well-developed clypeus or
with ostiolate, short papilla. Peridium composed of globose, brown or black cells. Pseudoparaphyses numerous,
cellular, hypha-like, narrow, indistinctly septate, branching
and anastomosing. Asci 8-spored, bitunicate, fissitunicate,
cylindrical-clavate. Ascospores fusiform, hyaline, septate,
with or without a bipolar sheath (Hashimoto et al. 2018).
Asexual morph: Undetermined.
Type species: Lentistoma bipolare (K.D. Hyde) A.
Hashim., K. Hiray. & Kaz. Tanaka
Notes: Lentistoma was introduced based on Massarina
bipolaris K.D. Hyde (Hashimoto et al. 2018), which was
collected from woody plant or submerged wood. We introduce the second Lentistoma species which was collected
from submerged wood in Thailand.
List of freshwater Lentistoma species
*Lentistoma aquaticum W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557918; Facesoffungi number: FoF09260; Fig. 70
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-1718
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 270–320 μm high, 580–620 μm
wide, black, scattered, immersed to erumpent, conical, carbonaceous, with ostiolate, short papilla. Peridium 50–70
μm at the sides, 20–30 μm at the base, comprising carbonaceous, occluded dark cells, easily cracked, flattened at the
base. Pseudoparaphyses 3.5 μm wide, numerous, cellular,
hypha-like, hyaline, distantly septate. Asci (90–)130–160 ×
(12–)15–19(–21) μm ( x̄ = 145 × 17 μm, n = 10), 8-spored,
bitunicate, narrowly to broadly clavate, short pedicellate,
apically rounded with well-developed ocular chamber.
445
Ascospores 38–43 × 6.5–8.5 μm ( x̄ = 40 × 7.5 μm, n = 20),
biseriate or overlapping triseriate, slightly curved, hyaline,
1-septate, constricted at the septa, fusiform, guttulate, thinwalled, smooth, lacking a mucilaginous sheath. Asexual
morph: Undetermined.
Culture characteristics: On PDA, colony irregular, reaching 15 mm in 15 days at 25 °C, white from above, white to
pale yellow from below, surface rough, dry, raised, edge
entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat3106-1 (MFLU 18-1718, holotype), ex-type living culture MFLUCC 18-1275; ibid., hat3106-2 (HKAS 105032,
isotype), ex-type living culture KUMCC 19-0047.
Notes: Lentistoma aquaticum clusters with L. bipolare
with strong bootstrap support (Fig. 74). Lentistoma bipolare has crest-like, elongate, laterally compressed neck,
while L. aquaticum has ostiolate and short papilla. Lentistoma aquaticum has longer and wider asci ((90–)130–160
× (12–)15–19(–21) μm vs. (82–) 105–140 × 8–15 μm)
and longer ascospores (38–43 × 6.5–8.5 μm vs. 20–33 ×
5.5–9(–11) μm) than L. bipolare (Hashimoto et al. 2018).
Additionally, an internal chamber and a narrow sheath
drawn out at both ends are described for the ascospores of
L. bipolare, but not L. aquaticum (Hashimoto et al. 2018).
There are 3, 31 and 13 nucleotide differences in LSU, ITS
and TEF sequence data, respectively, between L. aquaticum
MFLUCC 18-1275 and L. bipolare CBS 115375 (ex-type).
Thus, L. aquaticum sp. nov. is introduced according to the
guideline of Jeewon and Hyde (2016).
*Lentistoma bipolare (K.D. Hyde) A. Hashim., K. Hiray. &
Kaz. Tanaka, Stud. Mycol. 90: 169 (2018)
Basionym: Massarina bipolaris K.D. Hyde, Nova Hedwigia 61(1-2): 131 (1995)
Synonymy: Lophiostoma bipolare (K.D. Hyde) E.C.Y.
Liew, Aptroot & K.D. Hyde, Mycologia 94(5): 812 (2002)
Freshwater distribution: Australia (Hyde 1995c; Hyde
and Aptroot 1998b), Brunei (Ho et al. 2001), China (Hyde
1995a; Tsui et al. 2000; Ho et al. 2001, 2002; Luo et al.
2004), Japan (Hashimoto et al. 2018), Malaysia (Hyde and
Aptroot 1998b), Seychelles (Hyde and Goh 1998b), South
Africa (Hyde and Goh 1998b), Thailand (Sivichai et al.
2000, 2002), USA (Raja et al. 2009b)
Key to feshwater Lentistoma species
1. Ascomata with ostiolar, elongate, laterally compressed neck, surrounded by a well-developed clypeus,
ascospores 20–33 × 5.5–9(–11) μm, with a narrow
sheath………………………………………L. bipolare
1. Ascomata with ostiolar, short papilla, ascospores 38–43
× 6.5–8.5 μm, without sheath……………L. aquaticum
13
446
Fig. 70 Lentistoma aquaticum (MFLU 18-1718, holotype). a,
b Appearance of ascomata on host surface. c Vertical section of
ascoma. d–h Bitunicate asci. i Pseudoparaphyses. j–n Ascospores.
13
Fungal Diversity (2020) 105:319–575
o Ascospore in Indian Ink. p Germinated ascospore. q, r Colony on
PDA (left-front, right-reverse). Scale bars: c = 100 μm, d–o = 20 μm,
p = 50 μm
Fungal Diversity (2020) 105:319–575
Lophiostoma Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
219 (1863)
Saprobic on herbaceous and woody substrates in terrestrial and aquatic habitats. Sexual morph: Ascomata scattered to gregarious, immersed to semi-immersed, papilla
erumpent through host surface, uniloculate, globose to
subglobose, glabrous, dark brown to black, coriaceous to
carbonaceous, ostiolate. Ostiole slit-like, with crest-like
apex, usually with a pore opening, plugged by gelatinous
tissue. Peridium wider at the apex and thinner at the base,
composed of a single stratum, comprising several layers of
lightly pigmented to dark brown, thin-walled cells, outer
layer fusing and indistinguishable from the host tissues.
Pseudoparaphyses numerous, cellular, hypha-like, septate,
branched, embedded in a gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, cylindro-clavate, short to long
pedicellate, rounded at the apex with an ocular chamber.
Ascospores uni- or partially bi-seriate, fusiform, straight
to slightly curved, hyaline to yellowish brown, 1- to multiseptate, sometimes with 3–5-eusepta, constricted at the
central septum (Thambugala et al. 2015). Asexual morph:
Coelomycetous. Conidiomata scattered or clustered, subglobose with a flattened base, covered with brownish, septate
hyphae. Neck central, cylindrical, composed of dark brown,
polygonal to subglobose cells. Pycnidial wall slightly thinner at the base, composed of subglobose to polygonal cells.
Conidiophores septate, branched, hyaline. Conidiogenous
cells phialidic. Conidia ellipsoidal to obovoid, aseptate, hyaline, smooth (Tanaka and Harada 2003b).
Type species: Lophiostoma macrostomum (Tode) Ces. &
De Not.
Notes: Lophiostoma is an old genus with over 400 epithets recorded (Index Fungorum 2020). Lophiostoma was
confirmed in Lophiostomataceae based on multigene phylogenetic analyses (Thambugala et al. 2015). Hashimoto et al.
(2018) revisited 11 species that were classified as Lophiostoma bipolare complex, and proposed five new genera. The
asexual morph was found in the culture of L. semiliberum
(Desm.) Ces. & De Not. and characterized by subglobose
conidiomata covered by hyphae, phialidic conidiogenous
cells and ellipsoid to obovoid conidia (Tanaka and Harada
2003b). Six species have been found from freshwater habitats, but none of them is confirmed by molecular evidence.
Of the six freshwater species, L. purpurascens is the only
species staining the substrate purple (Hyde and Aptroot
1998b). Staining the substrate purple was also observed in
other freshwater species, e.g. Amniculicola parva, Jahnula
purpurea, Purpureofaciens aquatica (Zhang et al. 2009b;
Fournier et al. 2015; this study). Fresh collections are
needed to verify the taxonomic placement of these freshwater species.
List of freshwater Lophiostoma species
447
Lophiostoma aquaticum (J. Webster) Aptroot & K.D. Hyde,
Fungal Diversity Res. Ser. 7: 106 (2002)
Basionym: Massarina aquatica J. Webster, Trans. Br.
mycol. Soc. 48(3): 451 (1965)
Freshwater distribution: Brunei (Ho et al. 2001), England
(Webster 1965), Malaysia (Ho et al. 2001), South Africa
(Hyde and Aptroot 1998b; Hyde et al. 1998)
Lophiostoma frondisubmersum (K.D. Hyde) E.C.Y. Liew,
Aptroot& K.D. Hyde, Mycologia 94(5): 812 (2002)
Basionym: Massarina fronsisubmersa K.D. Hyde, Mycol.
Res. 98(7): 724 (1994)
Freshwater distribution: New Guinea (Hyde 1994), USA
(Raja et al. 2009b)
Lophiostoma lunisporum (K.D. Hyde & Goh) Aptroot&
K.D. Hyde, Fungal Diversity Res. Ser. 7: 107 (2002)
Basionym: Massarina lunispora K.D. Hyde & Goh,
Mycol. Res. 102(6): 741 (1998)
Freshwater distribution: Australia (Hyde and Goh 1998a)
Lophiostoma maquilingense K.D. Hyde and Aptroot, Nova
Hedwigia 66(3–4): 499 (1998)
Freshwater distribution: Philippines (Hyde and Aptroot
1998b)
Lophiostoma proprietunicatum K.M. Tsui, K.D. Hyde &
Hodgkiss, Fungal Diversity Res. Ser. 7: 107 (2002)
Basionym: Massarina proprietunicata K.M. Tsui, K.D.
Hyde & Hodgkiss, Mycol. Res. 103(12): 1575 (1999)
Freshwater distribution: China (Tsui et al. 1999)
Lophiostoma purpurascens (K.D. Hyde & Aptroot) Aptroot
& K.D. Hyde, Fungal Diversity Res. Ser. 7: 108 108(2002)
Basionym: Massarina purpurascens K.D. Hyde & Aptroot, Nova Hedwigia 66(3–4): 496 (1998)
Freshwater distribution: Australia (Hyde and Aptroot
1998b), China (Tsui et al. 2000), New Guinea (Hyde and
Aptroot 1998b)
Key to freshwater Lophiostoma species
1.
1.
2.
2.
3.
3.
4.
4.
5.
Often stain the substrate purple………L. purpurascens
Do not stain substrate purple……………………………2
Ascospores hyaline, 1-septate…………………………3
Ascospores initially hyaline, 1-septate, becoming pale
brown, 3-septate with age………………L. aquaticum
Ascospores with appendages…………………………4
Ascospores without appendages………L. lunisporum
Ascomata immersed beneath a darkened, slightly raised
pseudostromata……… ………………L. maquilingense
Ascomata not as above………………………………5
Ascospore appendages > 18 μm…L.proprietunicatum
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Fungal Diversity (2020) 105:319–575
Fig. 71 Neotrematosphaeria biappendiculata (Material examined:
JAPAN, Serisawa-park (pond), Matsukitai, Hirosaki, Aomori, on
submerged twigs, 7 December 2002, KT & NA 1124, HHUF 28026,
holotype; Mohei-pond, Aoki, Hirosaki, Aomori, on submerged twigs,
27 September 2003, KT & NA 1489a, HHUF 28306, paratype). a–c
from KT 1489a, paratype. a Ascomata on the host surface. b Section
of ascoma. c Structure of peridium. d–g from KT 1124, holotype. d
Pseudoparaphyses. e Ascus. f, g Ascospores (f in black-blue ink. Note
the pad-like appendages at both ends). Scale bars: a = 500 μm, b =
250 μm, c–g = 10 μm
5. Ascospore appendages < 18 μm…L. frondisubmersum
Type species: Neotrematosphaeria biappendiculata (Kaz.
Tanaka, Y. Harada & M.E. Barr) Thambug., Kaz. Tanaka
& K.D. Hyde
Notes: The monotypic genus Neotrematosphaeria was
introduced to accommodate Trematosphaeria biappendiculata Kaz. Tanaka et al. which was isolated from submerged
wood in freshwater in Northern Japan (Tanaka et al. 2005a;
Thambugala et al. 2015). The pale yellowish to olivaceous
brown, 5(–7)-septate ascospores with globose to conical terminal appendages can distinguish Neotrematosphaeria from
other genera in Lophiostomataceae.
Neotrematosphaeria Thambugala, Kaz. Tanaka & K.D.
Hyde, Fungal Diversity 74: 223 (2015)
Saprobic on submerged, dead twigs. Sexual morph:
Ascomata solitary, semi-immersed to erumpent, uniloculate,
subglobose, glabrous, black, coriaceous to carbonaceous,
with ostiolate papilla, with a crest-like apex and a pore-like
opening. Peridium broad at sides, poorly developed at the
base, composed of a single stratum, with several layers of
dark brown to black cells of textura prismatica. Pseudoparaphyses numerous, cellular, filiform, indistinctly septate,
embedded in a gelatinous matrix. Asci 8-spored, bitunicate,
fissitunicate, cylindric-clavate, with a long pedicel, rounded
at the apex, with an ocular chamber. Ascospores biseriate to
uniseriate near the base, fusiform, mostly curved, pale yellowish to olivaceous brown, but pale at the terminal cells,
septate, with bipolar appendages (Tanaka et al. 2005a;
Thambugala et al. 2015). Asexual morph: Coelomycetes,
pleurophomopsis-like. Conidiomata erumpent to superficial,
uniloculate, globose, with thick, brown, sparse hyphae at
sides, with ostiolate papilla. Peridium composed of polygonal to globose cells. Conidiophores cylindrical, branched,
0–1-septate. Conidiogenous cells phialidic. Conidia
extremely small-sized, subglobose to ellipsoid, straight,
hyaline, aseptate, smooth (Tanaka et al. 2005a).
13
List of freshwater Neotrematosphaeria species
*Neotrematosphaeria biappendiculata (Kaz.Tanaka, Y.
Harada & M.E. Barr) Thambugala, Kaz. Tanaka & K.D.
Hyde; Fig. 71
Basionym: Trematosphaeria biappendiculata Kaz.
Tanaka, Y. Harada & M.E. Barr, Fungal Diversity 19: 149
(2005)
Freshwater distribution: Japan (Tanaka et al. 2005a)
Neovaginatispora A. Hashim., K. Hiray. & Kaz. Tanaka,
Stud. Mycol. 90: 188 (2018)
Saprobic on dead herbaceous twigs and submerged
wood. Sexual morph: Ascomata solitary, semi-immersed
to erumpent, black, subglobose, coriaceous, ostiolate. Ostiole rounded or slit-like, central, with a pore-like opening.
Peridium uneven in width, thinner at the base, two-layered,
Fungal Diversity (2020) 105:319–575
outer layer fusing with the host cells, inner layer comprising
hyaline cells of textura angularis. Pseudoparaphyses numerous, cellular, hypha-like, septate. Asci 8-spored, bitunicate,
fissitunicate, cylindric-clavate, with a short, bulbous pedicel,
with an indistinct ocular chamber. Ascospores uni- to biseriate, fusiform with acute ends, septate, constricted at the
septum, hyaline, smooth-walled, with globose appendages at
both ends (Hashimoto et al. 2018; Bao et al. 2019c). Asexual
morph: Undetermined.
Type species: Neovaginatispora fuckelii (Sacc.) A.
Hashim., K. Hiray. & Kaz. Tanaka
Notes: Neovaginatispora was proposed for Vaginatispora fuckelii (Sacc.) Thambug. et al. due to its thinner (up
to 25 μm thick) and uniformly thick peridium (Hashimoto
et al. 2018). However, the freshwater collection of Neovaginatispora fuckelii found by Bao et al. (2019c) has thicker
peridium (up to 40 μm thick) which is thinner at the base.
The specimen HHUF 27325 examined by Thambugala et al.
(2015) also showed the uneven peridium width. Phylogenetic analyses based on four strains of N. fuckelii guaranteed
Neovaginatispora as a distinct genus (Hashimoto et al. 2018;
Bao et al. 2019c; this study, Fig. 74).
List of freshwater Neovaginatispora species
*Neovaginatispora fuckelii (Sacc.) A. Hashim., K. Hiray.
& Kaz. Tanaka, Stud. Mycol. 90: 188 (2018)
Basionym: Lophiostoma fuckelii Sacc., Michelia 1(no. 3):
336 (1878)
Synonymy: see Index Fungorum (2020)
Freshwater distribution: China (Bao et al. 2019c)
Pseudocapulatispora Mapook & K.D. Hyde, Fungal Diversity 101: 47 (2020)
Saprobic on submerged wood or dead stems. Sexual
morph: Ascomata brown to black, solitary or scattered,
immersed with papilla erumpent through host surface,
ellipsoidal or ovoid, coriaceous, ostiolate. Ostiolar papilla
short, blackened, compressed, crest-like. Peridium composed of several layers of brown to dark brown, thin-walled
cells. Pseudoparaphyses numerous, trabeculate, filiform,
hyaline, sparsely septate, branched. Asci 8-spored, bitunicate, mostly broadly clavate, rarely cylindric-clavate, apically narrowly rounded, with an ocular chamber and a short
pedicel. Ascospores biseriate, broadly fusiform, hyaline,
septate, mostly rounded or rarely sharply narrowed at two
ends, guttulate, thin-walled, smooth, with a narrow, hyaline, elongate sheath (Mapook et al. 2020). Asexual morph:
Undetermined.
Type species: Pseudocapulatispora longiappendiculata
Mapook & K.D. Hyde
Notes: Pseudocapulatispora longiappendiculata was
isolated from a dead stem of Chromolaena odorata in Thailand (Mapook et al. 2020). Pseudocapulatispora is distinct
449
in Lophiostomataceae by its sheath drawn out to form polar
appendages with tips of the sheath capped (Mapook et al.
2020). Capulatispora Thambug. & K.D. Hyde has a similar
sheath but it mostly has cylindric-clavate asci and fusiform
ascospores with narrowed and acute ends (Thambugala
et al. 2015). Phylogenetic analysis supported them as distinct genera (Thambugala et al. 2015; Mapook et al. 2020).
We collected a new specimen of P. longiappendiculata from
freshwater habitats.
List of freshwater Pseudocapulatispora species
*Pseudocapulatispora longiappendiculata Mapook & K.D.
Hyde, Fungal Diversity 101: 48 (2020); Fig. 72
Freshwater distribution: China (this study)
Saprobic on submerged wood. Sexual morph: Ascomata 270–310 μm high, 310–330 μm diam., black, scattered, immersed with papilla erumpent through host surface,
ellipsoidal, ostiolate. Ostiolar papilla short, blackened, compressed, 110–130 μm high, 160–180 μm wide, composed of
dark brown to black, thick-walled cells of textura angularis
or subglobose cells, 20–25 μm thick at the base, 70–80 μm
thick at the top. Peridium 15–30 μm thick, composed of
several layers of brown to dark brown, thin-walled cells of
textura angularis or subglobose cells. Pseudoparaphyses 2
μm wide, numerous, trabeculate, filiform, hyaline, sparsely
septate, branched. Asci 8-spored, bitunicate, mostly broadly
clavate, rarely cylindric-clavate, slightly to obviously
curved, (90–)105–130 × 12–17 μm ( x̄ = 115 × 14.5 μm, n
= 10), apically narrowly rounded, with an ocular chamber
and a short pedicel. Ascospores (20.5–)25.5–30 × 6–9 μm
( x̄ = 27 × 7 μm, n = 10), biseriate, ellipsoidal, mostly with
rounded or rarely acute ends, straight or slightly curved, hyaline, 1-septate, slightly constricted at the to septum, slightly
constricted at a quarter, guttulate, thin-walled, smooth, with
a narrow, hyaline, elongate sheath with globose ending, 30
μm long. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 15 days at 25 °C, white to grey from above, pale
yellow to pale brown from below, surface rough, with dense
mycelium, dry, raised, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H39A-1 (MFLU 18-1188), living culture MFLUCC
18-1027; ibid., H39A-2 (HKAS 101722), living culture
KUMCC 18-0072.
Notes: Phylogenetic analysis shows our collection
MFLUCC 18-1027 is P. longiappendiculata (Fig. 74).
MFLUCC 18-1027 has shorter appendages (30 μm vs. 40
μm) than the holotype (Mapook et al. 2020). This is a new
habitat and new geographical record for P. longiappendiculata from freshwater in China.
13
450
Fungal Diversity (2020) 105:319–575
Fig. 72 Pseudocapulatispora
longiappendiculata (MFLU
18-1188, new habitat and
geographical record). a,
b Appearance of ostiolar
papilla erumpent through host
substrate. c Vertical section of ascoma. d Structure
of peridium. e–h Bitunicate
asci. i Pseudoparaphyses. j–l
Ascospores. m Germinated
ascospore. n, o Colony on PDA
(up-front, down-reverse). Scale
bars: c = 100 μm, d–l = 20 μm,
m = 50 μm
Sigarispora Thambug. & K.D. Hyde, Fungal Diversity 74:
238 (2015)
Saprobic on decaying culms of grasses (Poaceae), dead
herbaceous stems or submerged wood. Sexual morph:
Ascomata solitary, scattered to gregarious, semi-immersed
to immersed, with papilla erumpent through host surface,
13
globose to subglobose, black, ostiolate. Ostiole crest- or slitlike, central, rounded, with a pore-like opening and plugged
by gelatinous tissue, comprising pseudoparenchymatous
cells. Peridium composed of light to dark brown, small,
thin-walled cells of textura angularis, fusing with the host
tissues in outer layer. Pseudoparaphyses numerous, cellular,
Fungal Diversity (2020) 105:319–575
long, hyaline, septate, branched. Asci 8-spored, bitunicate,
fissitunicate, cylindric-clavate, pedicellate, with an ocular
chamber. Ascospores uni- to bi-seriate, yellowish brown
to dark brown, cigar-shaped or ellipsoidal-fusiform, transversely septate or muriform, smooth-walled, with or without
a mucilaginous sheath, sometimes with terminal appendages
(Thambugala et al. 2015). Asexual morph: Undetermined.
Type species: Sigarispora ravennica (Tibpromma, Camporesi & K.D. Hyde) Thambug. & K.D. Hyde
Notes: Sigarispora was introduced for some lophiostomalike species (Thambugala et al. 2015). Sigarispora is phylogenetically separated from Lophiostoma (Thambugala et al.
(2015); this study, Fig. 74) and characterized by yellowish
brown to dark brown, cigar-shaped or ellipsoidal-fusiform
ascospores which are unlike the hyaline to yellowish brown,
fusiform ascospores of Lophiostoma. Bao et al. (2019c)
reported a freshwater species S. clavata D.F. Bao et al. from
China and characterized by dark brown to yellowish brown,
ellipsoidal to clavate ascospores.
List of freshwater Sigarispora species
*Sigarispora clavata D.F. Bao, Z.L. Luo, K.D. Hyde & H.Y.
Su, Mycosphere 10: 1090 (2019)
Freshwater distribution: China (Bao et al. 2019c)
Vaginatispora K.D. Hyde, Nova Hedwigia 61(1–2): 234
(1995)
Saprobic on submerged wood, intertidal wood, dead
twigs, endocarp or fallen fruit pericarp. Sexual morph:
Ascomata solitary or scattered, immersed to erumpent, uniloculate, subglobose, glabrous, dark brown to black. Ostiolar
neck crest-like, elongate, laterally compressed, composed of
globose to elongate, brown to black cells, with a pore-like
opening and hyaline periphyses. Peridium unequal in thickness, two-layered, outer layer comprising somewhat flattened
cells, fusing and indistinguishable from the host tissues,
inner layer comprising lightly pigmented to hyaline cells.
Pseudoparaphyses numerous, cellular, hypha-like, hyaline,
septate, anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical to clavate, with a short or long
pedicel, apically round with an ocular chamber. Ascospores
uni- to bi-seriate, narrowly ellipsoidal or fusiform, straight
to slightly curved, hyaline, 1-septate, occasionally producing pseudosepta, septum mostly median, upper cell slightly
broader than lower cell, smooth, thin-walled, with bipolar
appendages or entire sheath (Hyde 1995b). Asexual morph:
Undetermined.
Type species: Vaginatispora aquatica K.D. Hyde
Notes: Vaginatispora aquatica was collected from North
Queensland, Australia (Hyde 1995b). Zhang et al. (2014a)
transferred V. aquatica to Lophiostoma because it formed a
basal clade to Lophiostoma species with high bootstrap support and morphologically resembles L. macrostomum (type
451
species), in having immersed to erumpent ascomata with a
slit-like ostiole, an unequally thick peridium, clavate asci,
and hyaline ascospores (Hyde 1995b; Zhang et al. 2014a).
With more sequences added in Lophiostomataceae, Vaginatispora (typified by V. aquatica) was accepted as a distinct
genus in Lophiostomataceae (Thambugala et al. 2015). The
freshwater collections of V. armatispora (K.D. Hyde et al.)
Wanas. et al. were found from Thailand (Bao et al. 2019c;
Hyde et al. 2019).
List of freshwater Vaginatispora species
*Vaginatispora aquatica K.D. Hyde, Nova Hedwigia
61(1–2): 235 (1995); Fig. 73
Synonymy: Lophiostoma vaginatispora H. Zhang & K.D.
Hyde, Phytotaxa 176(1): 177 (2014)
Freshwater distribution: Australia (Hyde 1995b), China
(Tsui et al. 2000), Thailand (Zhang et al. 2014a)
*Vaginatispora armatispora (K.D. Hyde, Vrijmoed, Chinnaraj & E.B.G. Jones) Wanas., E.B.G. Jones & K.D. Hyde,
Stud. Fung. 1(1): 62 (2016)
Basionym: Massarina armatispora K.D. Hyde, Vrijmoed,
Chinnaraj & E.B.G. Jones, Bot. Mar. 35(4): 325 (1992)
Synonymy: Lophiostoma armatisporum (K.D. Hyde, Vrijmoed, Chinnaraj & E.B.G. Jones) E.C.Y. Liew, Aptroot &
K.D. Hyde, Mycologia 94(5): 812 (2002)
Freshwater distribution: Thailand (Hu et al. 2010b; Bao
et al. 2019c; Hyde et al. 2019)
Key to freshwater Vaginatispora species
1. Ascospores 33–45 × 11–12.5 μm, with a mucilaginous
collar around its equator, surrounded by a spreading
papilionaceous sheath………………………V. aquatica
1. Ascospores 22–34 × 5.5–9 μm, surrounded by a narrow
mucilaginous sheath, with distinct hyaline appendages at
both ends………………………………V. armatispora
Lophiotremataceae K. Hiray. & Kaz. Tanaka, Mycoscience
52(6): 405 (2011)
Lophiotrema Sacc., Michelia 1(no. 3): 338 (1878)
Saprobic on woody and herbaceous substrates or submerged wood. Sexual morph: Ascomata scattered to
crowded, mostly immersed, rarely erumpent, uniloculate,
globose, subglobose to ovoid, glabrous, dark brown to black,
opening compressed, with a slit-like ostiole. Peridium composed of pale brown, small, thin-walled cells of textura
angularis to globosa. Pseudoparaphyses numerous, cellular, hyaline, septate, occasionally anastomosing or branching. Asci 8-spored, bitunicate, fissitunicate, cylindrical,
sessile to minutely pedicellate, rounded at the apex, with
13
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Fungal Diversity (2020) 105:319–575
Fig. 73 Vaginatispora aquatica (Material examined: THAILAND,
Chiang Mai Province, Doi Inthanon, 16 November 2010, H. Zhang,
MFLU 11-1115). a, b Appearance of ascomata on wood. c Section
of ascoma. d Papilla of ascoma. e Ascus. f, g Ascospores with wide
papilionaceous sheath. (g in Indian Ink). Scale bars: c, d = 100 μm, e
= 30 μm, f = 20 μm, g = 40 μm
an ocular chamber. Ascospores uni- to bi-seriate, fusiform
to cylindrical, straight to slightly curved, hyaline to brown,
septate, symmetrical, smooth, thin-walled (Hirayama and
Tanaka 2011; Hyde et al. 2013). Asexual morph: probably
pleurophomopsis-like (Leuchtmann 1985).
Type species: Lophiotrema nucula (Fr.) Sacc.
Notes: Lophiotrema is distinguished from Lophiostoma
by subglobose to globose ascomata, peridium composed
thin-walled cells of textura angularis to globosa, cylindrical asci with a short pedicel, and hyaline to brown, septate
ascospores (Hirayama and Tanaka 2011; Hyde et al. 2013).
Hirayama and Tanaka (2011) mentioned that the ascal
shape, including length of the ascal pedicel, was a reliable
taxonomic indicator to separate these two genera (cylindrical, up to 15 µm long in Lophiotrema vs. clavate, mostly
(10–)15–30 µm long in Lophiostoma). Based on multigene
phylogenetic analysis, Lophiotrema was placed in Lophiotremataceae, which was distant from Lophiostomataceae
(Hirayama and Tanaka 2011). Pleurophomopsis Petr. was
linked to the asexual morph of Lophiotrema, which formed
in culture of L. nucula (Leuchtmann 1985), but Hyde et al.
(2013) considered that it was probably a spermatial state.
The freshwater species Lophiotrema rubi (Fuckel) Y.
Zhang ter et al. is characterized by hemispherical to pyriform
ascomata with black clypeus formed around the neck, broadly
cylindrical asci with a hoof-shaped pedicel, and broadly fusiform, hyaline ascospores with bipolar appendages (Fallah and
Shearer 2001). Lophiotrema rubi was transferred from Massarina rubi (CBS 691.95) based on multigene phylogenetic
analysis (Zhang et al. 2009c). Lophiotrema rubi is likely an
immigrant to freshwater because most reports are from terrestrial habitats (Fallah and Shearer 2001).
13
List of freshwater Lophiotrema species
Lophiotrema rubi (Fuckel) Y. Zhang ter, C.L. Schoch &
K.D. Hyde, Stud. Mycol. 64: 97 (2009)
Basionym: Massaria rubi Fuckel, Jb. nassau. Ver. Naturk.
25-26: 303 (1871)
Synonymy: Didymellina rhaphithamni Keissl., Nat. Hist.
Juan Fernandez Easter Isl. 2: 480 (1927)
Lophiostoma rubi (Fuckel) E.C.Y. Liew, Aptroot & K.D.
Hyde, Mycologia 94(5): 812 (2002)
Lophiotrema emergens P. Karst., Hedwigia 22: 42 (1883)
Massarina emergens (P. Karst.) L. Holm, Les Pleosporaceae: 149 (1957)
Massarina rubi (Fuckel) Sacc., Syll. fung. (Abellini) 2:
155 (1883)
Mycosphaerella rhaphithamni (Keissl.) Petr., Annls
mycol. 38(2/4): 221 (1940)
Freshwater distribution: USA (Fallah and Shearer 2001)
Massarinaceae Munk, Friesia 5(3-5): 305 (1956)
Key to freshwater genera of Massarinaceae
1. Ascospores basically symmetric, unevenly coloured,
brown in central two cells and paler in end cells at maturity………………………………………Byssothecium
Fungal Diversity (2020) 105:319–575
Fig. 74 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS, TEF and RPB2
sequence data for species of
Lophiostomataceae. Bootstrap
values for maximum likelihood
equal to or greater than 75% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree is
rooted to Teichospora rubriostiolata TR 7 and T. trabicola
C134 (Teichosporaceae)
453
Neotrematosphaeria biappendiculata KT 975 F
Neotrematosphaeria biappendiculata KT 1124 F
Coelodictyosporium pseudodictyosporium MFLUCC 13-0451
Coelodictyosporium muriforme MFLUCC 13-0351
Lophiostoma semiliberum KT 828
Lophiostoma macrostomum KT 635
Lophiostoma crenatum CBS 629.86
Lophiostoma caulium CBS 623.86
Lophiohelichrysum helichrysi MFLUCC 15-0701
76/-Platystomum crataegi MFLUCC 14-0925
Platystomum compressum MFLUCC 13-0343
81/0.99
Platystomum salicicola MFLUCC 15-0632
Platystomum actinidiae KT 521
89/-Sigarispora clavata MFLUCC 18-1316 F
Sigarispora caudata KT 530
Sigarispora arundinis KT 651
97/1.00
Sigarispora ravennica MFLUCC 14-0005
Sigarispora ononidis MFLU 15-2667
80/1.00
98/0.99
Lophiopoacea winteri KT 740
91/1.00
Lophiopoacea winteri KT 764
Lophiopoacea paramacrostoma MFLUCC 11-0463
91/0.99
Guttulispora crataegi MFLUCC 14-0993
Guttulispora crataegi MFLUCC 13-0442
Capulatispora sagittiformis KT 1934
90/0.99
Biappendiculispora japonica KT 686-1
99/1.00
Biappendiculispora japonica KT 573
Biappendiculispora japonica MFLUCC 17-2450 F
Lophiostoma macrostomoides CBS 123097
Lophiostoma heterosporum CBS 644.86
100/1.00
Pseudoplatystomum scabridisporum BCC 22835
Pseudoplatystomum scabridisporum BCC 22836
Pseudolophiostoma obtusisporum CBS 143655
100/1.00
100/1.00
Pseudolophiostoma obtusisporum CBS 143656
100/1.00
Pseudolophiostoma cornisporum CBS 143654
100/1.00
82/0.99
Pseudolophiostoma tropicum CBS 143660
92/0.99
Pseudolophiostoma
tropicum CBS 143659
100/1.00
Pseudolophiostoma vitigenum HH 26931
91/1.00
Pseudolophiostoma vitigenum HH 26930
Alpestrisphaeria terricola SC 12
Pseudocapulatispora
longiappendiculata MFLUCC 17-1457
100/0.99
--/0.99
Pseudocapulatispora longiappendiculata MFLUCC 17-1452
100/1.00
Pseudocapulatispora longiappendiculata MFLUCC 18-1027 F
Lophiostoma quadrinucleatum GKM 1233
100/1.00
1. Ascospores asymmetric, upper cell longer and larger
than lower cell, evenly coloured, medium to dark brown
at maturity…………………………Helminthosporium
Byssothecium Fuckel, Bot. Ztg. 19: 251 (1861)
Saprobic on submerged wood in freshwater or decaying wood in terrestrial habitats. Sexual morph: Ascomata
Neotrematosphaeria
Coelodictyosporium
Lophiostoma
Lophiohelichrysum
Platystomum
Sigarispora
Lophiopoacea
Guttulispora
Guttulispora
Biappendiculispora
Pseudoplatystomum
Pseudolophiostoma
Alpestrisphaeria
Pseudocapulatispora
gregarious, erumpent to superficial, uniloculate, subglobose,
ovoid to obpyriform, black, coriaceous, with a central ostiolate papilla. Peridium composed of brown, rectangle, pseudoparenchymatous cells, somewhat thinner at base. Pseudoparaphyses dense, cellular, hypha-like, hyaline, septate,
embedded in a gelatinous matrix, anastomosing between
and above the asci. Asci 8-spored, bitunicate, fissitunicate,
cylindrical to clavate, pedicellate. Ascospores biseriate at
13
454
Fungal Diversity (2020) 105:319–575
Fig. 74 (continued)
100/1.00
--/1.00
100/1.00
Lentistoma bipolare CBS 143648
Lentistoma bipolare CBS 143647 F
Lentistoma bipolare CBS 110448 F
Lentistoma bipolare CBS 143649 F
Lentistoma
100/1.00 Lentistoma bipolare CBS 115375
Lentistoma bipolare CBS 143651
99/1.00
Lentistoma bipolare CBS 143652
100/1.00
Lentistoma bipolare CBS 115370
Lentistoma aquaticum MFLUCC 18-1275 F
Vaginatispora armatispora HKTLCC 1562
99/-100/1.00
Vaginatispora armatispora MFLUCC 18-0213 F
100/1.00
100/1.00
Vaginatispora armatispora MFLUCC 18-0247 F
100/1.00
Vaginatispora scabrispora CBS 143663
Vaginatispora
100/1.00
Vaginatispora amygdali CBS 143662
100/1.00
Vaginatispora aquatica MFLUCC 11-0083 F
Vaginatispora appendiculata MFLUCC 16-0314
88/-Neovaginatispora fuckelii CBS 101952
89/-Neovaginatispora fuckelii KH 161
100/1.00
Neovaginatispora
Neovaginatispora fuckelii KT 634
Neovaginatispora fuckelii MFLUCC 17-1334 F
100/1.00 Flabellascoma minimum CBS 143646
98/1.00
Flabellascoma minimum CBS 143645
79/0.95
Flabellascoma cycadicola CBS 143644
Flabellascoma
100/1.00
Flabellascoma aquaticum KUMCC 15-0258 F
Flabellascoma fusiforme MFLUCC 18-1019 F
83/1.00 100/1.00
Flabellascoma fusiforme MFLUCC 18-1584 F
--/1.00 100/1.00 Crassiclypeus aquaticus CBS 143643 F
Crassiclypeus
--/0.99
Crassiclypeus aquaticus CBS 143642 F
Leptoparies
Leptoparies palmarum CBS 143653
--/1.00
Parapaucispora
Parapaucispora pseudoarmatispora KT 2237
Lophiostoma alpigenum GKM 1091b
100/1.00 Paucispora quadrispora KT 843
Paucispora quadrispora KH 448
Paucispora
100/1.00
Paucispora versicolor KH 110
Dimorphiopsis brachystegiae CPC 22679
Pseudopaucispora
Pseudopaucispora brunneospora CBS 143661
Teichospora trabicola C 134
Outgroup
Teichospora rubriostiolata TR 7
0.05
the apex, becoming uniseriate near the base, ellipsoidal to
fusiform, straight to slightly curved, hyaline to brown, usually paler at end cells, septate, basically symmetric, smooth
to minutely verrucose, thin-walled (Crane et al. 1992; Pem
et al. 2019b). Asexual morph: chaetophoma-like (Boise
1983).
Type species: Byssothecium circinans Fuckel
Notes: Byssothecium is characterized by ellipsoidal to
fusiform, hyaline to brown ascospores usually with bipolar paler cells (Crane et al. 1992; Pem et al. 2019b). The
13
asexual morph of B. circinans was produced in culture and
characterized by minute conidiomata, phialidic conidiogenous cells and hyaline, aseptate conidia, which are similar
to Chaetophoma (Boise 1983).
Savoryella E.B.G. Jones & R.A. Eaton shares similar
morphological characters with Byssothecium in having
cylindrical asci, ellipsoidal, 3-septate ascospores with brown
central cells and paler end cells (Jones and Eaton 1969;
Dayarathne et al. 2019). The ascomata have a long neck
in Savoryella, but it is lacking in Byssothecium. Phylogeny
Fungal Diversity (2020) 105:319–575
separates them in different classes: Byssothecium in Dothideomycetes (Lumbsch and Lindemuth 2001), and Savoryella in Sordariomycetes (Dayarathne et al. 2019) (Fig. 74).
One putative strain of Byssothecium circinans (CBS
675.92) clustered in Massarinaceae in Schoch et al. (2009).
Zhang et al. (2009d) found that B. circinans clustered basal
to Melanommataceae based on combined LSU and SSU
dataset, while clustered in Massarinaceae based on RPB2
dataset. Because B. circinans considerably differs from Massarina in morphology, such as black, erumpent ascomata,
ellipsoidal to fusiform ascospores with pigmented central
cells and hyaline end cells (Crane et al. 1992), and had an
unstable relationships in the phylognetic tree (Zhang et al.
2009d). Byssothecium circinans was not assigned to Melanommataceae and treated as an unresolved species (Zhang
et al. 2009c, d). Based on combined LSU, SSU and RPB2
dataset, B. circinans formed a stable clade in Melanommataceae (Zhang et al. 2012b; Hyde et al. 2013; Tanaka et al.
2015; Voglmayr and Jaklitsch 2017; Pem et al. 2019b; this
study, Fig. 2).
The morphology of strain Byssothecium circinans (CBS
675.92) is unavailable, a specimen G-K 18367 of B. circinans was examined by Pem et al. (2019b) and also examined in this study (Fig. 75). Two Byssothecium species were
reported from freshwater habitats (see list below), but not
confirmed by molecular data.
List of freshwater Byssothecium species
*Byssothecium circinans Fuckel, Bot. Ztg. 19: 251 (1861);
Fig. 75
Synonymy: Possible synonymy see Index Fungorum
(2020)
Freshwater distribution: Hungary (Révay and Gönczöl
1990; Gönczöl and Révay 1993)
Byssothecium flumineum J.L. Crane, Shearer & Huhndorf,
Mycologia 84(2): 236 (1992)
Freshwater distribution: USA (Crane et al. 1992)
Key to freshwater Byssothecium species
1. Ascospores 7.3−9.2 µm wide……………B. flumineum
1. Ascospores 10−12 µm wide………………B. circinans
Helminthosporium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1-2): 10 (1809)
Synonym: Exosporium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 9 (1809)
Saprobic on submerged wood, various dead plant material
or rarely parasitic on plants or fungicolous. Sexual morph:
Pseudostromata dark brown or reddish brown, pseudoparenchymatous, of thick-walled dark brown cells; margin mostly
455
composed of dark brown, verrucose hyphae. Ascomata
immersed in pseudostromata, solitary or in small groups,
large, often strongly depressed, or globose, dark brown to
black, with a central, inconspicuous ostiole. Peridium pseudoparenchymatous. Pseudoparaphyses numerous, cellular,
septate, branched, anastomosing, narrow, usually embedded in a gel matrix. Asci 8-spored, bitunicate, fissitunicate,
clavate or fusoid, short pedicellate. Ascospores irregularly
biseriate, fusoid, ellipsoidal, obovoid, occasionally oblong,
asymmetric, with one primary, eccentric, deeply constricted
euseptum, upper cell longer and larger than lower cell, often
with several transverse or oblique distosepta, rarely with a
longitudinal distoseptum in the larger cell, slightly or not
constricted at the secondary distosepta, hyaline when young,
becoming medium to dark brown when mature, with subacute to rounded end cells, smooth or verruculose, sometimes
with longitudinal striae, surrounded by a thick gelatinous
sheath (Voglmayr and Jaklitsch 2017). Asexual morph:
Colonies on natural substrate conspicuous, effuse to punctiform, hairy, brown to black. Mycelium mostly immersed,
composed of branched, septate hyphae. Conidiophores
macronematous, mononematous, arising solitarily or in fascicles, erect, mostly unbranched, straight or flexuous, cylindrical or subulate, with a well-defined small pore at the apex,
and often with several lateral pores beneath the upper septa,
ceasing growth with the formation of a terminal conidium,
usually not proliferating, pale to dark brown, smooth or
occasionally verruculose. Conidiogenous cells enteroblastic,
polytretic, integrated, determinate, terminal and intercalary,
subcylindrical. Conidia mostly solitary, acropleurogenous,
obclavate, obpyriform to lageniform, mostly rostrate, straight
or curved, pale brown to brown, distoseptate, smooth, with a
paler apical cell and truncate base, often with a prominent,
dark brown or black scar at the base (Ellis 1971; Zhu et al.
2016; Voglmayr and Jaklitsch 2017).
Type species: Helminthosporium velutinum Link [as
‘Helmisporium’], Mag. Gesell. naturf. Freunde, Berlin
3(1−2): 10 (1809)
Notes: Helminthosporium is an old genus which was
introduced in 1809 and currently comprises over 750 epithets
in Index Fungorum (2020). Helminthosporium is morphologically similar to Corynespora Güssow and Exosporium
Link as all of them have distoseptate conidia with tapering apex and truncate base, growing through a wide pore
at the apex of the conidiophores (Ellis 1971; Voglmayr and
Jaklitsch 2017). Exosporium has been synonymized with
Helminthosporium based on their very similar morphological characters (Fries 1832) and this was accepted by Voglmayr and Jaklitsch (2017) with molecular data supported.
Corynespora differs from Helminthosporium as conidiophores have successive proliferations (up to four), while
they are not described in Helminthosporium (Voglmayr and
Jaklitsch 2017). Phylogenetically, Corynespora has been
13
456
Fig. 75 Byssothecium circinans (Material examined: GERMANY,
Sabine, M. Huhndorf, 10 December 1993, G-K 18367). a, b Ascomata on the host surface. c Section of ascoma. d Peridium. e Pseudo-
13
Fungal Diversity (2020) 105:319–575
paraphyses. f−i Bitunicate asci. j−l Ascospores. Scale bars: b = 200
µm, c = 100 µm, d = 30 µm, e = 5 µm, f−h = 40 µm, i−l = 15 µm
Fungal Diversity (2020) 105:319–575
placed in Corynesporascaceae, while Helminthosporium
clustered in Massarinaceae (Voglmayr and Jaklitsch 2017;
this stuy, Fig. 2).
The massaria- and splachnonema-like sexual morphs
have been linked to Helminthosporium based on cultural
studies and sequence data (Tanaka et al. 2015; Voglmayr
and Jaklitsch 2017). Both sexual and asexual morphs can
be observed in H. massarinum Kaz. Tanaka et al., H. microsorum D. Sacc., H. oligosporum (Corda) S. Hughes, H.
quercicola (M.E. Barr) Voglmayr & Jaklitsch, H. quercinum
Voglmayr & Jaklitsch. and H. tiliae (Link) Fr. (Tanaka et al.
2015; Voglmayr and Jaklitsch 2017). The sexual morph of
the type species H. velutinum Link is still unknown. The
sexual genus Byssothecium has a relationship with Helminthosporium in Massarinaceae (Fig. 2), however, they
can be distinguished by ascospore characters (see key to
freshwater genera of Massarinaceae).
Helminthosporium species are commonly collected from
terrestrial habitats worldwide as mentioned in Voglmayr
and Jaklitsch (2017). Zhu et al. (2016) reported two species
H. aquaticum H.Y. Su et al. and H. velutinum (type) from
freshwater habitats in China (see list below) and no sexual
morphs were observed. We do not make a key for these two
species as they are morphologically very similar and they
have overlapping size in all structures. Even though, multigene phylogenetic analyses supported them to be different
species (Zhu et al. 2016; Voglmayr and Jaklitsch 2017). We
believe that sexual morphs of H. aquaticum and H. velutinum will separate them clearly in future.
List of freshwater Helminthosporium species
*Helminthosporium aquaticum H.Y. Su, Z.L. Luo & K.D.
Hyde, Phytotaxa 253(3): 184 (2016)
Freshwater distribution: Yunnan, China (Zhu et al. 2016)
*Helminthosporium velutinum Link [as ‘Helmisporium’],
Mag. Gesell. naturf. Freunde, Berlin 3(1-2): 10 (1809)
Freshwater distribution: Yunnan, China (Zhu et al. 2016)
Melanommataceae G. Winter [as ‘Melanommeae’],
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 220 (1885)
Key to freshwater genera of Melanommataceae
1.
1.
2.
2.
3.
3.
Asexual morph………………………Camposporium
Sexual morph…………………………………………2
Ascospores dimorphic………………Mamillisphaeria
Ascospores monomorphic……………………………3
Ascomata glabrous………………………Melanomma
Ascomata spread on substrate as a subiculum or covered
by hyphae………………………………………………4
4. Ascomata immersed or erumpent………Herpotrichia
457
4. Ascomata superficial, with red or orange at pore………
…………………………………………Byssosphaeria
Byssosphaeria Cooke, Grevillea 7(no. 43): 84 (1879)
Saprobic on woody angiosperms or submerged wood.
Sexual morph: Ascomata solitary, scattered to gregarious,
superficial, uniloculate, globose, subglobose or ovoid, with
basal subiculum and covered by thick, branched hyphae,
dark brown to black, coriaceous, central ostiolate, red or
orange at pore, without papilla. Peridium two-layered, outer
layer composed of irregular, thick-walled, brown to dark
brown cells of textura epidermoidea, inner layer composed
of small thin-walled, pale brown cells. Pseudoparaphyses
dense, trabeculate, long, embedded in mucilage, anastomosing between and above asci. Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate to clavate, with short pedicel,
apically rounded with an ocular chamber. Ascospores uni- to
bi-seriate, fusiform, tapering at both ends, mostly straight,
sometimes slightly curved, hyaline to pale brown, 1-septate,
verrucose (Tian et al. 2015). Asexual morph: Coelomycetous. Pycnidia formed in culture. Conidiogenous cells phialidic, lining cavity. Conidia hyaline, ellipsoid or subglobose
(Barr 1984).
Type species: Byssosphaeria keithii (Berk. & Broome)
Cooke [as ‘keitii’]
Notes: Byssosphaeria is characterized by superficial ascomata with bright yellow, orange or red, flat apices around
the ostiole, with thick, branched hyphae, and hyaline to pale
brown ascospores (Tian et al. 2015). Byssosphaeria differs
from Herpotrichia Fuckel in having superficial ascomata
with red or orange at pore (Zhang et al. 2012b; Tian et al.
2015). Byssosphaeria was shown to belong to Melanommataceae based on molecular data of some species (Zhang
et al. 2012b; Tian et al. 2015). Sequence data of the type
species B. keithii are unavailable. The freshwater species
B. schiedermayeriana (Fuckel) M.E. Barr was found from
Seychelles, but not supported by molecular data. Three terrestrial strains of B. schiedermayeriana clustered in Melanommataceae with good bootstrap support (Liu et al. 2015;
Tian et al. 2015).
List of freshwater Byssosphaeria species
*Byssosphaeria schiedermayeriana (Fuckel) M.E. Barr,
Mycotaxon 20(1): 34 (1984)
Basionym: Herpotrichia schiedermayeriana Fuckel, Jb.
nassau. Ver. Naturk. 27-28: 27 (1874)
Synonymy: see Index Fungorum (2020)
Freshwater distribution: Seychelles (Hyde and Goh
1998b)
Camposporium Harkn., Bull. Calif. Acad. Sci. 1(no. 1): 37
(1884)
13
458
Saprobic on decaying wood, bark and fruit of various
trees and shrubs in terrestrial or freshwater habitats. Sexual
morph: Undetermined. Asexual morph: Colonies effuse,
grey, brown to black. Mycelium immersed or partly superficial on host substrate. Conidiophores macro- or micronematous, mononematous, often short, straight or irregularly bent,
unbranched, pale brown to dark brown. Conidiogenous cells
mono- or polyblastic, integrated, terminal, sympodial, cylindrical or subulate, denticulate. Conidia solitary, dry, multiseptate, pale brown, often unevenly coloured, the end cells
subhyaline, smooth, mostly cylindrical, rounded or tapering
at the apex, sometimes rostrate, with or without appendages
(Ellis 1971; Hyde et al. 2020b).
Type species: Camposporium antennatum Harkn.
Notes: Camposporium is an old genus and characterized
by short conidiophores, denticulate conidiogenous cells, and
cylindrical, multi-septate, uneven coloured conidia often
with single or several cylindrical appendages (Ellis 1971;
Hyde et al. 2020b). Camposporium had a relationship with
Fusiconidium Jun F. Li et al., but they can be distinguished
by conidiogenous cells and conidial shape (Li et al. 2017;
Hyde et al. 2020b). They are treated as distinct genera until
their relationships are well-resolved (Hyde et al. 2020b).
The type species, C. antennatum, represented by a strain
CBS 113441 was placed in Melanommataceae (Hyde et al.
2020b). Other Camposporium species clustered with C.
antennatum, but with low bootstrap support (Hyde et al.
2020b). A freshwater species C. septatum N.G. Liu et al.
formed a well-supported clade with Fusiconidium species
(Hyde et al. 2020b). However, C. septatum was placed in
Camposporium, rather than Fusiconidium, based on its
monoblastic conidiogenous cells and cylindrical conidia
with several appendages at the apex (Hyde et al. 2020b).
Both C. appendiculatum D.F. Bao et al. and C. multiseptatum D.F. Bao et al. have tapering apical cell with truncate
ends, while C. antennatum has rounded apex (Ellis 1971;
Hyde et al. 2020b). Camposporium hyalinum Abdullah and
C. marylandicum Shearer have the typical apical appendages of Camposporium, but they have hyaline conidia which
are different from the brown, unevenly coloured conidia of
the other freshwater species (Shearer 1974; Abdullah 1980;
Hyde et al. 2020b). The molecular data of these two species
are needed to confirm their phylogenetic placement. Camposporium species need further study with more collections.
List of freshwater Camposporium species
*Camposporium appendiculatum D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Fungal Diversity 100: 77 (2020)
Freshwater distribution: China (Hyde et al. 2020b)
Camposporium hyalinum Abdullah, Trans. Br. mycol. Soc.
75(3): 514 (1981) [1980]
Freshwater distribution: UK (Abdullah 1980)
13
Fungal Diversity (2020) 105:319–575
Camposporium marylandicum Shearer, Mycologia 66(1):
16 (1974)
Freshwater distribution: USA (Shearer 1974)
*Camposporium multiseptatum D.F. Bao, Z.L. Luo, K.D.
Hyde & H.Y. Su, Fungal Diversity 100: 82 (2020)
Freshwater distribution: China (Hyde et al. 2020b)
*Camposporium pellucidum (Grove) S. Hughes, Mycol.
Pap. 36: 9 (1951)
Freshwater distribution: China (Hyde et al. 2020b)
*Camposporium septatum N.G. Liu, J.K. Liu & K.D. Hyde,
Fungal Diversity 100: 85 (2020)
Freshwater distribution: Thailand (Hyde et al. 2020b)
Key to freshwater Camposporium species
1.
1.
2.
2.
3.
3.
4.
4.
5.
Conidial apex without apppendages……C. pellucidum
Conidial apex with appendages………………………2
Conidia with 2–3 appendages………………C. septatum
Conidia with a single appendage………………………3
Conidia 2–4(–6)-septate …………………C. hyalinum
Conidia > 5-septate……………………………………4
Conidia 5–10-septate…………………C. marylandicum
Conidia 10–13-septate…………………………………5
Conidial apppendages 72–114 μm long…………………
……………………………………C. appendiculatum
5. Conidial apppendages 11–17 μm long…………………
………………………………………C. multiseptatum
Herpotrichia Fuckel, Fungi rhenani exsic., suppl., fasc.
7(nos 2101-2200): no. 2171 (1868)
Saprobic, parasitic or hyperparasitic in terrestrial or
freshwater habitats. Sexual morph: Ascomata scattered
to gregarious, immersed, erumpent to nearly superficial,
subglobose to pyriform, dark brown to black, spreading
on substrate as a subiculum, coriaceous, roughened, with
ostiolate papilla. Peridium two-layered, outer layer composed of pigmented, thick-walled cells of textura angularis,
inner layer composed of light pigmented, thin-walled cells
of textura prismatica. Pseudoparaphyses dense, cellular,
hypha-like, hyaline, septate, branched and anastomosing
above asci, embedded in a gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to cylindro-clavate, with
a furcate pedicel, apically rounded with an ocular chamber.
Ascospores uni- to bi-seriate, fusoid, ellipsoidal or oblong,
straight, hyaline to pale brown, 1-septate, thin-walled,
smooth or verruculose (Tian et al. 2015). Asexual morph:
Coelomycetous. pyrenochaeta- or pyrenochaeta-like. Pycnidia solitary or gregarious, superficial, globose to pyriform or hemispherical, with or without hairs, with ostiolate
Fungal Diversity (2020) 105:319–575
papilla, with a long or short neck. Peridium composed of
cells of textura angularis. Conidiophores absent. Conidiogenous cells phialidic, simple, cylindrical, branched. Conidia
globose to ovoid or cylindrical, straight or curved, hyaline,
aseptate (Bose 1961; Sivanesan 1984; Tian et al. 2015).
Type species: Herpotrichia herpotrichoides (Fuckel) P.F.
Cannon
Notes: The type species of Herpotrichia was not indicated when it was introduced (Fuckel 1868). Herpotrichia
herpotrichoides was accepted as the type species (Cannon
1982; Wijayawardene et al. (2017). The holotype of H. rubi
Fuckel (an earlier name of H. herpotrichoides) was examined by Tian et al. (2015) and characterized by ascomata
spreading on substrate as a subiculum and with a black,
roughened wall. Herpotrichia is polyphyletic (Tian et al.
2015). Herpotrichia dalisayi K.D. Hyde & Aptroot is the
only freshwater species in the genus (Hyde and Aptroot
1998a). Herpotrichia dalisayi is characterized by superficial
or partially immersed ascomata with a tangle of dark brown
subiculum and 1–5-septate ascospores, which are characteristics of Herpotrichia (Hyde and Aptroot 1998a). However,
the phylogenetic placement of H. dalisayi in Herpotrichia is
pending confirmation with molecular data. Herpotrichia dalisayi has larger ascospores (53–60 × 16–18 μm) than other
Herpotrichia species (Barr 1984; Hyde and Aptroot 1998a).
List of freshwater Herpotrichia species
Herpotrichia dalisayi K.D. Hyde & Aptroot, Nova Hedwigia 66(1-2): 249 (1998)
Freshwater distribution: Philippines (Hyde and Aptroot
1998a)
Mamillisphaeria K.D. Hyde, S.W. Wong & E.B.G. Jones,
Nova Hedwigia 62(3–4): 514 (1996)
Saprobic on submerged wood. Sexual morph: Ascomata
scattered to gregarious, superficial, conical, dark brown to
black, carbonaceous, with ostiolate papilla, under a pseudostroma that forms a thin layer on the host surface. Peridium
thick, composed of several layers of compressed, hyaline
cells, with palisade-like cells at the sides. Pseudoparaphyses dense, trabeculate, long, hyaline, septate, branching
and anastomosing above the asci, embedded in a gelatinous matrix. Two types of asci and ascospores: TYPE 1:
Asci 8-spored, bitunicate, fissitunicate, cylindro-clavate,
short pedicellate, apically rounded with an ocular chamber. Ascospores 2–4-seriate, ellipsoidal, hyaline, 1-septate,
symmetric, with pad-like, mucilaginous appendage at each
end and with some mucilage associated around the spore.
TYPE 2: Asci 8-spored, bitunicate, fissitunicate, cylindrical,
pedicellate, with an ocular chamber and faint apical ring.
Ascospores uni- to bi-seriate, ellipsoidal-fusoid, brown,
459
1-septate, asymmetric, with a thin mucilaginous sheath
(Tian et al. 2015). Asexual morph: Undetermined.
Type species: Mamillisphaeria dimorphospora K.D.
Hyde, S.W. Wong & E.B.G. Jones
Notes: The monotypic genus Mamillisphaeria is typified
by a freshwater species M. dimorphospora which is strikingly characterized by dimorphic ascospores (Hyde et al.
1996b), reminiscent of Aliquandostipite siamensis (Pang
et al. 2002). However, they differ in ascomata (globose to
subglobose, with stalk in A. siamensis vs. conical, without
stalk in M. dimorphospora) and sheath (ascospores without
sheath in A. siamensis vs. both types of ascospores with
mucilaginous sheaths in M. dimorphospora) (Hyde et al.
1996b; Pang et al. 2002). Aliquandostipite siamensis clustered in Aliquandostipitaceae (Fig. 14), while Tian et al.
(2015) placed M. dimorphospora in Melanommataceae, but
based on morphology. Sequence data of M. dimorphospora
is needed to clarify its phylogenetic placement.
List of freshwater Mamillisphaeria species
Mamillisphaeria dimorphospora K.D. Hyde, S.W. Wong &
E.B.G. Jones, Nova Hedwigia 62(3–4): 515 (1996); Fig. 76
Freshwater distribution: Australia (Bareen and Iqbal
1994; Hyde et al. 1996b; Vijaykrishna and Hyde 2006),
Brunei (Ho et al. 2001), Malaysia (Ho et al. 2001)
Melanomma Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
23–24: 159 (1870) [1869–70]
Saprobic or parasitic on wood and bark in terrestrial and
freshwater habitats. Sexual morph: Ascomata gregarious,
immersed, erumpent to nearly superficial, uniloculate, globose to subglobose, conical, often laterally flattened, black,
coriaceous, often puckered or sulcate, with ostiolate papilla.
Peridium two-layered, arranged in textura angularis, outer
layer comprising small, heavily pigmented, thick-walled
cells, inner layer comprising of hyaline to lightly pigmented,
thin-walled cells. Pseudoparaphyses dense, trabeculate, filamentous, hyaline, indistinctly septate, branching and rarely
anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, cylindrical to fusoid, with a short pedicel, apically
rounded with an ocular chamber. Ascospores uniseriate,
fusoid, pale brown to brown, septate, the second cell from
the apex slightly wider than the others, smooth-walled (Tian
et al. 2015). Asexual morph: Coelomycetous. Conidiomata
superficial, globose, black, ostiolate. Mycelium immersed.
Peridium two-layered. Conidiophores cylindrical, septate,
hyaline. Conidiogenous cells phialidic, hyaline, smooth.
Conidia cylindrical or ellipsoidal, hyaline, aseptate (Sutton
1980).
Type species: Melanomma pulvis-pyrius (Pers.) Fuckel
Notes: As the holotype of Melanomma pulvis-pyrius
was not validly designated, a neotype and epitype were
designated (Holm 1957; Zhang et al. 2008b). Melanomma
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Fig. 76 Mamillisphaeria dimorphospora (BRIP 22967a). (g, j Type
1, h, i, k Type 2) a Specimen and herbarium label. b, c Ascomata
on host surface with a small papilla. d Vertical section of ascoma. e
Structure of peridium. f Trabeculate pseudoparaphyses. g–i Asci. j, k
Ascospores. Scale bars: b = 1 mm, c = 500 μm, d = 100 μm, e = 50
μm, f–i = 20 μm, j, k = 10 μm
pulvis-pyrius clustered with an asexual morphic species
Aposphaeria populina Died., but they are not recognized as
congeneric until the type species of Aposphaeria is obtained
(Tian et al. 2015). The freshwater species M. australiense
K.D. Hyde & Goh normally has larger ascospores than other
Melanomma species (Hyde and Goh 1999b). The phylogenetic placement of M. australiense needs confirmation by
molecular data.
Key to freshwater genera of Morosphaeriaceae
List of freshwater Melanomma species
Melanomma australiense K.D. Hyde & Goh, Nova Hedwigia 68(1–2): 256 (1999); Fig. 77
Freshwater distribution: Australia (Hyde and Goh 1999b;
Vijaykrishna and Hyde 2006)
Morosphaeriaceae Suetrong, Sakay., E.B.G. Jones & C.L.
Schoch, Stud. Mycol. 64: 161 (2009)
13
1. Asci with uncoiled endoascus and hyaline ascospores…
…………………………………………Aquihelicascus
1. Asci with coiled endoascus and brown ascospores……
…………………………………………Neohelicascus
Helicascus Kohlm., Can. J. Bot. 47: 1471 (1969)
Saprobic on rotten roots in sea water, intertidal wood
of mangrove. Sexual morph: Pseudostromata immersed
to semi-immersed, lenticular, solitary to clustered, dark
brown to black, carbonaceous, enclosing single or multilocules, which are horizontally arranged under a black pseudoclypeus, ostiolate. Pseudoclypeus composed of host cells
enclosed in black fungal hyphae, isodiametric or elongate
cells formed between locules. Pseudoparaphyses numerous,
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461
two new genera, Aquihelicascus and Neohelicascus, based
on strong morphological and molecular evidence.
Fig. 77 Melanomma australiense (redrawn from (Hyde and Goh
1999b), HKU(M) 403, holotype). a Section of ascoma. b Bitunicate
ascus. c, d Ascospores. e Pseudoparaphyses. Scale bars: a, b = 50
μm, c, d = 20 μm, e = 10 μm
trabeculate, filiform, persistent, hyaline, septate, anastomosing, embedded in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, subcylindrical, pedicellate, with an ocular
chamber, endoascus coiled at first and finally stretching in
water. Ascospores uniseriate, obovoid, straight to slightly
curved, brown, asymmetrical, unequally 2-celled, with larger
apical cells and smaller basal cells, sometimes at one or both
ends apiculate, septate, smooth or verrucose, with or without
a mucilaginous sheath. Asexual morph: Undetermined.
Type species: Helicascus kanaloanus Kohlm.
Notes: Helicascus kanaloanus was isolated from rotten
roots in sea water (Kohlmeyer 1969). Sequences are available in GenBank for all Helicascus species. Helicascus
comprises three species, viz. H. kanaloanus, H. mangrovei
Preedanon et al. and H. nypae K.D. Hyde, representing a distinct marine genus which is characterized by subcylindrical
asci, uniseriate, obovoid, brown, asymmetrical ascospores.
We exclude other species to Helicascus and place them in
Aquihelicascus W. Dong, H. Zhang & Doilom, gen. nov.
Index Fungorum number: IF557806; Facesoffungi number: FoF08721
Etymology: referring to the aquatic habitat and its similarity to Helicascus
Saprobic on submerged wood. Sexual morph: Pseudostromata scattered, comprising brown to black fungal material growing in cortex of host cells, to uni- or multi-loculate,
flattened at the basal region, horizontally arranged under the
pseudostroma, carbonaceous or coriaceous, visible on the
host surface as blackened ostiolar dots. Locules immersed,
compressed subglobose to lenticular or sometimes triangular, with ostiolate papilla. Ostiole central, rounded, periphysate. Peridium comprising several layers of thin-walled
angular cells, hyaline inwardly and dark at the outside, fusing with the host cells. Pseudoparaphyses numerous, cellular, hyaline, indistinctly septate, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, clavate, long pedicellate,
endoascus uncoiled, apically rounded, with an indistinct
or distinct, trapezoidal ocular chamber. Ascospores mostly
biseriate, sometimes overlapping uniseriate, straight or
slightly curved, ellipsoidal to fusiform, with rounded ends,
1-septate, almost symmetrical, constricted at the septum,
hyaline, smooth, thin-walled, lacking a mucilaginous sheath
or appendage. Asexual morph: Undetermined.
Type species: Aquihelicascus thalassioideus (K.D. Hyde
& Aptroot) W. Dong & H. Zhang
Notes: Aquihelicascus is introduced to accommodate
one new comination A. thalassioideus and two new species
A. songkhlaensis and A. yunnanensis. Aquihelicascus differs from Helicascus in having clavate asci with uncoiled
endoascus, and biseriate, ellipsoidal, symmetrical, hyaline
ascospores with rounded ends. In contrast, Helicascus has
subcylindrical asci with coiled endoascus, and uniseriate,
obovoid, asymmetrical, brown ascospores with apiculate
ends (Kohlmeyer 1969; Hyde 1991; Preedanon et al. 2017).
All Aquihelicascus species were collected from freshwater
habitats, whereas Helicascus species are from marine habitats. Aquihelicascus and Helicascus clustered separately in
phylogenetic analyses (Zhang et al. 2013a; Luo et al. 2016b;
Preedanon et al. 2017; this study, Fig. 84) which supports
the placement of two different genera.
Aquihelicascus species have hyaline ascospores and are
morphologically very similar. Among them, A. songkhlaensis possesses two-loculate pseudostromata and A. thalassioideus has the widest peridium. The phylogenetic analysis and
single gene comparison can easily separate them.
List of freshwater Aquihelicascus species
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Fig. 78 Aquihelicascus songkhlaensis (MFLU 18-1511,
holotype). a Immersed ascomata with papilla erumpent
through host surface. b Vertical
section of pseudostroma. c
Structure of medium peridium.
d Structure of lateral peridium.
e–g Bitunicate asci. h Pedicel. i
Pseudoparaphyses. j Ascospore
in Indian Ink. k–m Ascospores.
n, o Colony on PDA (left-front,
right-reverse). Scale bars: b =
200 μm, c–g, i = 20 μm, h = 10
μm, j–m = 5 μm
*Aquihelicascus songkhlaensis W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557919; Facesoffungi number: FoF09261; Fig. 78
Etymology: referring to songkhla, where the holotype was
collected
Holotype: MFLU 18-1511
Saprobic on decaying wood submerged in freshwater. Sexual morph: Pseudostromata 380–430 μm high,
820–870 μm wide, scattered, comprising brown to black
13
fungal tissues growing in cortex of host, with two locules,
flattened at the basal region, horizontally arranged under the
pseudostroma, visible on the host surface as blackened ostiolar dots. Locules 170–270 × 350–430 μm, immersed, lenticular, coriaceous, with ostiolate papilla. Ostiole converging at the centre, subcylindrical, black. Peridium of locules,
25–35 μm thick, comprising 8–11 layers of brown to dark
brown, thin-walled, compressed cells of textura angularis,
fusing with the host cells. Pseudoparaphyses 3 μm diam.,
numerous, cellular, hypha-like, hyaline, distantly septate.
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463
Fig. 79 Aquihelicascus thalassioideus (MFLU 18-1705). a, b Immersed pseudostromata with blackened ostiolar dots. c Ascus embedded in
pseudoparaphyses. d, e Bitunicate asci. f, g Ascospores. Scale bars: c–e = 50 μm, f, g = 20 μm
Asci 110–145(–160) × 16–18 μm ( x̄ = 130 × 17 μm, n =
10), 8-spored, bitunicate, clavate, long pedicellate, up to 50
μm long, apically rounded with an indistinct ocular chamber.
Ascospores 24–28 × 8–10 μm ( x̄ = 25.7 × 9.4 μm, n = 20),
overlapping biseriate, straight or slightly curved, hyaline,
1-septate, constricted at the septum, symmetrical, ellipsoidal
with rounded ends, with 2–4 prominent big guttules when
immature, smooth, thin-walled, lacking a mucilaginous
sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 13 days at 25 °C, white to pale grey from
above, dark brown from below, surface rough, dry, raised,
edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
20180508-1 (MFLU 18-1511, holotype), ex-type living culture MFLUCC 18-1154; ibid., 20180508-2 (HKAS 105006,
isotype), ex-type living culture KUMCC 19-0022; Nakhon
Si Thammarat Province, on submerged wood in a stream,
10 May 2018, W. Dong, hat6107-1 (MFLU 18-1716), living culture MFLUCC 18-1273; ibid., hat6107-2 (HKAS
105028), living culture KUMCC 19-0043; Songkhla Province, on submerged wood in a stream, 10 May 2018, W.
Dong, hat8114-1 (MFLU 18-1702), living culture MFLUCC
18-1278; ibid., hat8114-2 (HKAS 105036), living culture
KUMCC 19-0051.
Notes: Aquihelicascus songkhlaensis clusters with A.
yunnanensis in our phylogenetic tree (Fig. 84). They share
similar morphological characters of clavate asci, ellipsoidal, hyaline, 1-septate, symmetrical ascospores and are from
freshwater habitats and they have overlapping ascospore
size. However, A. songkhlaensis can be distinguished by its
two-loculate pseudostromata contrasting with one-loculate
in A. yunnanensis. There are 12, 16 and 14 nucleotide differences between ex-type strains of the two species in LSU,
ITS and TEF sequence data, respectively. Following the
guideline of Jeewon and Hyde (2016), A. songkhlaensis sp.
nov. is introduced here.
*Aquihelicascus thalassioideus (K.D. Hyde & Aptroot) W.
Dong & H. Zhang, comb. nov.
Index Fungorum number: IF557920; Facesoffungi number: FoF09262; Fig. 79
Basionym: Massarina thalassioidea K.D. Hyde & Aptroot, Nova Hedwigia 66(3–4): 498 (1998)
Synonymy: Helicascus thalassioideus (K.D. Hyde & Aptroot) H. Zhang & K.D. Hyde, Sydowia 65(1): 159 (2013)
Freshwater distribution: Australia (Hyde and Aptroot
1998b), Brunei (Hyde and Aptroot 1998b), China (Tsui
et al. 2000; Ho et al. 2001; Luo et al. 2004), French West
Indies, Martinique (Zhang et al. 2015), Peru (Shearer et al.
2015), Janpan (Tanaka et al. 2015), Philippines (Hyde and
Aptroot 1998b), Thailand (Kurniawati et al. 2010; Zhang
et al. 2013a; this study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Pseudostromata clustered, immersed, visible on the host surface as blackened ostiolar dots. Pseudoparaphyses 1–3 μm diam., abundant, cellular, hyphalike, hyaline, septate, embedded in a gelatinous matrix.
Asci 150–190 × 14.5–23 μm ( x̄ = 175 × 17.5 μm, n = 5),
8-spored, bitunicate, fissitunicate, clavate, long pedicellate,
up to 90 μm long. Ascospores 24–28 × 7–10 μm ( x̄ = 25.5
× 8.5 μm, n = 25), overlapping uni- to bi-seriate, sometimes overlapping tri-seriate, ellipsoidal, hyaline, 1-septate,
constricted at septum, curved, thin-walled, smooth. Asexual
morph: Undetermined (detailed description see Hyde and
Aptroot (1998b) and Zhang et al. (2013a)).
Material examined: THAILAND, Chiang Rai Province, on submerged wood in a stream, 1 July 2018, W.
Dong, CR134-1 (MFLU 18-1705), living culture KUMCC
19-0094; ibid., CR134-2 (HKAS 105076).
Notes: Our new collection KUMCC 19-0094 is identified as Aquihelicascus thalassioideus based on multigene
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phylogenetic analysis (Fig. 84). The morphological features of our collection also fits well with A. thalassioideus
except for the long pedicel, which is probably because of the
pedicel spreading in water (Fig. 79). Most publications do
not report an ascospore sheath for A. thalassioideus except
Zhang et al. (2015) who noted some fugacious mucilaginous
remnants visible in Indian Ink when ascospores were just
released from the asci. This character was not observed in
our collection.
Aquihelicascus yunnanensis is morphologically similar
to A. thalassioideus in having clavate asci, and ellipsoidal,
hyaline, 1-septate, guttuate ascospores (Zhang et al. 2013a).
However, A. yunnanensis has a thinner peridium (15–25
μm vs. 70 μm) than A. thalassioideus. There are 10 and
32 nucleotide differences in LSU and ITS sequence data
between A. yunnanensis (MFLUCC 18-1025) and A. thalassioideus (MFLUCC 10-0911), respectively, which indicates
them to be different species.
*Aquihelicascus yunnanensis W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557921; Facesoffungi number: FoF09263; Fig. 80
Etymology: referring to Yunnan, where the holotype was
collected
Holotype: MFLU 18-1170
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Pseudostromata 150–180 μm high, 420–480
μm diam., scattered, comprising brown to black fungal tissues growing in cortex of host, uniloculate, lenticular,
immersed with blackened ostiolar dots on the host surface.
Ostiole central, opening rounded, periphysate. Peridium
15–25 μm thick, comprising several layers of brown to dark
brown, thin-walled, compressed, elongate cells of textura
angularis, pale brown to hyaline inwardly. Pseudoparaphyses 1.5–2.5 μm diam., numerous, cellular, hypha-like,
short, hyaline, sparsely septate, unbranched. Asci 120–140
× 13.5–19 μm ( x̄ = 132 × 17 μm, n = 10), 8-spored, bitunicate, clavate, apically narrowly rounded, with a distinct, trapezoidal chamber, long pedicellate, up to 40 μm. Ascospores
25–28 × 9–10.5 μm ( x̄ = 27 × 10 μm, n = 10), biseriate,
ellipsoidal, rounded at both ends, straight or slightly curved,
hyaline, 1-septate, constricted at the septum, symmetrical,
guttulate, two prominent guttules near the septa and with
additionally smaller ones beside, smooth, thin-walled, lacking a mucilaginous sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 15 days at 25 °C, white to grey from above,
brown from below, surface rough, fluffy, with dense mycelium, dry, raised, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H35A-1 (MFLU 18-1170, holotype), ex-type living
culture MFLUCC 18-1025; ibid., H35A-2 (HKAS 101719,
isotype), ex-type living culture KUMCC 18-0069.
Notes: Aquihelicascus yunnanensis clusters with A. songkhlaensis in our multigene phylogenetic analysis (Fig. 84).
However, the number of locules in pseudostromata, single
gene comparison (see notes under A. songkhlaensis) and
phylogenetic analysis (Fig. 84) separate them to be different
species.
Key to freshwater Aquihelicascus species
13
1.
1.
2.
2.
Pseudostromata multi-loculate………A. songkhlaensis
Pseudostromata uni-loculate…………………………2
Peridium 15–25 μm thick………………A. yunnanensis
Peridium up to 70 μm thick…………A. thalassioideus
Neohelicascus W. Dong, H. Zhang, K.D. Hyde & Doilom,
gen. nov.
Index Fungorum number: IF557807; Facesoffungi number: FoF08722
Etymology: referring to its morphological similarity to
Helicascus
Saprobic on submerged wood. Sexual morph: Pseudostromata scattered, comprising brown to black fungal
material growing in cortex of host, unil- or multi-loculate,
flattened at the basal region, horizontally arranged under
the pseudostroma, visible on the host surface as blackened
ostiolar dots. Locules immersed, lenticular to ampulliform,
ostiolate. Ostiole converging at the centre, uniting into one
common, central pore, periphysate. Peridium of locules
comprising several layers of brown, thin-walled angular
cells, paler inwardly, fusing with the host cells. Pseudoparaphyses numerous, cellular, hypha-like, hyaline, septate,
embedded in a gelatinous matrix. Asci 8-spored, bitunicate,
fissitunicate, clavate, long pedicellate, base of endoascus
long, narrow and coiled within ectoascus, ectoascus uncoiling to form a long tail-like extension, apically rounded with
a cylindrical ocular chamber. Ascospores mostly biseriate,
straight or slightly curved, ellipsoidal to broadly fusiform,
with rounded ends, 1–(2–3)-septate, apical cell slightly
larger than basal cell, brown, thin-walled, smooth- or roughwalled, with or without a deliquescing sheath (Zhang et al.
2013a). Asexual morph: Coelomycetous. Pycnidia solitary,
superficial, with the base immersed, uniloculate, globose to
subglobose, brown, coriaceous, central ostiolate, with minute papilla. Peridium composed of brown, thin-walled cells
of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, determinate,
cylindrical to subcylindrical, hyaline, smooth. Conidia ellipsoid to obovoid, hyaline, aseptate, occasionally two-celled,
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Fig. 80 Aquihelicascus yunnanensis (MFLU 18-1170,
holotype). a, b Immersed
ascomata with blackened
ostiolar dots on the host surface.
c Vertical section of ascoma. d
Structure of ostiole. e Structure
of peridium. f–i Bitunicate
asci. j Pseudoparaphyses. k–n
Ascospores. o Germinated
ascospores. p Colony on PDA
(up-front, down-reverse). Scale
bars: c = 100 μm, d = 50 μm, e,
i–n = 10 μm, f–h, o = 20 μm
smooth, thin-walled, without sheath or appendage (Zhang
et al. 2013a).
Type species: Neohelicascus aquaticus (H. Zhang & K.
D. Hyde) W. Dong & H. Zhang
Notes: Neohelicascus is introduced to accommodate the
new species N. submersus H. Yang et al. and another seven
new combinations as listed below. Neohelicascus is distinguished from Aquihelicascus in producing brown ascospores
and the base of endoascus is long, narrow and coiled within
ectoascus which uncoils to form a long tail-like extension.
In contrast, Aquihelicascus possesses hyaline ascospores and
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Fig. 81 Neohelicascus
aquaticus (HKAS 102145). a,
b Immersed pseudostromata
with blackened ostiolar dots. c
Pseudoparaphyses. d Bitunicate ascus. e, f Ascospores. g,
h Colony on PDA (up-front,
down-reverse). Scale bars: c, f
= 10 μm, d = 30 μm, e = 20 μm
an uncoiled endoascus. Phylogenetic analysis supports two
genera, Aquihelicascus and Neohelicascus (Fig. 84).
The ascospores of Neohelicascus egyptiacus and N. elaterascus are surrounded by a defined gelatinous sheath, which is
absent in N. unilocularis (Zhang et al. 2013a, 2015). Neohelicascus gallicus is surrounded by some fugacious mucilaginous
remnants, but only seen in ascospores that are just released
from the asci (Zhang et al. 2014b). Neohelicascus aquaticus
was characterized by 1–(2– or 3–) -septate ascospores with
a deliquescing sheath (Zhang et al. 2013a), although Tanaka
et al. (2015) observed only 1-septate ascospores without a
sheath from a Japanese collection. Neohelicascus aquaticus is
the only species producing asexual morph in culture (Zhang
et al. 2013a).
List of freshwater Neohelicascus species
*Neohelicascus aquaticus (H. Zhang & K. D. Hyde) W.
Dong, K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557922; Facesoffungi number: FoF09264; Fig. 81
Basionym: Helicascus aquaticus H. Zhang & K.D. Hyde,
Sydowia 65(1): 155 (2013)
Synonymy: Helicascus alatus M. Zeng, S.K. Huang, Q.
Zhao & K.D. Hyde, Phytotaxa 351(3): 215 (2018)
Freshwater distribution: China (Zeng et al. 2018; this
study), Japan (Tanaka et al. 2015), Thailand (Zhang et al.
2013a)
Saprobic on decaying wood submerged in freshwater. Sexual morph: Pseudostromata clustered, immersed,
ampulliform, coriaceous, with ostiolate papilla, visible on
the host surface as blackened ostiolar dots. Pseudoparaphyses 1–3 μm diam., abundant, cellular, hypha-like, hyaline,
septate, embedded in a gelatinous matrix. Asci 90–120 ×
14–21 μm ( x̄ = 110 × 19 μm, n = 5), 8-spored, bitunicate,
fissitunicate, clavate, pedicellate. Ascospores 20–23 × 8–10
μm ( x̄ = 22 × 8.5 μm, n = 15), mostly biseriate, ellipsoidal, brown, 1-septate, slightly constricted at septum, asymmetrical, apical cell usually longer than basal cell, curved,
thin-walled, smooth, with a deliquescing sheath. Asexual
morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 40 mm in 50 days at 25 °C, reddish brown to black
13
from above, reddish brown from below, surface rough, dry,
umbonate, edge entire.
Material examined: CHINA, Yunnan Province, Qujing,
on submerged wood in a stream, 10 May 2017, C.X. Liu,
L25 (HKAS 102145), living culture KUMCC 19-0107.
Notes: The LSU and ITS sequence data from our collection HKAS 102145 shows 100% identity with the sequences
of Helicascus aquaticus (MFLUCC 10-0918). HKAS
102145 fits well with the morphological features of H.
aquaticus (Zhang et al. 2013a). However, we did not observe
the 3-septate ascospores and asexual morph in culture.
Helicascus alatus was introduced by Zeng et al. (2018)
based on multigene phylogenetic analysis and morphological characters. Zeng et al. (2018) considered that H. alatus
differs from H. aquaticus in smaller ascomata, short pedicels and presence of a sheath. However, a comparion of
nucleotides shows that H. alatus (MFLUCC 17-2300) and
H. aquaticus (MFLUCC 10-0918) have identical sequences
in LSU and ITS sequence data. Additionally, there are only
two nucleotide differences in TEF sequence data between
H. alatus (MFLUCC 17-2300) and H. aquaticus (MAFF
243866), thus H. alatus is a synonymy of H. aquaticus.
Phylogenetic analysis supports these strains to be the same
species (Fig. 84).
*Neohelicascus chiangraiensis (Z.L. Luo, J.K Liu, H.Y. Su
& K.D. Hyde) W. Dong, K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557923; Facesoffungi number: FoF09265
Basionym: Helicascus chiangraiensis Z.L. Luo, H.Y. Su
& K.D. Hyde, Phytotaxa 270(3): 185 (2016)
Freshwater distribution: Thailand (Luo et al. 2016b)
*Neohelicascus egyptiacus (Abdel-Wahab & Abdel-Aziz)
W. Dong, K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557924; Facesoffungi number: FoF09266
Basionym: Helicascus egyptiacus Abdel-Wahab & AbdelAziz [as ‘aegyptiacus’], Sydowia 65(1): 153 (2013)
Freshwater distribution: Egypt (Zhang et al. 2013a)
Fungal Diversity (2020) 105:319–575
467
Fig. 82 Neohelicascus elaterascus (c, d, h, j–m from MFLU
18-1007, e–g, i, n–p from
MFLU 18-0997). a, b Immersed
pseudostromata with blackened
ostiolar dots. c Vertical section
of pseudostroma. d Structure of
peridium. e–h Bitunicate asci.
i–m Ascospores. (l, m in Indian
Ink). n Germinated ascospore.
o, p Colony on PDA (left-front,
right-reverse). Scale bars: c =
50 μm, d–n = 20 μm
*Neohelicascus elaterascus (Shearer) W. Dong, K.D. Hyde
& H. Zhang, comb. nov.
Index Fungorum number: IF557925; Facesoffungi number: FoF09267; Fig. 82
Basionym: Kirschsteiniothelia elaterascus Shearer, Mycologia 85:963 (1993).
Synonymy: Morosphaeria elaterascus (Shearer) S. Boonmee & K.D. Hyde, Mycologia 103: 705 (2012)
Helicascus elaterascus (Shearer) H. Zhang & K.D. Hyde,
Sydowia 65: 158 (2013)
Freshwater distribution: Brunei (Ho et al. 2001), Chile
(Shearer 1993b), China (Tsui et al. 2000; Luo et al. 2004),
Japan (Tanaka et al. 2015), Peru (Shearer et al. 2015), South
Africa (Hyde et al. 1998), Thailand (Hyde et al. 2020a; this
study), USA (Shearer 1993b; Raja et al. 2009b)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Pseudostromata 205–250 μm high, 170–200
μm diam., clustered, immersed, unilocular, mammiform,
coriaceous, with ostiolate papilla, visible on the host surface as blackened ostiolar dots. Peridium 20–30 μm thick,
13
468
comprising several layers of brown, thin-walled, pseudoparenchymatous cells, outer layer partially occluded with
the host cells, deeply pigmented, inwardly hyaline, forming
a textura angularis in the upper regions and a textura prismatica at sides and angles of base. Pseudoparaphyses 1–3
μm diam., abundant, cellular, cylindrical, hyaline, sparsely
septate. Asci 95–130 × 17–24 μm ( x̄ = 112 × 21 μm, n =
15), 8-spored, bitunicate, fissitunicate, clavate, endoascus
separating from ectoascus at time of ectoascus dehiscence,
base of endoascus long, narrow and coiled within ectoascus,
up to 210 μm long. Ascospores 23–28 × 8–11 μm (x̄ = 25
× 9 μm, n = 25), overlapping uni- or bi-seriate, sometimes
overlapping tri-seriate, ellipsoidal, brown, 1-septate, constricted at septum, asymmetrical, apical cell slightly longer
than basal cell, curved, thin-walled, smooth in optical microscope, surrounded by a gelatinous sheath. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 40 mm in 45 days at 25 °C, dark brown to black from
above, black from below, surface rough, dry, raised, edge
undulate.
Material examined: THAILAND, Phayao Province, on
submerged wood in a stream, 23 February 2018, X.D. Yu,
Y19 (MFLU 18-0997), living culture MFLUCC 18-0985;
Chiang Mai Province, Mushroom Research Center, on
submerged wood in a stream, 2 April 2018, X.D. Yu, Y29
(MFLU 18-1007), living culture MFLUCC 18-0993.
Notes: Our new collections MFLUCC 18-0985 and
MFLUCC 18-0993 are identified as Neohelicascus elaterascus based on morphological and molecular evidence
(Fig. 84). Shearer (1993b) described verrucose ascospores
from the holotype (C-76-1), while our collections have
smooth ascospores which are also described by Hyde et al.
(2020a). Helicascus elaterascus is commonly reported on
submerged wood from many countries as listed above.
*Neohelicascus gallicus (Y. Zhang ter & J. Fourn) W. Dong,
K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557926; Facesoffungi number: FoF09268
Basionym: Helicascus gallicus Y. Zhang ter & J. Fourn.,
Phytotaxa 183: 185 (2014)
Freshwater distribution: France (Zhang et al. 2014b)
*Neohelicascus submersus H. Yang, W. Dong, K.D. Hyde
& H. Zhang, sp. nov.
Index Fungorum number: IF557927; Facesoffungi number: FoF09269; Fig. 83
Etymology: referring to submerged habitat of this fungus
Holotype: MFLU 20-0436
Saprobic on submerged wood. Sexual morph: Pseudostromata 485–540 μm high, 315–510 μm diam., scattered,
comprising dark brown to black fungal tissues growing in
13
Fungal Diversity (2020) 105:319–575
cortex of host, immersed, unilocular, lenticular, flattened
at the base, black, with wide ostiole, visible as elongate,
prominent, black neck erumpent on the host surface. Neck
310–320 × 125–165 μm, central, cylindrical, with periphyses. Peridium 20–45 μm thick, comprising several layers
of dark brown cells of textura epidermoidea. Pseudoparaphyses 1.5–2.5 μm wide, numerous, cellular, cylindrical,
hyaline, indistinctly septate. Asci 120–165 × 17.5–21 μm
( x̄ = 143 × 19 μm, n = 10), 8-spored, bitunicate, fissitunicate, cylindrical to narrowly clavate, apically rounded, base
of endoascus long, narrow and coiled within ectoascus, up
to 150 μm long. Ascospores 23.5–31 × 8.5–11.5 μm, ( x̄
= 27 × 10 μm, n = 30), obliquely arranged, overlapping
uniseriate or occasionally biseriate, ellipsoidal to fusiform,
slightly curved, 1-septate, constricted at the septum, upper
cell slightly wider than lower cell, hyaline when young,
become brown when mature, guttulate, thin-walled, smooth,
covered by a distinct gelatinous sheath, 1.5–5.5 μm wide,
showing 15–27 μm wide in Indian Ink. Asexual morph:
Undetermined.
Cultural characteristics: Conidia germinated on PDA
within 24 hours. Germ tubes arising from end of the
ascospore. Colonies on PDA reaching 30 mm diameter in 30
days at 20–25 °C, initially white, turning reddish brown to
dark brown after 15 days, with dense, hairy mycelium on the
surface, with undulate, red margin, reverse reddish brown.
Material examined: CHINA, Yunnan Province, a small
river in Puzhehei wetland, on submerged wood, 23 June
2019, H. Yang, P36 (MFLU 20-0436, holotype), ex-type
living culture KUMCC 20–0153.
Notes: Neohelicascus submersus forms a well-supported
clade with four species N. elaterascus, N. egyptiacus, N.
unilocularis and N. uniseptatus (Fig. 84). Neohelicascus
submersus resembles N. elaterascus in having unilocular pseudostromata, and ellipsoidal to fusiform, brown
ascospores with a distinct gelatinous sheath (Shearer
1993b). However, N. elaterascus has shorter and wider asci
(95–149 × (14–)18–25(–38) μm vs. 120–165 × 17.5–21 μm)
and ascospores are mostly biseriate and verrucose in the
holotype C-76-1 (Shearer 1993b). Our two collections of
N. elaterascus (MFLUCC 18-0993 and MFLUCC 18-0985)
have smooth ascospores, which are similar to N. submersus,
but phylogenetic analysis separate them as distinct species
(Fig. 84).
Neohelicascus submersus is phylogenetically closest to
N. unilocularis (Fig. 84). They have overlapping ascospore
size, however, N. submersus has longer and thinner asci
(120–165 × 17.5–21 μm vs. 70–75 × 22–27 μm) than N.
unilocularis (Zhang et al. 2015). Neohelicascus submersus
has a distinct gelatinous ascospore sheath, 1.5–5.5 μm wide,
showing 15–27 μm wide in Indian Ink, but it is lacking in
N. unilocularis. There are two and 39 nucleotide differences in LSU and ITS sequence data between N. submersus
Fungal Diversity (2020) 105:319–575
Fig. 83 Neohelicascus submersus (MFLU 20-0436, holotype). a, b
Immersed pseudostromata with blackened neck. c, d Vertical section
of pseudostroma. e Structure of peridium. f Pseudoparaphyses. g–k
469
Bitunicate asci. l–o Ascospores. p Germinated ascospore. q, r Colony
on PDA (up-front, down-reverse). Scale bars: c = 100 μm, d = 50
μm, e–j, l–p = 20 μm, k = 10 μm
13
470
Fungal Diversity (2020) 105:319–575
Fig. 84 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS and TEF sequence
data for species of Morosphaeriaceae. Bootstrap values for
maximum likelihood equal to or
greater than 75% and Bayesian
posterior probabilities equal to
or greater than 0.95 are placed
near the branches as ML/BYPP.
Newly generated sequences are
in red and ex-type strains are
in bold. The new species introduced in this study are indicated
with underline. Freshwater
strains are indicated with a red
letter “F”. The tree is rooted to
Latorua caligans CBS 576.65
(Latoruaceae)
Neohelicascus elaterascus MAFF 243867 F
Neohelicascus elaterascus CBS 139689 F
Neohelicascus elaterascus HKUCC 7769
Neohelicascus elaterascus MFLUCC 18-0993 F
Neohelicascus elaterascus MFLUCC 18-0985 F
Neohelicascus aegyptiacus FWCC 99 F
Neohelicascus uniseptatus MFLUCC 15-0057 F
89/0.99 Neohelicascus unilocularis MJF 14020-1 F
Neohelicascus unilocularis MJF 14020 F
Neohelicascus submersus KUMCC 20–0153 F
Neohelicascus aquaticus MFLUCC 17-2300 F
100/1.00
Neohelicascus aquaticus KUMCC 17-0145 F
100/1.00
Neohelicascus aquaticus MFLUCC 10-0918 F
Neohelicascus aquaticus MAFF 243866 F
89/1.00
Neohelicascus aquaticus KUMCC 19-0107 F
97/0.99
Neohelicascus gallicus CBS 123118 F
94/1.00
Neohelicascus gallicus BJFUCC 200228 F
76
Neohelicascus gallicus BJFUCC 200224 F
Neohelicascus chiangraiensis MFLUCC 13-0883 F
78/0.99 Aquihelicascus thalassioideus MJF 14020-2 F
Aquihelicascus thalassioideus KUMCC 19-0094 F
Aquihelicascus thalassioideus CBS 110441 F
100/1.00
Aquihelicascus thalassioideus KH 242 F
100/1.00
Aquihelicascus thalassioideus MFLUCC 10-0911 F
100/1.00
Aquihelicascus songkhlaensis MFLUCC 18-1154 F
100/1.00
Aquihelicascus songkhlaensis MFLUCC 18-1273 F
Aquihelicascus songkhlaensis MFLUCC 18-1278 F
Aquihelicascus yunnanensis MFLUCC 18-1025 F
Helicascus mangrovei BCC 68260
100/1.00
Helicascus mangrovei BCC 74471
100/1.00
Helicascus mangrovei BCC 68258
81/-100/1.00
Helicascus kanaloanus ATCC 18591
Helicascus nypae BCC 36752
100/1.00
Helicascus nypae BCC 36751
Morosphaeria ramunculicola BCC 18404
98/0.98
Morosphaeria ramunculicola KH 220
100/1.00
Morosphaeria ramunculicola BCC 18405
Morosphaeria ramunculicola JK 5304B
98/1.00
100/1.00 Morosphaeria velatispora KH 221
Morosphaeria velatispora BCC 17059
Clypeoloculus hirosakiensis CBS 139682
100/1.00
Clypeoloculus microsporus CBS 139683
100/1.00
Clypeoloculus akitaensis CBS 139681
Clypeoloculus
towadaensis CBS 139685
99/0.99
Aquilomyces patris CBS 135661
100/0.99
Aquilomyces patris CBS 135760
Aquilomyces patris CBS 135662
100/1.00
Aquilomyces rebunensis CBS 139684
Latorua caligans CBS 576.65
Neohelicascus*
Aquihelicascus*
Helicascus
Morosphaeria
Clypeoloculus
Aquilomyces
Outgroup
0.03
(KUMCC 20–0153) and N. unilocularis (MJF 14020), which
indicates them to be different species (Jeewon and Hyde
2016).
*Neohelicascus unilocularis (J. Fourn. & Y. Zhang ter) W.
Dong, K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557928; Facesoffungi number: FoF09270
13
Basionym: Helicascus unilocularis J. Fourn. & Y. Zhang
ter, Mycol. Progr. 14(7/47): 3 (2015)
Freshwater distribution: French West Indies, Martinique
(Zhang et al. 2015)
Fungal Diversity (2020) 105:319–575
*Neohelicascus uniseptatus (J. Yang, J.K. Liu & K.D.
Hyde) W. Dong, K.D. Hyde & H. Zhang, comb. nov.
Index Fungorum number: IF557929; Facesoffungi number: FoF09271
Basionym: Helicascus uniseptatus J. Yang, J.K. Liu &
K.D. Hyde, Phytotaxa 270(3): 187 (2016)
Freshwater distribution: Thailand (Luo et al. 2016b)
Key to freshwater Neohelicascus species
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
Pseudostromata uni-loculate…………………………2
Pseudostromata mostly multi-loculate…………………6
Ascospores rough-walled………………………………3
Ascospores smooth-walled……………………………4
Peridium 60–70 μm, ascospores 1–(2–3)-septate………
…………………………………………N. elaterascus
Peridium 30–50 μm, ascospores 1-septate………………
………………………………………N. chiangraiensis
Ascospores with sheath…………………N. submersus
Ascospores lacking sheath or with deliquescing
sheath…………………………………………………5
Peridium 50–65 μm, ascospores (21.6–)23–25.8(–27.3)
× (9.2–)10.2–11.4(–11.7) μm……… …N. unilocularis
Peridium 40–48 μm, ascospores 25–32 × 9–13 μm……
…………………………………………N. uniseptatus
Ascospores rough-walled………………N. aegyptiacus
Ascospores smooth-walled……………………………7
Ascospores (24.2–)25.1–31.3(–33) × (8.8–)9.3–12.1(–
12.6) μm……………………………………N. gallicus
Ascospores 19−26 × 8−11 μm……………N. aquaticus
471
solitary to gregarious, immersed to partly-immersed, globose
to subglobose, black, with a short, black papilla. Peridium
comprising several layers of hyaline to dark cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells enteroblastic, phialidic, determinate,
ampulliform to cylindrical, hyaline, smooth-walled. Conidia
oblong, rounded at the apex, aseptate, hyaline to pale brown,
straight, smooth-walled (Dai et al. 2017).
Type species: Occultibambusa bambusae D.Q. Dai &
K.D. Hyde
Notes: Occultibambusa species commonly occur on
culms of bamboo (Hyde et al. 2016b; Dai et al. 2017;
Zhang et al. 2017b). Occultibambusa is characterized by
uniloculate ascostromata, broadly cylindrical to clavate asci
and fusiform, 1-septate ascospores (Dai et al. 2017). The
asexual morph developed in a culture of O. fusispora Phook.
et al. with oblong, hyaline to pale brown, aseptate conidia
(Dai et al. 2017). Occultibambusa species cluster in a wellsupported clade, except O. fusispora and O. maolanensis Jin
F. Zhang et al. cluster with Versicolorisporium triseptatum
Sat. Hatak. et al. in a separated clade (Fig. 89).
Occultibambusa aquatica H. Zhang & K.D. Hyde was
isolated from submerged bamboo in Thailand and was characterized by 1-septate ascospores which become 3-septate
when germinated (Hyde et al. 2016b). The two additional
septa formed at maturity and they were also observed in two
other freshwater species, O. kunmingensis sp. nov. and O.
pustula D.Q. Dai & K.D. Hyde. Three freshwater species
can be distinguished by ascospore size (see key below).
List of freshwater Occultibambusa species
Occultibambusaceae D.Q. Dai & K.D. Hyde, Fungal
Diversity 82: 25 (2016)
Occultibambusa D.Q. Dai & K.D. Hyde, Fungal Diversity
82: 26 (2016)
Saprobic on dead bamboo culms or submerged wood.
Sexual morph: Ascostromata solitary, scattered, or in small
groups, immersed under host tissue, visible as dark, round,
black spots on host surface, uniloculate, subglobose or conical in section, black at ostiolar region, coriaceous, with a
central papilla and rounded ostiole, internally lined with periphyses. Peridium comprising host and fungal tissues, composed of brown and thick-walled cells in outer layer, hyaline
and thin-walled cells inwardly, arranged in textura angularis.
Pseudoparaphyses numerous, cellular, long, hypha-like, septate. Asci 8-spored, bitunicate, broadly cylindrical to clavate,
with a short pedicel and an ocular chamber. Ascospores bito tri-seriate, broadly fusiform to narrowly fusiform, some
with acute ends, 1-septate, slightly constricted at the septum,
pale brown to brown, straight to curved (Dai et al. 2017).
Asexual morph: Coelomycetous. Pycnidia eustromatic,
*Occultibambusa aquatica H. Zhang & K.D. Hyde, Fungal
Diversity 80: 81 (2016); Fig. 85
Freshwater distribution: Thailand (Hyde et al. 2016b)
*Occultibambusa kunmingensis C.X. Liu, H. Zhang &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557930; Facesoffungi number: FoF09272; Fig. 86
Etymology: referring to Kunming, where the holotype
was collected
Holotype: HKAS 102151
Saprobic on decaying bamboo submerged in freshwater.
Sexual morph: Ascomata 110–150 μm high, 220–260 μm
diam., black, scattered, semi-immersed to superficial, with a
short neck, ellipsoidal, ostiolate, flattened at the base. Ostiolar neck black, short, always broken off, with a large opening on the surface of ascomata. Peridium 30–50 μm thick,
thin at the base and becoming wider laterally, composed
of several layers of brown to dark brown, thin-walled cells
of textura angularis. Pseudoparaphyses 2.2 μm wide, cellular, hypha-like, hyaline, septate. Asci 8-spored, bitunicate,
13
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Fig. 85 Occultibambusa aquatica (Material examined: THAILAND,
Chiang Rai Province, on submerged bamboo, 16 November 2010, H.
Zhang, a50, MFLU 11-1141, holotype). a Appearance of ascomata
on host surface. b Vertical section through ascoma. c Ascus. d Pseudoparaphyses. e–f Ascospores. f Ascospore in Indian Ink. Scale bars:
b = 100 μm, c = 20 μm, d–f = 10 μm
clavate or cylindric-clavate, 110–140(–160) × 13–16.5 μm
( x̄ = 127 × 15.5 μm, n = 10), narrowly rounded at the apex,
with a distinct ocular chamber, pedicellate. Ascospores
32–40 × 5–6.5 μm ( x̄ = 35.5 × 5.5 μm, n = 30), 3–4-seriate, fusiform, straight or curved, brown, 1-septate, becoming
3-septate when germinated, constricted at the septa, upper
cell slightly shorter and wider than lower part, guttulate,
thin-walled, smooth, without any mucilaginous sheaths and
appendages. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 30 mm in 30 days at 25 °C, dark brown from above,
black from below, surface rough, with dense mycelium, dry,
raised, edge entire.
Material examined: CHINA, Yunnan Province, Kunming University of Science and Technology, on submerged
bamboo in a stream, 10 May 2017, C.X. Liu, L62 (HKAS
102151, holotype).
Notes: Occultibambusa kunmingensis is morphologically
similar to O. jonesii in having cylindrical to clavate asci, fusiform, curved, brown, guttulate ascospores without any mucilaginous sheaths and appendages (Zhang et al. 2017b). However,
O. kunmingensis has longer asci (110–140(–160) × 13–16.5 μm
vs. (65–)75–89(–105) × 13.5–19 μm) and ascospores (32–40 ×
5–6.5 μm vs. 27–33.5 × 5.5–6.5 μm). Occultibambusa kunmingensis has a prominent, black, short neck, which is absent in O.
jonesii (Zhang et al. 2017b). Phylogenetic analysis supports our
collection to be a new species (Fig. 89).
60–80 μm high, 25–70 μm wide, obvious short cylindrical,
central ostiolate, periphysate. Peridium 25–90 μm thick,
composed of several layers of brown to black, thin-walled
cells of textura angularis, flattened and thin at the base, with
palisade-shaped cells near the base. Pseudoparaphyses 2–3
μm wide, numerous, cellular, hypha-like, hyaline, septate.
Asci 8-spored, bitunicate, mostly broadly clavate or sometimes narrowly clavate, (60–)78–125 × 12.5–15.5 μm (x̄ = 90
× 14 μm, n = 15), apically rounded, with an ocular chamber,
2.8 μm high × 3.8 μm wide, short or long pedicellate and up
to 37 μm long when mature. Ascospores 22–29 × 6–8 μm (x̄
= 27 × 7.5 μm, n = 20), uni- to bi-seriate, fusiform, mostly
slightly curved towards different directions at each end, pale
brown and 1-septate when young, brown, 3-septate and deeply
pigmented at septa when mature, becoming dark brown in
central two cells and hyaline to pale brown in bipolar cells,
constricted at the septa, guttulate, thin-walled, smooth, without sheath. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 15 mm in 25 days at 25 °C, dark grey from above, black
from below, surface rough, with dense mycelium, dry,
raised, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H1B-1 (MFLU 18-1208), living culture MFLUCC
18-1028; ibid., H1B-2 (HKAS 101724), living culture
KUMCC 18-0074.
Notes: Our collection MFLUCC 18-1028 clusters with
Occultibambusa pustula MFLUCC 18-1028 with high bootstrap support (Fig. 89). Occultibambusa pustula only has
LSU and ITS sequence data in GenBank. They have identical
LSU sequence data and three nucleotide differences in ITS
sequence data between MFLUCC 18-1028 and MFLUCC
11-0502, which indicate them to be the same species.
*Occultibambusa pustula D.Q. Dai & K.D. Hyde, Fungal
Diversity 82: 30 (2016); Fig. 87
Freshwater distribution: China (this study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 190–220 μm high, 320–350 μm
diam., black, scattered, immersed with proliferating papilla
erumpent through host surface, ellipsoidal, ostiolate. Papilla
13
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Fig. 86 Occultibambusa kunmingensis (HKAS 102151, holotype). a–d Ascomata on host substrate. e Vertical section of ascoma.
f Structure of peridium. g Pseudoparaphyses. h–k Bitunicate asci. l
473
Ocular chamber. m–p Ascospores. q Germinated ascospore. r, s Colony on PDA (left-front, right-reverse). Scale bars: e = 50 μm, f, h–k,
m–q = 20 μm, g, l = 10 μm
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Fungal Diversity (2020) 105:319–575
Fig. 87 Occultibambusa pustula (MFLU 18-1208, new habitat and geographical record).
a, b Ascomata erumpent
through host substrate. c, d
Vertical section of ascoma. e
Structure of peridium showing
the palisade-shaped cells near
the base. f Structure of lateral
peridium. g Asci embedded
in pseudoparaphyses. h–k
Bitunicate asci. l–q Ascospores.
r, s Colony on PDA (up-front,
down-reverse). Scale bars: d =
100 μm, e–i, k = 20 μm, j, l–q
= 10 μm
Occultibambusa pustula was collected from dead culm of
bamboo in Thailand and characterized by immersed ascomata
with central ostiole, cylindrical asci, and fusiform, 1-septate,
hyaline to pale brown ascospores (Dai et al. 2017). Our collection has very similar morphological characters with O.
pustula and they have overlapping ascus and ascospore size.
13
The holotype of O. pustula studied by Dai et al. (2017) was
immature as shown in their photo plate, and thus we describe
more details for this species from our mature collection
MFLUCC 18-1028. In our collection, the ascospores are
initially pale brown and 1-septate, which are similar to the
holotype. At maturity, ascospores become brown, 3-septate,
Fungal Diversity (2020) 105:319–575
deeply pigmented at septa, and with two dark brown central
two cells and hyaline to pale brown end cells. In addition, we
observed prominent cylindrical papilla and palisade-shaped
cells near the ascomatal base, which were not described in the
holotype. Based on phylogenetic analysis (Fig. 89) and morphological characters, we identify our collection MFLUCC
18-1028 as O. pustula. This is a new habitat and geographical
record for O. pustula from freshwater in China.
Key to freshwater Occultibambusa species
1. Ascospores 32–40 × 5–6.5 μm………O. kunmingensis
1. Ascospores < 32 μm long………………………………2
2. Ascospores have two dark brown central cells and hyaline to pale brown end cells when mature……O. pustula
2. Ascospores are evenly coloured when mature…………
……………………………………………O. aquatica
Seriascoma Phook., D.Q. Dai & K.D. Hyde, Fungal Diversity 82: 30 (2016)
Saprobic on decaying or submerged bamboo culms.
Sexual morph: Ascostromata gregarious, immersed beneath
clypeus, coriaceous, uni- to multi-loculate, ostiolate, with
slit-like opening. Locules arranged in rows, immersed,
subglobose to ampulliform, or quadrilateral, with ostiolate
papilla. Peridium thick-walled, composed of several layers
of small, dark brown, pseudoparenchymatous cells of textura
angularis. Pseudoparaphyses numerous, cellular, hyphalike, broadly filamentous, hyaline, septate. Asci 8-spored,
bitunicate, fissitunicate, clavate, long pedicellate, apically
rounded, with a well-developed, ocular chamber. Ascospores
uni- to tri-seriate, clavate to fusiform, hyaline, septate (Dai
et al. 2017). Asexual morph: Coelomycetous. Conidiomata
eustromatic, solitary to gregarious, immersed, conical, uniloculate, black. Peridium comprising host and fungal tissue,
with several layers of dark brown to black, pseudoparenchymatous cells of textura angularis. Conidiophores Conidiogenous cells reduced. phialidic, determinate, discrete,
cylindrical to ampulliform or lageniform, hyaline. Conidia
oblong, hyaline, aseptate (Dai et al. 2017).
Type species: Seriascoma didymosporum Phook., D.Q.
Dai, Karun. & K.D. Hyde
Notes: The monotypic genus Seriascoma, typified by S.
didymosporum, was introduced by Dai et al. (2017). The
asexual morph was produced on WA with oblong, hyaline,
aseptate conidia, which are similar to Occultibambusa fusispora (Dai et al. 2017). However, the sexual morph of Seriascoma is characterized by elongate, multiloculate ascostromata with a slit-like opening, which are different to the
uniloculate ascostromata with a central papilla of Occultibambusa. Seriascoma formed a distinct clade and nested in
Occultibambusaceae with high bootstrap support (Dai et al.
475
2017; Tibpromma et al. 2018). We re-collected the type species S. didymosporum from submerged wood in China.
List of freshwater Seriascoma species
*Seriascoma didymosporum Phook., D.Q. Dai, Karun. &
K.D. Hyde [as ‘didymospora’], Fungal Diversity 82: 32
(2016); Fig. 88
Freshwater distribution: China (this study)
Saprobic on submerged wood. Sexual morph: Ascomata
120–170 μm high, 200–250 μm diam., gregarious, immersed
to erumpent in linear rows, raised, coriaceous, uniloculate,
ostiolate. Peridium 30–50 μm wide, unequal in thickness,
composed of several layers of large, dark brown pseudoparenchymatous cells of textura angularis. Pseudoparaphyses 2.5–4 μm wide, numerous, cellular, hypha-like, broadly
filamentous, hyaline, septate. Asci 70–95 × 9–11 μm ( x̄ =
83 × 9.5 μm, n = 10), 8-spored, bitunicate, fissitunicate,
clavate, long pedicellate, apically rounded, with an ocular
chamber. Ascospores 10.5–14.5 × 3.5–5 μm ( x̄ = 12.5 ×
4 μm, n = 20), uni- to tri-seriate, clavate to fusiform, with
rounded ends, 1-septate, constricted at the septum, asymmetrical, upper cell shorter and wider than lower cell, hyaline, straight or curved, smooth-walled. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 15 days at 25 °C, white to pale brown from
above, black from below, surface rough, with dense mycelium, dry, raised, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H158C-1 (MFLU 18-1185), living culture MFLUCC
18-1026; ibid., H158C-2 (HKAS 101720), living culture
KUMCC 18-0070.
Notes: Our new collection MFLUCC 18-1026 clusters
with Seriascoma didymosporum with high bootstrap support
(Fig. 89). MFLUCC 18-1026 is identified as S. didymosporum based on sequence data which has one, four (includes
two gaps) and three nucleotide differences in LSU, ITS
and TEF sequence data, respectively, between MFLUCC
18-1026 and MFLUCC 11-0179 (ex-holotype). We only
found uniloculate ascomata in our collection, while the holotype MFLU 11–0215 has multi-loculate ascomata (Dai et al.
2017). Our collection MFLUCC 18-1026 has longer asci
(70–95 × 9–11 μm vs. (56–)60–75(−80) × 8–11(−13) μm)
than the holotype (Dai et al. 2017). This is a new habitat and
geographical record for S. didymosporum from freshwater
in China.
Parabambusicolaceae Kaz. Tanaka & K. Hiray., Stud.
Mycol. 82: 115 (2015)
13
476
Fungal Diversity (2020) 105:319–575
Fig. 88 Seriascoma didymosporum (MFLU 18-1185, new
habitat and geographical
record). a Ascostromata
immersed in host substrate. b
Vertical section of ascoma. c
Structure of peridium. d–g Bitunicate asci. h Pseudoparaphyses.
i–l Ascospores. m Germinated
ascospore. n, o Colony on PDA
(up-front, down-reverse). Scale
bars: b = 50 μm, c, m = 20 μm,
d–h = 10 μm, i–l = 5 μm
Aquastroma Kaz. Tanaka & K. Hiray., Stud. Mycol. 82: 115
(2015)
Saprobic on submerged twigs of woody plant. Sexual
morph: Ascomata scattered to gregarious, immersed to
erumpent, depressed globose. Ostiolar neck central, compressed, with a wide ostiole. Peridium composed of pale
13
brown, thin-walled cells. Pseudoparaphyses septate,
branched and anastomosed. Asci 8-spored, bitunicate, fissitunicate, clavate, pedicellate. Ascospores clavate to fusiform,
slightly curved, multiseptate, slightly constricted at septa,
hyaline, smooth, with an entire sheath (Tanaka et al. 2015).
Asexual morph: Undetermined.
Fungal Diversity (2020) 105:319–575
477
Fig. 89 Phylogram generated
from maximum likelihood
analysis of combined LSU,
SSU, ITS, TEF and RPB2
sequence data for species of
Occultibambusaceae. Bootstrap
values for maximum likelihood
equal to or greater than 75% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree is
rooted to Torula herbarum CBS
111855 (Torulaceae)
Occultibambusa chiangraiensis MFLUCC 16-0380
96/0.99
Occultibambusa sp. NCYUCC 19-0370
100/0.98
Occultibambusa bambusae MFLUCC 11-0394
100/1.00
96
Occultibambusa bambusae MFLUCC 13-0855
Occultibambusa pustula MFLUCC 18-1028 F
86/1.00
100/1.00
Occltibambusa pustula MFLUCC 11-0502
Occultibambusa aquatica MFLUCC 11-0006 F
100/1.00
89/0.99
Occultibambusa
Occultibambusa jonesii GZCC 16-0117
Occultibambusa kunmingensis HKAS 102151 F
Versicolorisporium triseptatum JCM 14775
82/0.99
Versicolorisporium
Occultibambusa maolanensis GZCC 16-0116
Occultibambusa fusispora MFLUCC 11-0127
77/0.96
100/0.96
100/1.00
Occultibambusa
Seriascoma didymospora MFLUCC 11-0194
Seriascoma didymospora MFLUCC 11-0179
Seriascoma
Seriascoma didymospora MFLUCC 18-1026 F
67/0.95
Massarina rubi MUT 4323
Massarina rubi CBS 691.95
100/1.00
Brunneofusispora sinensis KUMCC 17-0030
Brunneofusispora
Massarina rubi MUT 4887
Neooccultibambusa jonesii MFLUCC 16-0643
Neooccultibambusa pandanicola KUMCC 17-0179
Neooccultibambusa
Neooccultibambusa thailandensis MFLUCC 16-0274
Torula herbarum CBS 111855
Outgroup
0.03
Type species: Aquastroma magniostiolatum Kaz. Tanaka
& K. Hiray.
Notes: The monotypic genus Aquastroma was introduced for a freshwater species A. magniostiolatum collected from Japan (Tanaka et al. 2015). Aquastroma fits
well with the familial concept of Parabambusicolaceae and
differs from other genera by its ascomata with a compressed
neck and wide ostiole, and clavate to fusiform, multiseptate ascospores with an entire sheath (Tanaka et al. 2015).
Aquastroma clustered within Parabambusicolaceae with
high bootstrap support (Tanaka et al. 2015; Phookamsak
et al. 2019). We recollected the type species A. magniostiolatum from submerged wood in Thailand.
List of freshwater Aquastroma species
*Aquastroma magniostiolata Kaz. Tanaka & K. Hiray.,
Stud. Mycol. 82: 115 (2015)
Facesoffungi number: FoF09273; Fig. 90
Freshwater distribution: Japan (Tanaka et al. 2015), Thailand (this study)
Saprobic on submerged wood in freshwater. Sexual morph:
Ascomata 190–230 μm high, 200–240 μm diam., scattered to
gregarious, immersed to erumpent, subglobose, black, coriaceous, with wide ostiole. Peridium 15–20 μm thick at sides,
composed of several layers of polygonal, thin-walled, pale
brown cells, poorly developed at the base, 5–10 μm thick.
Pseudoparaphyses 1.5–2 μm wide, numerous, cellular, hyphalike, hyaline, septate, branching and anastomosing. Asci
100–125 × 19–22 μm (x̄ = 107 × 21 μm, n = 10), 8-spored,
bitunicate, fissitunicate, clavate, sessile or short pedicellate.
Ascospores 32–39 × 8–11 μm (x̄ = 36 × 9.5 μm, n = 20), bi- to
tri-seriate, clavate to fusiform, mostly curved, 5–8-septate, constricted at the septa, hyaline, become brown in old specimen,
smooth, with an entire sheath, 5–9 μm wide. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 50 mm in 60 days at 25 °C, grey to black from above,
black from below, surface rough, dry, raised, with dense
mycelium, edge entire.
Material examined: THAILAND, Chiang Mai Province,
on submerged wood in a stream, 9 February 2018, X.D. Yu,
Y2-1 (MFLU 18-0988), living culture MFLUCC 18-0976;
ibid., Y2-2 (HKAS 105097).
Notes: Our collection MFLUCC 18-0976 is identified as
Aquastroma magniostiolata based on phylogenetic analysis
(Fig. 2). MFLUCC 18-0976 has identical morphological
characters with the holotype of A. magniostiolata, except for
slightly thinner asci (100–125 × 19–22 μm vs. 112.5–137.5
× 25–29.5 μm) (Tanaka et al. 2015). This is a new geographical record for A. magniostiolatum in Thailand.
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Fungal Diversity (2020) 105:319–575
Fig. 90 Aquastroma magniostiolata (MFLU 18-0988, new geographical record). a Appearance of ascomata immersed in host
substrate. b Vertical section of ascoma. c Structure of peridium. d
Pseudoparaphyses. e–h Bitunicate asci. i–l Ascospores. (l ascospore
in Indian Ink). m Germinated ascospore. n Colony on PDA (from
above). Scale bars: b = 50 μm, c–h = 20 μm, i–l = 10 μm, m = 30
μm
Parabambusicola Kaz. Tanaka & K. Hiray., Stud. Mycol.
82: 115 (2015)
Saprobic on dead stems or submerged wood. Sexual
morph: Ascomata scattered or clustered in group, immersed
to erumpent, hemispherical with flattened base, ostiolate,
with short, central papilla. Peridium composed of pale
brown, flattened cells at the sides and parallel rows of rectangular to polygonal cells at the rim. Pseudoparaphyses
numerous, cellular, hypha-like, hyaline, septate. Asci
8-spored, bitunicate, broadly cylindrical to clavate, short
pedicellate. Ascospores triseriate, fusiform, multiseptate,
hyaline, smooth, with an entire sheath (Tanaka et al. 2015).
Asexual morph: Undetermined.
Type species: Parabambusicola bambusina (Teng) Kaz.
Tanaka & K. Hiray.
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Fungal Diversity (2020) 105:319–575
Notes: Parabambusicola was established to accommodate
a known species Massarina bambusina Teng (Teng 1934;
Tanaka and Harada 2003c). Both P. bambusina and P. thysanolaenae Goonas. et al. were isolated from dead stems in
terrestrial habitats (Teng 1934; Tanaka and Harada 2003c),
while our new species P. aquatica was found on submerged
wood in freshwater habitats.
List of freshwater Parabambusicola species
*Parabambusicola aquatica W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557931; Facesoffungi number: FoF09274; Fig. 91
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-1524
Saprobic on submerged wood in freshwater. Sexual
morph: Ascomata 150–180 μm high, 350–400 μm wide,
scattered, immersed, with black spot on host surface, flat
cone, brown, coriaceous. Peridium 10–25 μm thick at the
sides, comprising several layers of brown, thin-walled, large
cells of textura angularis, flattened and poorly developed
at the base, 10–15 μm thick. Pseudoparaphyses 1.5–2 μm
diam., numerous, cellular, hypha-like, hyaline, indistinctly
septate, embedded in a gelatinous matrix. Asci 110–175
× 30–37 μm ( x̄ = 147 × 34 μm, n = 10), 8-spored, bitunicate, broadly clavate or narrowly ellipsoidal, apically
rounded with well-developed ocular chamber. Ascospores
55–60(–67) × 12–17 μm (x̄ = 56 × 14 μm, n = 10), overlapping bi- to tri-seriate, straight or slightly curved, hyaline,
5-septate, clearly shown when old, constricted at the septa,
fusiform with rounded ends, with minute guttules thinwalled, smooth, with a globose mucilaginous sheath, 30–50
μm wide. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 5 days at 25 °C, grey to dark grey from above,
brown from below, surface rough, dry, raised, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
20180503-1 (MFLU 18-1524, holotype), ex-type living culture MFLUCC 18-1140; ibid., 20180503-2 (HKAS 104993,
isotype), ex-type living culture KUMCC 19-0010.
Notes: Parabambusicola aquatica is similar to P. bambusina and P. thysanolaenae in the characters of peridium,
asci and ascospores (Tanaka and Harada 2003c; Phookamsak
et al. 2019). However, P. aquatica has wider asci (110–175
× 30–37 μm vs. 86–150 × 22–30 μm) and ascospores
(55–60(–67) × 12–17 μm vs. 42–64.5 × 7–10 μm) than P.
bambusina (Tanaka and Harada 2003c). Parabambusicola
aquatica also has larger ascospores than P. thysanolaenae
(55–60(–67) × 12–17 μm vs. 45–55 μm × 7.5–11 μm). They
are well separated in our phylogenetic tree (Fig. 2).
479
Paramonodictys N.G. Liu, K.D. Hyde & J.K. Liu, Fungal
Diversity 100: 90 (2020)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Colonies on natural substrate superficial,
scattered, black. Mycelium partly immersed, partly superficial. Stroma present, erumpent, erect, subcylindrical or
truncated-cone-form, pale brown to olivaceous brown.
Conidiophores absent. Conidiogenous cells holoblastic,
monoblastic. Conidia dictyosporous, muriform, globose or
subglobose, broadly rounded at the apex, olivaceous brown
to dark brown, solitary, smooth-walled (Hyde et al. 2020b).
Type species: Paramonodictys solitarius N.G. Liu, K.D.
Hyde & J.K. Liu
Notes: Paramonodictys was introduced to accommodate
a monodictys-like species collected on decaying wood in
China (Hyde et al. 2020b). Monodictys S. Hughes is polyphyletic (Mouzouras and Jones 1985; Han et al. 2014; Tanaka et al. 2015) and typified by M. putredinis (Wallr.) S.
Hughes. Due to the dearth of sequences of M. putredinis,
Monodictys sensu stricto could not be clarified. Based on the
presence of stroma, Paramonodictys was established (Hyde
et al. 2020b).
List of freshwater Paramonodictys species
*Paramonodictys solitarius N.G. Liu, K.D. Hyde & J.K.
Liu, Fungal Diversity 100: 91 (2020); Fig. 92
Freshwater distribution: Thailand (this study)
Saprobic on submerged wood in freshwater. Sexual
morph: Undetermined. Asexual morph: Colonies superficial, scattered, black. Mycelium mostly superficial, composed of branched, brown, septate, rough hyphae. Stroma
not observed. Conidiophores absent. Conidiogenous cells
holoblastic, monoblastic. Conidia 30–55 × 25–50 μm ( x̄
= 40 × 37 μm, n = 15), solitary, acrogenous, simple, dry,
globose or subglobose, ellipsoidal, muriform, multiseptate,
dark brown to black, easily breaking when in water, smooth,
thin-walled.
Culture characteristics: On PDA, colony circular, reaching 15 mm in 20 days at 25 °C, grey from above, black from
below, surface rough, with dense mycelium, dry, raised, edge
entire.
Material examined: THAILAND, Nan Province, on
submerged wood in a stream, 4 August 2017, S. Boonmee,
DP10 (MFLU 17-1714), living culture MFLUCC 17-2353.
Notes: Our new collection MFLUCC 17-2353 has
identical LSU sequence data with the ex-type strain of
Paramonodictys solitarius, with two and one nucleotide
differences in ITS and TEF sequence data, respectively.
Phylogenetic analysis also supports our collection as P.
solitarius (Fig. 2). Our collection has smaller conidia
(30–55 × 25–50 μm vs. 50–87 × 40–61 μm) than the holotype (Hyde et al. 2020b), and lacks a stroma. This is a
13
480
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Fungal Diversity (2020) 105:319–575
◂Fig. 91 Parabambusicola aquatica (MFLU 18-1524, holotype). a–c
Appearance of ascomata immersed in host substrate. d Vertical section of ascoma. e Structure of peridium. f Ascus apex with an ocular
chamber. g–j Bitunicate asci. k Pseudoparaphyses. l–p Ascospores.
q Ascospore in Indian Ink. r Germinated ascospore. s, t Colony on
PDA (up-front, down-reverse). Scale bars: d = 100 μm, e, f, k = 10
μm, g–j, l–p = 20 μm, q, r = 50 μm
new habitat and geographical record for P. solitarius from
freshwater in Thailand.
Periconiaceae Nann., Repert. mic. uomo: 482 (1934)
Periconia Tode, Fung. mecklenb. sel. (Lüneburg) 2: 2
(1791)
Saprobic on decaying wood from terrestrial, freshwater,
mangrove and marine habitats. Sexual morph: Ascomata
scattered to gregarious, immersed to erumpent, globose,
with ostiolate papilla. Ostiolar neck central, periphysate.
Peridium composed of several layers of pale brown to brown,
thin-walled cells. Pseudoparaphyses numerous, cellular,
hyaline, septate, branched, anastomosing. Asci bitunicate,
fissitunicate, 8-spored, oblong to cylindrical. Ascospores
broadly fusiform, 1-septate, hyaline, smooth, with an entire
sheath (Tanaka et al. 2015). Asexual morph: Hyphomycetous. periconia- or noosia-like. Conidiophores macronematous, mononematous, sometimes absent, branched. Branches
pale brown to brown, smooth to slightly echinulate. Conidial
heads spherical. Conidiogenous cells mono- to poly-blastic,
discrete on stipe and branches. Conidia variable in shape,
globose to ellipsoidal, aseptate, solitary or catenate, brown,
verruculose to echinulate (Tanaka et al. 2015; Hyde et al.
2017).
Type species: Periconia lichenoides Tode
Notes: Periconia is characterized by mostly macronematous conidiophores which often have a spherical head,
monoblastic or polyblastic conidiogenous cells, and catenate, aseptate, verruculose or echinulate conidia (Tode
1791; Cai et al. 2006a). After re-examination of six collections of P. byssoides Pers., it was deemed conspecific with
P. lichenoides, but without molecular evidence (Mason and
Ellis 1953). Periconia species are widely distributed and
have been reported as endophytes, plant pathogenic (Odvody
et al. 1977; Romero et al. 2001) and saprobic fungi from
terrestrial (Liu et al. 2017b; Jayasiri et al. 2019), mangrove
(Alias and Jones 2000), marine (Kohlmeyer 1977) and freshwater habitats (Luo et al. 2004; Hyde et al. 2017).
Two species have been linked to sexual morphs based on
cultural and phylogenetic methods, viz. Periconia igniaria
E.W. Mason & M.B. Ellis (Massarina in Dothideomycetes)
(Booth 1968; Aptroot 1998) and P. prolifica Anastasiou
(Okeanomyces in Sordariomycetes) (Kohlmeyer 1969;
Pang et al. 2004). Periconia is, therefore, recognized as a
polyphyletic genus (Tanaka et al. 2015; Liu et al. 2017b).
481
The phylogenetic analysis of Phookamsak et al. (2019)
showed that some genera, viz. Bambusistroma D.Q. Dai
& K.D. Hyde, Flavomyces D.G. Knapp et al. and Noosia
Crous et al., nested within Periconia in Periconiaceae, which
also indicated the polyphyletic nature of Periconia. The
sequences of the type species P. lichenoides are unavailable,
but Periconia is placed in Dothideomycetes based on some
species, e.g. P. aquatica Z.L. Luo et al., P. byssoides Pers.
and P. submersa Z.L. Luo et al, which have the typical characters of Periconia (Tanaka et al. 2015; Hyde et al. 2017).
Tanaka et al. (2015) introduced a freshwater sexual species P. pseudodigitata Kaz. Tanaka & K. Hiray., which
formed a sister clade to P. digitata (Cooke) Sacc. The freshwater species P. prolifica was suggested to be excluded from
Periconia based on the distinct molecular characters and
basipetal production of conidial chains, contrasting with the
acropetal conidial chains of Periconia sensu stricto (Markovskaja and Kacergius 2014; Tanaka et al. 2015). We exclude
this species in the key below.
List of freshwater Periconia species
*Periconia aquatica Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Diversity 87: 71 (2017); Fig. 93b, f–i
Freshwater distribution: China (Hyde et al. 2017)
*Periconia byssoides Pers., Syn. meth. fung. (Göttingen)
1: 18 (1801)
Synonymy: Sporocybe byssoides (Pers.) Fr., Syst. mycol.
(Lundae) 3: 343 (1832)
Periconia pycnospora Fresen., Beitr. Mykol. 1: 20 (1850)
Periconia pycnospora f. citri Penz., Michelia 2: 469
(1882)
Cephalotrichum byssoides (Pers.) Kuntze, Revis. gen. pl.
(Leipzig) 3: 453 (1898)
Periconia pycnospora f. israelitica Rayss & Borut, Mycopath. Mycol. appl. 10: 167 (1958)
Freshwater distribution: USA (Shearer 1972), China (Luo
et al. 2004)
*Periconia digitata (Cooke) Sacc., Syll. fung. (Abellini) 4:
274 (1886)
Basionym: Sporocybe digitata Cooke, Grevillea 12: 33
(1883)
Freshwater distribution: China (Luo et al. 2004)
*Periconia minutissima Corda, Icon. fung. (Prague) 1: 19
(1837)
Freshwater distribution: China (Cai et al. 2002a; Luo
et al. 2004; Hyde et al. 2017)
*Periconia pseudodigitata Kaz. Tanaka & K. Hiray., Stud.
Mycol. 82: 119 (2015)
Freshwater distribution: Japan (Tanaka et al. 2015)
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Fungal Diversity (2020) 105:319–575
Fig. 92 Paramonodictys
solitarius (MFLU 17-1714,
new habitat and geographical
record). a Colonies on natural
substrate. b Appearance of
conidial wall. c–f Conidia. g,
h Colony on PDA (left-front,
right-reverse). Scale bars: b–f
= 20 μm
*Periconia submersa Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Diversity 87: 73 (2017); Fig. 93a, c–e, j, k
Freshwater distribution: China (Hyde et al. 2017)
Key to freshwater Periconia species
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
Conidiophores branched………………………………2
Conidiophores unbranched……………………………4
Conidia solitary………………………………P. digitata
Conidia catenate………………………………………3
Conidia 7–9.5 μm diam.………………P. pseudodigitata
Conidia 4.5–5.5 μm diam.………………P. minutissima
Conidia spherical…………………………P. byssoides
Conidia non-spherical…………………………………5
Conidia 10–12 × 6–7 μm……………………P. aquatica
Conidia 6.5–9.5 × 4.5–5.5 μm……………P. submersa
Phaeoseptaceae S. Boonmee, Thambugala & K.D. Hyde,
Mycosphere 9(2): 323 (2018)
Key to freshwater genera of Phaeoseptaceae
1. Sexual morph……………………………Phaeoseptum
13
1. Asexual morph……………………………Pleopunctum
Phaeoseptum Ying Zhang, J. Fourn. & K.D. Hyde, Mycologia 105(3): 606 (2013)
Saprobic on submerged branch or dead stems. Sexual
morph: Ascomata solitary, scattered to gregarious, sometimes immersed under pseudoclypeus, erumpent when
mature, appearing as black, elongate regions on host surface, depressed globose, sometimes covered with some
brown hyphae, dark brown, with ostiolate papilla. Peridium
pseudoparenchymatous, comprising several layers of somewhat flattened cells of textura angularis. Pseudoparaphyses numerous, cellular, narrow, hyaline, cylindrical, septate,
anastomosing. Asci 8-spored, bitunicate, cylindrical-clavate,
pedicellate, with a small ocular chamber and apical ring.
Ascospores uni- to tri-seriate, fusiform, slightly curved,
transversally septate, with a vertical septum in nearly all
cells, sometimes with a Y-shaped septum in the end cell, pale
brown to brown, with thickened and darkened septa at maturity, smooth- or rough-walled, without sheath or appendages
(Zhang et al. 2013b). Asexual morph: Undetermined.
Type species: Phaeoseptum aquaticum Ying Zhang, J.
Fourn. & K.D. Hyde
Fungal Diversity (2020) 105:319–575
483
Fig. 93 Periconia spp.
(Material examined: CHINA,
Yunnan Province, saprobic on
submerged decaying wood in
Nujiang River, 3 May 2015,
X.C. Tao, N7–4–1, HKAS
92754 holotype; ibid., X.J.
Su, N4–8–1, HKAS 92738). a,
c–e, j, k Periconia submersa
(HKAS 92738). b, f–i Periconia aquatica (HKAS 92754).
a, b Colonies on the host. c
Conidiophore with conidia. d,
e Conidiogenous cells bearing
conidia. f–k Conidia. Scale
bars: c = 100 μm, d, e = 20 μm,
f–k = 5 μm
Notes: Phaeoseptum was introduced by Zhang et al.
(2013b) to accommodate the freshwater species P. aquaticum collected from France. Phaeoseptum was placed in
Halotthiaceae based on LSU sequence data (Zhang et al.
2013b). With more sequence data added, Phaeoseptum was
transferred to Phaeoseptaceae (Hyde et al. 2018a). The type
species P. aquaticum is characterized by ascomata fully
immersed under a small blackened pseudoclypeus, cylindrical-clavate asci with a small ocular chamber and an apical
ring, and fusiform ascospores with transverse and longitudinal septa, exceptionally with a Y-shaped septum in the end
cell (Zhang et al. 2013b).
List of freshwater Phaeoseptum species
*Phaeoseptum aquaticum Ying Zhang, J. Fourn. & K.D.
Hyde, Mycologia 105(3): 606 (2013); Fig. 94
Freshwater distribution: France (Zhang et al. 2013b)
Pleopunctum N.G. Liu, K.D. Hyde & J.K. Liu, Mycosphere
10 (1): 767 (2019)
Saprobic on decaying wood in terrestrial or submerged
wood in freshwater habitats. Sexual morph: Undetermined.
Asexual morph: Hyphomycetous. Colonies clustered in several small groups or gregarious, sporodochial, punctiform,
brown to black, shining. Mycelium immersed in the substratum, composed of septate, branched, subhyaline to greyish
brown hyphae. Conidiophores macronematous, mononematous, cylindrical, branched or unbranched, septate, medium
13
484
Fig. 94 Phaeoseptum aquaticum (redrawn from Zhang et al.
(2013b), IFRD 8986, holotype). a Habitat section of an ascoma
showing the clypeus. b Structure of lateral peridium. c Apex of
immature ascus showing the thickened apex and minute subapical
ring. d Mature ascospore. e Bitunicate asci. Scale bars: a = 80 μm, b
= 20 μm, c = 10 μm, d = 10 μm, e = 20 μm
brown, smooth-walled, thick-walled. Conidiogenous cells
holoblastic, monoblastic, cylindrical, brown. Conidia acrogenous, solitary, muriform, constricted at septa, broadly oval
to ellipsoidal, variable in shape, smooth-walled, pale brown
to dark brown, often with a hyaline, ellipsoidal to globose
basal cell (Liu et al. 2019).
Type species: Pleopunctum ellipsoideum N.G. Liu, K.D.
Hyde & J.K. Liu
Notes: Pleopunctum was introduced to accommodate two
terrestrial species, P. ellipsoideum N.G. Liu et al. and P.
pseudoellipsoideum N.G. Liu et al., collected from decaying
wood in China (Liu et al. 2019). Pleopunctum is characterized by muriform, oval to ellipsoidal conidia often with a
hyaline, elliptical to globose basal cell (Liu et al. 2019). Pleopunctum, the only hyphomycetous genus, formed a wellsupported and distinct clade in Phaeoseptaceae (Liu et al.
2019). We collected P. pseudoellipsoideum from submerged
wood in China.
List of freshwater Pleopunctum species
*Pleopunctum pseudoellipsoideum N.G. Liu, K.D. Hyde &
J.K. Liu, Mycosphere 10 (1): 768 (2019); Fig. 95
Freshwater distribution: China (this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies clustered in several small
groups, sporodochial, punctiform, black, shining. Mycelium
partly immersed in natural substratum, composed of hyaline
13
Fungal Diversity (2020) 105:319–575
to pale brown, septate hyphae. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells up to 19.5 µm
long, 3.5 µm wide, holoblastic, monoblastic, integrated,
determinate, cylindrical, easily broken off, pale brown to
brown, smooth-walled, thick-walled. Conidia 35.5–66 ×
26.5–40 µm (subtending cells included) (x̄ = 51.5 × 30 µm,
n = 15), solitary, dry, ellipsoidal, subglobose, subcylindrical
or obovoid, rounded at the apex, multi-septate, muriform,
slightly constricted at the septa, brown to black, paler to
hyaline at the base; basal cells (subtending cells) subhyaline
to hyaline, cuneiform, always swollen to globose, up to 15
µm diam., straight, smooth, thin-walled.
Material examined: CHINA, Yunnan Province, Qujing,
on submerged wood in a stream, 10 May 2017, C.X. Liu,
L43 (HKAS 102147).
Notes: HKAS 102147 clusters with Pleopunctum pseudoellipsoideum with high bootstrap support (Fig. 2). HKAS
102147 has identical LSU and ITS sequence data with
the ex-type strain of P. pseudoellipsoideum (MFLUCC
19-0391) and seven nucleotide differences in TEF sequence
data, which indicate they are conspecific. Our collection
has wider conidia than the holotype (35.5–66 × 26.5–40
µm vs. 39–59 × 19–28 μm) (Liu et al. 2019). This is a new
habitat record for P. pseudoellipsoideum from freshwater.
We directly sequenced from conidia because the culture was
contaminated.
Pleomassariaceae M.E. Barr, Mycologia 71(5): 949 (1979)
Splanchnonema Corda, 3 Abt. (Pilze Deutschl.) 2: 115
(1829)
Saprobic on wood or lichens in terrestrial or submerged
wood in freshwater habitats. Sexual morph: Ascomata
medium to large, solitary or scattered, immersed in a pseudostroma or superficial, uniloculate, globose to flattened
subglobose, lobate to generally reniform, dark brown to
black, with a small ostiole appearing on the host surface,
sometimes carbonized at the whole upper. Peridium thin.
Pseudoparaphyses dense, cellular, hyaline, septate, anastomosing and branching, embedded in a matrix. Asci 8-spored,
bitunicate, fissitunicate, clavate to broadly cylindrical, short
pedicellate. Ascospores uni- to bi-seriate, clavate with a
rounded apex and acute base, or obovoid, reddish brown,
septate, constricted at the septa, usually asymmetrical, thinwalled, with or without sheath (Zhang et al. 2012b; Aptroot
and Moon 2014). Asexual morph: Myxocyclus Riess, Stegonsporium Corda (Barr 1982; Sivanesan 1984), Macrodiplodiopsis Petr. (Glawe 1985) and Helminthosporium Link
(Subramanian and Sekar 1987).
Type species: Splanchnonema pustulatum Corda
Notes: Splanchnonema, an old genus, comprises over
40 epithets in Index Fungorum (2020). Pleomassaria was
treated as a synonym of Splanchnonema (Barr 1993), but not
Fungal Diversity (2020) 105:319–575
485
Fig. 95 Pleopunctum pseudoellipsoideum (HKAS 102147,
new habitat record). a, b
Colonies on submerged wood.
c–f Conidiogenous cells with
conidia. g, h Conidia. Scale
bars: c–h = 20 μm
accepted by some authors (Zhang et al. 2012b). Pleomassaria was suppressed and replaced by Prosthemium Kunze
(Wijayawardene et al. 2014) based on the phylogenetic
results where the type species of both genera (P. siparia
(Berk. & Broome) Sacc. and P. betulinum Kunze) grouped
together in a monophyletic clade (Tanaka et al. 2010).
Splanchnonema was accepted as a member of Pleomassariaceae based on phylogeny (Hyde et al. 2013; Liu et al. 2015)
and followed by Wijayawardene et al. (2018, 2020). The
asexual morphs of this genus are generally coelomycetous
although some are hyphomycetous (Subramanian and Sekar
1987), but not shown by molecular data.
The freshwater species Splanchnonema britzelmayrianum
(Rehm) Boise is characterized by spherical ascomata with
a short papilla, cylindric-clavate asci, and obovoid, brown,
7–8-septate, minutely punctate ascospores (Boise 1985). The
phylogenetic placement of S. britzelmayrianum in Splanchnonema needs to be confirmed with molecular data.
List of freshwater Splanchnonema species
Splanchnonema britzelmayrianum (Rehm) Boise [as ‘britzelmayriana’], Mycotaxon 22(2): 480 (1985)
Basionym: Melanomma megalosporum var. britzelmayrianum Rehm, Hedwigia 20(3): 51 (1881)
Synonymy: Melanomma britzelmayrianum (Rehm) Sacc.,
Syll. fung. (Abellini) 2: 120 (1883)
Trematosphaeria britzelmayriana (Rehm) Sacc., Syll.
fung. (Abellini) 2: 120 (1883)
Freshwater distribution: Hungary (Révay and Gönczöl
1990)
Pleomonodictydaceae Hern.-Restr., J. Mena & Gené, Stud.
Mycol. 86: 76 (2017)
Pleomonodictys Hern.-Restr., J. Mena & Gené, Stud. Mycol.
86: 77 (2017)
Saprobic on submerged wood or dead bark in freshwater or terrestrial habitats. Sexual morph: Undetermined.
Asexual morph: Colonies effuse, black. Mycelium partly
immersed or partly superficial, composed of branched,
septate, nodulose, smooth or verruculose hyphae. Conidiophores micronematous or semi-macronematous, often
reduced to conidiogenous cells, terminal or intercalary.
Conidiogenous cells blastic. Conidia muriform, variable
in shape, dark brown to black, solitary or in short chains,
verrucose or tuberculate (Hernández-Restrepo et al. 2017).
Type species: Pleomonodictys descalsii Hern.-Restr. et al.
13
486
Notes: Pleomonodictys was introduced to accommodate
two monodictys-like species which differ from Monodictys
S. Hughes by verrucose to tuberculate conidia and/or hyphae
(Cai et al. 2006a; Hernández-Restrepo et al. 2017). The type
species of Monodictys, M. putredinis (Wallr.) S. Hughes,
lacks sequence data and was reported as the asexual morph
of Ohleria brasiliensis Starbäck based on culture methods
(Samuels 1980); molecular data are needed to clarify the
placement of Monodictys. Pleomonodictys capensis (R.C.
Sinclair, Boshoff & Eicker) Hern.-Restr. et al. was found
from freshwater habitats, but not confirmed by molecular
data (Ho et al. 2001).
List of freshwater Pleomonodictys species
*Pleomonodictys capensis (R.C. Sinclair, Boshoff & Eicker)
Hern.-Restr., J. Mena & Gené, Stud. Mycol. 86: 77 (2017)
Basionym: Monodictys capensis R.C. Sinclair, Boshoff &
Eicker, Mycotaxon 59: 359 (1996)
Freshwater distribution: Brunei (Ho et al. 2001)
Pseudoastrosphaeriellaceae Phook. & K.D. Hyde, Fungal
Diversity 74: 181 (2015)
Pseudoastrosphaeriella Phook., Z.L. Luo & K.D. Hyde,
Fungal Diversity 74: 182 (2015)
Saprobic on bamboo and palms in freshwater or terrestrial habitats. Sexual morph: Ascostromata dark brown,
scattered to gregarious, immersed to erumpent, slightly
conical to lenticular or hemisphaerical, with flattened base,
uni- to bi-loculate, glabrous, coriaceous, with ostiolate
papilla. Ostioles central, cylindrical, brittle, carbonaceous,
with short or long neck. Peridium thick-walled, unequal
in thickness, composed of pseudoparenchymatous cells,
outer layers mostly comprising host cells and fungal tissue,
poorly developed at the base. Pseudoparaphyses numerous,
trabeculate, filiform, hyaline, branching and anastomosing,
embedded in a hyaline gelatinous matrix. Asci 8-spored,
bitunicate, clavate to cylindric-clavate, pedicellate, with
ocular chamber. Ascospores overlapping uni- to tri-seriate, brown to reddish brown, fusiform to clavate, septate,
rough-walled or smooth-walled, with or without striations,
or with longitudinal ridges, with or without a mucilaginous
sheath (Phookamsak et al. 2015b). Asexual morph: Coelomycetous. Conidiomata pycnidial, immersed, solitary to
scattered, conical or hemisphaerical to globose, ostiolate.
Peridium thick-walled, unequal in thickness, composed of
dark, pseudoparenchymatous cells. Conidiophores reduced
to conidiogenous cells. Conidiogenous cells phialidic, discrete, cylindrical or cylindric-clavate or ampulliform, septate, hyaline, smooth-walled. Conidia globose to subglobose,
or oblong, aseptate, hyaline, smooth-walled (Phookamsak
et al. 2015b).
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Fungal Diversity (2020) 105:319–575
Type species: Pseudoastrosphaeriella thailandensis
Phook., Z.L. Luo & K.D. Hyde
Notes: Pseudoastrosphaeriella was introduced to accommodate astrosphaeriella-like species (Phookamsak et al.
2015b). Pseudoastrosphaeriella is distinguished from Astrosphaeriella Syd. & P. Syd. by its immersed ascostromata
beneath the host epidermis, with short or long necks, and
the peridium comprising host cells and fungal tissue. In contrast, Astrosphaeriella has erumpent ascostromata through
the host epidermis, with a star-like or rounded flange and
small papilla, and the peridium comprising an opaque, black,
amorphous, brittle layer (Liu et al. 2011; Phookamsak et al.
2015b). Seven species are accepted in Pseudoastrosphaeriella, five of which are from freshwater habitats. All freshwater Pseudoastrosphaeriella species (see list below), have
rough-walled ascospores with minute striations or longitudinal ridges (Phookamsak et al. 2015b), except our new
species P. aquatica which has smooth-walled ascospores.
List of freshwater Pseudoastrosphaeriella species
*Pseudoastrosphaeriella africana (D. Hawksw.) Phook. &
K.D. Hyde, Fungal Diversity 74: 183 (2015)
Basionym: Astrosphaeriella africana D. Hawksw.,
Sydowia 38: 116 (1986) [1985]
Freshwater distribution: Thailand (Phookamsak et al.
2015b)
*Pseudoastrosphaeriella aquatica W. Dong, H. Zhang &
K.D. Hyde, sp. nov.
Index Fungorum number: IF557932; Facesoffungi number: FoF09275; Fig. 96
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-0996
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascostromata 250–350 μm high, 280–320
μm diam., black, scattered or gregarious, immersed with
neck erumpent through host tissue, becoming raised, subglobose, coriaceous, ostiolate. Neck 450–500 μm high, 100–110
μm diam., black, central, cylindrical, oblique, brittle, thickwalled, comprising some layers of blackened cells, carbonaceous. Peridium 15–25 μm thick, of unequal thickness,
composed of thin-walled, hyaline to brown, compressed
cells of textura angularis or textura prismatica, outer layers
comprising host cells and fungal tissue. Pseudoparaphyses 1 μm diam., numerous, trabeculate, filiform, hyaline,
branching and anastomosing, embedded in a gelatinous
matrix. Asci 145–260 × 19–25 μm ( x̄ = 213 × 22 μm, n =
15), 8-spored, bitunicate, distinctly clavate, or cylindrical,
with elongate pedicel, up to 115 μm long, with indistinct
ocular chamber. Ascospores 40–48 × 8–10.5 μm ( x̄ = 44.5
× 9.5 μm, n = 30), overlapping uni-seriate or uni-seriate at
the base and bi- to tri-seriate near the apex, fusiform, with
acute or rounded ends, brown, 3-septate, constricted at the
Fungal Diversity (2020) 105:319–575
487
Fig. 96 Pseudoastrosphaeriella aquatica (MFLU 18-0996,
holotype). a, b Ascomata
immersed with neck erumpent
through host substrate. c, d
Vertical section of ascomata.
e Structure of peridium. f,
g Asci embedded in numerous of pseudoparaphyses. h, i
Bitunicate asci. j–l Ascospores.
m Germinated ascospores. n,
o Colony on PDA (up-front,
down-reverse). Scale bars: c,
d = 100 μm, e = 10 μm, f, g =
30 μm, h, i, m = 50 μm, j–l =
20 μm
septa, swollen above central septum, guttulate, thin-walled,
smooth. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 20 mm in 45 days at 25 °C, grey from above, black from
below, surface rough, with dense mycelium, dry, edge entire.
Material examined: THAILAND, Chiang Mai Province,
on submerged wood in a stream, 9 February 2018, X.D.
Yu, Y18 (MFLU 18-0996, holotype), ex-type living culture MFLUCC 18-0984; Phayao Province, on submerged
wood, 23 February 2018, X.D. Yu, Y27 (MFLU 18-1005),
living culture MFLUCC 18-0991; Chiang Rai Province, on
submerged wood, 10 April 2018, X.D. Yu, Y41 (MFLU
18-1016), living culture KUMCC 19-0096.
Notes: Pseudoastrosphaeriella aquatica is similar to P.
papillata in having black ascomata with long necks and
similar in shape and overlapping size of ascospores, but the
smooth-walled, 3-septate ascospores of the former differs
from the rough-walled, mostly 1-septate ascospores with
small, pad-like appendages of the latter (Hyde and Frohlich
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488
Fungal Diversity (2020) 105:319–575
Fig. 97 Pseudoastrosphaeriella bambusae (MFLU 18-1704). a, b Ascomata immersed with neck erumpent through host substrate. c–g Bitunicate asci. h–k Ascospores. i–k Ascospores in Indian Ink. l Germinated ascospore. Scale bars: c–g = 30 μm, h, j, k = 10 μm, i, l = 20 μm
1998; Phookamsak et al. 2015b). Phylogenetic analysis supports our collections to be a new species (Fig. 2).
*Pseudoastrosphaeriella bambusae Phook. & K.D. Hyde,
Fungal Diversity 74: 186 (2015); Figs. 97, 98
Freshwater distribution: Thailand (Phookamsak et al.
2015b; this study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascostromata 200–300 µm high, 200–400
µm diam., black, scattered to gregarious, immersed with
neck erumpent through host tissue, becoming raised,
13
hemispherical to conical, with flattened base and long neck,
unilocular, coriaceous, ostiolate. Peridium 10–30 µm thick,
composed of dark, thick-walled, angular, pseudoparenchymatous, occluded cells. Pseudoparaphyses 1.5 μm diam.,
numerous, trabeculate, filiform, hyaline, anastomosing at
the apex, embedded in a gelatinous matrix. Asci 120–240
× 12–17 μm ( x̄ = 188 × 14 μm, n = 10), including elongate
pedicel, 8-spored, bitunicate, cylindric-clavate to clavate,
with furcate or foot-like pedicel, become elongate when in
water, with an ocular chamber. Ascospores 31–50 × 5.5–8.2
μm ( x̄ = 41 × 7.2 μm, n = 20), biseriate, fusiform, with
Fungal Diversity (2020) 105:319–575
489
Fig. 98 Pseudoastrosphaeriella
bambusae (MFLU 10-0160).
a, b Ascomata immersed with
neck erumpent through host
substrate. c Vertical section of
ascoma. e Pseudoparaphyses.
d, f, g Bitunicate asci. h–k
Ascospores. (h, k ascospores
show minute striations; i
ascospore in Indian Ink). l
Germinated ascospore. Scale
bars: c = 50 μm, d, f, l = 20 µm,
e, g–k = 5 µm
acute or rounded ends, slightly reddish brown, 1(–3)-septate,
constricted at the central septum, slightly swollen above central septum, guttulate, thin-walled, smooth or rough, with a
mucilaginous sheath; sheath elongate at both ends and winglike at the middle. Asexual morph: Undetermined.
Material examined: THAILAND, Chiang Rai Province,
on submerged wood in a stream, 1 July 2018, W. Dong,
CR131 (MFLU 18-1698), living cultue KUMCC 19-0091;
ibid., CR133 (MFLU 18-1704), living cultue KUMCC
19-0093; ibid., CR142 (MFLU 18-1699), living cultue
KUMCC 19-0095; ibid., 18 January 2010, H. Zhang, a4
(MFLU 10-0160), living cultue MFLUCC 10-0885.
Notes: Our collections are identified as Pseudoastrosphaeriella bambusae based on the identical LSU and TEF
sequence data with the ex-type strain MFLUCC 11-0205,
and phylogenetic analysis (Fig. 2). Phookamsak et al.
(2015b) described P. bambusae as having rough-walled
ascospores with minute striations surrounded by mucilaginous sheath. However, the minute striations are not found in
our three collections (KUMCC 19-0091, KUMCC 19-0093,
KUMCC 19-0095), but obvious in MFLUCC 10-0885
(Fig. 98). The ascospore sheaths in our four collections are
distinctly elongate at both ends and wing-like at the middle
(Figs. 97, 98), a feature not fully described in the holotype
(Phookamsak et al. 2015b).
13
490
*Pseudoastrosphaeriella longicolla Phook. & K.D. Hyde,
Fungal Diversity 74: 186 (2015)
Freshwater distribution: Thailand (Phookamsak et al.
2015b)
Pseudoastrosphaeriella papillata (K.D. Hyde & J. Fröhl.)
Phook. & K.D. Hyde, Fungal Diversity 74: 188 (2015)
Basionym: Astrosphaeriella papillata K.D. Hyde & J.
Fröhl., Sydowia 50(1): 109 (1998)
Freshwater distribution: Brunei (Hyde and Frohlich
1998), Philippines (Cai et al. 2003a), Yunnan, China (Luo
et al. 2004)
Key to freshwater Pseudoastrosphaeriella species
1. Ascospores smooth-walled…………………P. aquatica
1. Ascospores rough-walled………………………………2
2. Ascospores with mucilaginous sheaths or appendages……………………………………………………3
2. Ascospores without mucilaginous sheaths……………4
3. Ascospores surrounded by irregular sheaths……………
……………………………………………P. bambusae
3. Ascospores with small, pad-like, mucilaginous appendages at both ends…………………………P. papillata
4. Ascospores (39–)40–48(–50) × 5–7 μm…P. africana
4. Ascospores (48–)50–60(–63.5) × 5.5–7(–7.5) μm……
……………………………………………P. longicolla
Roussoellaceae J.K. Liu, Phook., D.Q. Dai & K.D. Hyde,
Phytotaxa 181(1): 7 (2014)
Roussoella Sacc., Atti dell’Istituto Veneto Scienze, 6: 410,
1888.
Saprobic on various flowering plants in terrestrial or
submerged wood in freshwater habitats, mainly on bamboo
and palms. Sexual morph: Ascostromata solitary to gregarious, visible as black, raised, elongate to hemisphaerical or
dome-shaped ascostroma, uni- to multi-loculate, glabrous,
coriaceous. Locules clustered, immersed under a clypeus,
subglobose to lenticular or quadrilateral dome-shaped, central ostiolate, with a minute papilla. Peridium slightly thick
at the side, composed of several layers of flattened, brown
to dark brown pseudoparenchymatous cells, arranged in
textura angularis to textura prismatica. Pseudoparaphyses dense, trabeculate, filamentous, hyaline, branching at
the apex, anastomosing, embedded in a gelatinous matrix.
Asci 8-spored, bitunicate, cylindrical to cylindric-clavate,
short pedicellate, with obtuse to knob-like pedicel, apically
rounded with well-developed ocular chamber. Ascospores
overlapping uniseriate, ellipsoidal to fusiform, pale brown
to dark brown, 1-septate, ornamented, with or without a
mucilaginous sheath (Liu et al. 2014). Asexual morph:
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Fungal Diversity (2020) 105:319–575
Coelomycetous, reported as Cytoplea (Hyde et al. 1996a).
Conidiomata pycnidial, scattered to clustered, solitary to
gregarious, semi-immersed to superficial, globose to subglobose, dark brown to black, single- or multi-loculate, locules separated by dark, palisade-like cells. Pycnidial wall
slightly thick at the base, composed of several layers of
brown to dark brown, pseudoparenchymatous cells, arranged
in textura angularis to textura prismatica. Conidiophores
arising from the basal cavity, reduced to conidiogenous cells.
Conidiogenous cells enteroblastic, phialidic, determinate,
discrete, ampulliform, hyaline, smooth-walled. Conidia globose to subglobose, oblong or ellipsoidal, brown, aseptate or
septate, smooth- or rough-walled.
Type species: Roussoella nitidula Sacc. & Paol.
Notes: Roussoella was introduced by Saccardo and Paoletti (1888) with R. nitidula as the type species. An epitype
of R. nitidula was designated by Liu et al. (2014) who also
found the Cytoplea asexual morph in culture. Roussoella
accommodates species having large ascostromata with
multi-loculate, bitunicate asci and brown, fusiform to ellipsoidal, ornamented, didymosporous ascospores which are
mostly associated with bamboo and palms (Tanaka et al.
2009; Hyde et al. 2013; Liu et al. 2014; Dai et al. 2017;
Jiang et al. 2019).
The freshwater species Roussoella minutella (Penz.
& Sacc.) Aptroot was treated as a synonym of R. pustulans (Ellis & Everh.) Y.M. Ju et al. by Hyde (1997), but
not shown in Index Fungorum (2020). We follow Cai et al.
(2003a), using the name R. minutella. Roussoella minutella
has shorter ascospores than another freshwater species R.
intermedia (see key below). We introduce a new species R.
aquatica collected from submerged wood in China.
List of freshwater Roussoella species
*Roussoella aquatica W. Dong, H. Zhang & K.D. Hyde,
sp. nov.
Index Fungorum number: IF557933; Facesoffungi number: FoF09276; Fig. 99
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-1169
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Stromata immersed under an
epidermis, becoming raised at maturity, 1–2 locules. Locules 150–180 μm high, 210–240 μm diam., immersed in
the stromata, conical or hemispherical, flattened at the base,
dark brown to black, ostiolate. Peridium 10–30 μm thick,
comprising 3–5 layers of brown, thick-walled cells of textura
angularis. Conidiophores reduced to conidiogenous cells.
Conidiogenous cells 3–4 × 1.5–2 μm, enteroblastic, annellidic, integrated, determinate, ampulliform, hyaline, smoothwalled. Conidia 2.7–3.5 × 2–2.5 μm ( x̄ = 3.2 × 2.2 μm, n
= 30), ellipsoidal, aseptate, straight, rounded at both ends,
brown to greenish brown, smooth-walled.
Fungal Diversity (2020) 105:319–575
491
Fig. 99 Roussoella aquatica
(MFLU 18-1169, holotype). a
Conidiomata on host substrate.
b Conidiomata cut through
vertically. c Vertical section
of conidioma. d Structure of
peridium. e Conidiogenous cells
bearing conidia. f–i Conidia. j
Germinated conidium. k Colony
on PDA (from front). Scale
bars: c = 50 μm, d, j = 20 μm, e
= 5 μm, f = 10 μm, g–i = 3 μm
Culture characteristics: On PDA, colony circular, reaching 15 mm in 25 days at 25 °C, white from above, yellowish
brown from below, surface rough, felt, with dense mycelium,
dry, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H1A-1 (MFLU 18-1169, holotype), ex-type living
culture MFLUCC 18-1040; ibid., H1A-2 (HKAS 101739,
isotype), ex-type living culture KUMCC 18-0091.
Notes: Roussoella aquatica is similar to R. tuberculata
D.Q. Dai & K.D. Hyde in having ellipsoidal, aseptate, brown
conidia. However, the latter has small, roughened tubercules
sparsely covering the conidia (Dai et al. 2017), while R.
aquatica has smooth-walled conidia. Additionally, R. aquatica has smaller conidia than R. tuberculata (2.7–3.5 × 2–2.5
μm vs. 8.5–10 × 4.5–5.5 μm). The ellipsoidal, brown, aseptate conidia of R. aquatica are also similar to R. chiangraina
Phook. et al, but the latter has obvious verrucose conidia
(Liu et al. 2014; Wijayawardene et al. 2016), and molecular
evidence shows them to be different species (Fig. 102).
*Roussoella intermedia Y.M. Ju, J.D. Rogers & Huhndorf,
Mycotaxon 58: 447 (1996)
Freshwater distribution: North America (Fallah and
Shearer 2001)
Roussoella minutella (Penz. & Sacc.) Aptroot, Nova Hedwigia 60(3–4): 367 (1995)
Basionym: Didymosphaeria minutella Penz. & Sacc.,
Malpighia 11(9-10): 396 (1897)
Freshwater distribution: Philippines (Cai et al. 2003a)
Key to freshwater Roussoella species
1. Coelomycetous……………………………R. aquatica
1. Ascomycetous…………………………………………2
13
492
2. Ascospores 9.5–13 × 3.5–4.5 μm…………R. minutella
2. Ascospores 12–16 × 4–6 μm……………R. intermedia
Neoroussoella J.K. Liu, Phook. & K.D. Hyde, Phytotaxa
181(1): 21 (2014)
Saprobic on decaying bamboo culms, submerged wood or
parasitic on human diseased eyes. Sexual morph: Ascostromata immersed under a clypeus or epidermis, raised, visible
as black dome-shaped or flattened ovoid areas on host surface, coriaceous, solitary to gregarious, black, central ostiolate. Peridium composed of several layers of light brown to
brown, thin-walled cells of textura angularis, with flattened
base. Pseudoparaphyses numerous, trabeculate, filiform,
hyaline, anastomosing, branching at the apex, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, cylindrical,
short to long pedicellate, apically rounded with an ocular
chamber. Ascospores uniseriate, ellipsoidal to fusiform,
1-septate, constricted at the septum, pale brown to brown or
yellowish brown, rough-walled (Liu et al. 2014). Asexual
morph: Coelomycetous. Conidiomata immersed to superficial, solitary to gregarious, globose to subglobose or irregular shape, uni- to multi-loculate. Pycnidial wall composed of
several layers of cells of textura angularis. Conidiophores
arising from the basal cavity, unbranched, aseptate. Conidiogenous cells annellidic, ampulliform or lageniform or
cylindrical or cylindric-clavate, hyaline. Conidia oblong
to ellipsoidal, hyaline, aseptate, smooth-walled (Liu et al.
2014).
Type species: Neoroussoella bambusae Phook., J.K. Liu
& K.D. Hyde
Notes: Neoroussoella was introduced to accommodate the
holomorphic species N. bambusae Phook. et al. collected
on dead branch in Thailand (Liu et al. 2014). The asexual
morph of N. bambusae was produced on bamboo pieces on
WA and characterized by annellidic conidiogenous cells
and oblong to ellipsoidal, hyaline conidia (Liu et al. 2014).
Neoroussoella species formed a well-supported and distinct
clade in Roussoellaceae (Jayasiri et al. 2019; Karunarathna
et al. 2019; Phookamsak et al. 2019). Neoroussoella was
considered to be different from Roussoella by relatively
smaller, hyaline and smooth conidia, and phylogenetic
analyses supported this conclusion (Liu et al. 2014; Jayasiri
et al. 2019; Phookamsak et al. 2019; this study, Fig. 102).
We re-collected N. bambusae Phook. et al. and N. leucaenae
Jayasiri et al. from submerged wood in China. Neoroussoella
bambusae has longitudinally ribbed ascospores, while N.
leucaenae has smooth-walled ascospores.
List of freshwater Neoroussoella species
*Neoroussoella bambusae Phook., Jian K. Liu & K.D.
Hyde, Phytotaxa 181(1): 23 (2014)
Facesoffungi number: FoF02414; Fig. 100
13
Fungal Diversity (2020) 105:319–575
Freshwater distribution: China (this study)
Saprobic on submerged wood. Sexual morph: Ascostromata 300–330 μm high, 370–400 diam., immersed under an
epidermis, erumpent to semi-immersed, scattered to gregarious, raised, visible as black dome-shaped or shield-shaped
on the host surface, coriaceous, black, central ostiolate, with
elongate, distinct papilla. Peridium composed of several
layers of textura angularis, thick-walled and dark brown to
black in outer layer, thin-walled and hyaline inwardly, flattened and thicker at the base, 50 μm wide, thinner at sides,
30 μm wide. Pseudoparaphyses 1.5–2 μm wide, numerous,
anastomosing cellular, embedded in a gelatinous matrix.
Asci (105–)113–115 × 6–8 μm ( x̄ = 114 × 7 μm, n = 10),
8-spored, bitunicate, cylindrical, short to long pedicellate, apically rounded with an ocular chamber. Ascospores
11–14.5 × 4–5.5 μm ( x̄ = 12.5 × 5 μm, n = 15), uniseriate
to overlapping uniseriate, ellipsoidal to broadly fusiform,
1-septate, constricted at the septum, pale brown to brown
or yellowish brown, guttulate, rough-walled, longitudinally
ribbed. Asexual morph: Undetermined.
Culture characteristics: On PDA, colony circular, reaching 15 mm in 25 days at 25 °C, white from above, yellowish
brown from below, surface rough, felt, with dense mycelium,
dry, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, HH1A-1 (MFLU 18-1187), living culture MFLUCC
18-1048; ibid., HH1A-2 (HKAS 101750), living culture
KUMCC 18-0102.
Notes: Our collection MFLUCC 18-1048 clusters with
Neoroussoella bambusae with high bootstrap support
(Fig. 102). MFLUCC 18-1048 is identified as N. bambusae
based on sequence data which has three, seven (includes
three gaps) and three (includes one gap) nucleotide differences in LSU, ITS and TEF sequence data, respectively, with
the ex-type strain MFLUCC 11-0124.
Our collection has longer asci (105–115 × 6–8 μm vs.
60–85 × 5–6.5 μm) and ascospores (11–14.5 × 4–5.5 μm
vs. 7.5–10 × 3–4 μm) than the holotype (Liu et al. 2014).
The elongate, distinct papilla is observed in our collection,
but lacking in the holotype (Liu et al. 2014). This is a new
habitat and geographical record for N. bambusae from freshwater in China.
*Neoroussoella leucaenae Jayasiri, E.B.G. Jones & K.D.
Hyde, Mycosphere 10(1): 106 (2019); Fig. 101
Freshwater distribution: China (this study)
Saprobic on submerged wood. Sexual morph: Ascostromata 140–180 μm high, 110–170 diam., immersed to
erumpent, scattered, raised, visible as black, dome-shaped
spots and with prominent, elongate neck, coriaceous, black,
ostiolate. Ostiole 60–90 μm diam., rounded, brown to dark
brown, central. Peridium 10–15 μm wide, composed of
Fungal Diversity (2020) 105:319–575
493
Fig. 100 Neoroussoella bambusae (MFLU 18-1187, new habitat
and geographical record). a, b Ascostromata on host substrate. c
Vertical section of ascoma. d Structure of peridium. e Pseudopara-
physes. f–i Bitunicate asci. j–l Ascospores. m, n Colony on PDA
(left-front, right-reverse). Scale bars: c = 100 μm, d–i = 20 μm, j–l
= 5 μm
several layers of compressed cells of textura angularis, dark
brown to black in outer layer, hyaline inwardly, flattened at
the base. Pseudoparaphyses 3 μm wide, numerous, cellular, narrow, filamentous, indistinctly septate, anastomosing,
embedded in a gelatinous matrix. Asci 65–145 × 4.7–6.5 μm
( x̄ = 98 × 5.5 μm, n = 10), 8-spored, bitunicate, cylindrical, pedicellate, apically rounded with an ocular chamber,
elongate when in water. Ascospores 7.5–10.7 × 3–4.5 μm ( x̄
= 8.8 × 3.5 μm, n = 30), overlapping uniseriate, ellipsoidal
to broadly fusiform, 1-septate, constricted at the septum,
pale brown to brown, smooth-walled. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony circular, reaching 50 mm in 30 days at 25 °C, white from above, become
reddish with age, dark brown from below, surface rough,
felt, with dense mycelium, dry, edge undulate.
Material examined: CHINA, Yunnan Province, Pingbian,
on submerged wood in a stream, 20 September 2017, W.
Dong, WF11A-1 (HKAS 101701), living culture KUMCC
18-0050; ibid., WF11A-2 (MFLU 18-1175).
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Fungal Diversity (2020) 105:319–575
Fig. 101 Neoroussoella
leucaenae (HKAS 101701,
new habitat and geographical record). a, b Ascostromata
on host substrate. c, d Vertical
section of ascomata. e Structure
of peridium. f Pseudoparaphyses. g–j Bitunicate asci. k–m
Ascospores. n Colony on PDA
(from front). Scale bars: c, d =
50 μm, e, g–j = 20 μm, f = 10
μm, k–m = 5 μm
Notes: Our collection KUMCC 18-0050 clusters with
Neoroussoella leucaenae with high bootstrap support
(Fig. 102). KUMCC 18-0050 is identified as N. leucaenae based on sequence data which has 100% similarity in
ITS sequence data, and one nucleotide difference in TEF
sequence data, with the ex-type strain MFLUCC 18-1544.
13
Our collection has longer asci (65–145 × 4.7–6.5 μm vs.
50–60 × 4.5–5.5 μm) than the holotype (Jayasiri et al. 2019).
This is a new habitat and geographical record for N. leucaenae from freshwater in China.
Key to freshwater Neoroussoella species
Fungal Diversity (2020) 105:319–575
495
Fig. 102 Phylogram generated from maximum likelihood
analysis of combined LSU,
SSU, ITS, TEF and RPB2
sequence data for species of
Roussoellaceae. Bootstrap
values for maximum likelihood
equal to or greater than 75% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree is
rooted to Torula herbarum CBS
111855 (Torulaceae)
98/0.99
97/0.96
80/--
Roussoella japanensis MAFF 239636
Roussoella hysterioides CBS 546.94
Roussoella verrucispora CBS 125434
Roussoella scabrispora MFLUCC 11-0624
98/0.99
Roussoellopsis macrospora MFLUCC 12-0005
Roussoellopsis tosaensis KT 1659
98/-Roussoella intermedia NBRC 106245
Roussoella pustulans MAFF 239637
Elongatopedicellata lignicola MFLUCC 15-0642
98/0.99
3x
Roussoella pseudohysterioides MFLUCC 13-0852
Roussoella nitidula MFLUCC 11-0182
Roussoella nitidula MFLUCC 11-0634
Roussoella tuberculata MFLUCC 13-0854
Roussoella thailandica MFLUCC 11-0621
100/0.99
Roussoella aquatica MFLUCC 18-1040 F
Roussoella neopustulans MFLUCC 12-0003
Roussoella neopustulans MFLUCC 11-0609
Roussoella kunmingensis KUMCC 18-0128
81/0.99
Roussoella magnatum MFLUCC 15-0185
Roussoella chiangraina MFLUCC 10-0556
Roussoella angustior MFLUCC 15-0186
Roussoella siamensis MFLUCC 11-0149
Roussoella doimaesalongensis MFLUCC 14-0584
Roussoella yunnanensis KUMCC 18-0115
80/-- Roussoella intermedia CBS 170.96
89/1.00
Arthopyrenia salicis UTHSC DI 16-220
Roussoella mexicana CPC 25355
Arthopyrenia salicis CBS 368.94
Neoroussoella leucaenae MFLUCC 17-0927
Neoroussoella leucaenae KUMCC 18-0050 F
Neoroussoella leucaenae MFLUCC 17-0346
98/-- Neoroussoella leucaenae KT 3264
79/-- Neoroussoella leucaenae KT 3265
87/0.99 Neoroussoella solani CPC 26331
Neoroussoella leucaenae MFLUCC 18-1544
96/0.99
91/0.99
Neoroussoella entadae MFLUCC 15-0098
92/-- Neoroussoella entadae MFLUCC 18-0243
Neoroussoella lenispora GZCC 16-0020
75/-Neoroussoella heveae MFLUCC 17-0338
100/1.00 Neoroussoella alishanense FU31016
100/1.00
Neoroussoella alishanense FU31018
Neoroussoella bambusae MFLUCC 18-1048 F
100/1.00 Neoroussoella bambusae MFLUCC 11-0124
92/1.00
100/1.00 Roussoella elaeicola MFLUCC 15-0276a
Roussoella elaeicola MFLUCC 15-0276b
Pseudoneoconiothyrium rosae MFLUCC 15-0052
99/1.00
100/1.00 Roussoella euonymi CBS 143426
97/0.99
Pararoussoella rosarum MFLUCC 17-0796
Pararoussoella mukdahanensis MFLUCC 11-0201
Roussoella mangrovei MFLUCC 16-0424
100/0.99 Thyridaria broussonetiae CBS 141481
98/1.00
Thyridaria broussonetiae CBS 121895
Thyridaria acaciae CBS 138873
100/1.00 Parathyridaria ramulicola MF4
98/1.00
Parathyridaria ramulicola CBS 141479
Parathyridaria robiniae MFLUCC 14-1119
76/0.97 100/1.00 Parathyridaria percutanea CBS 868.95
Parathyridaria percutanea CBS 128203
Torula herbarum CBS 111855
Roussoella
Neoroussoella
Outgroup
0.07
1. Ascospores longitudinally ribbed…………N. bambusae
1. Ascospores smooth-walled………………N. leucaenae
Testudinaceae Arx, Persoonia 6(3): 366 (1971)
Angustospora Abdel-Aziz, Fungal Diversity 78: 52 (2016)
Saprobic on submerged wood. Sexual morph: Ascomata solitary, immersed to erumpent, uniloculate, globose
to subglobose, glabrous, dark brown to black, coriaceous
to sub-carbonaceous, with ostiolate papilla. Ostiole central,
circular, periphysate. Peridium two-layered, outer layer
comprising dark brown to black cells of textura angularis,
inner layer comprising hyaline, thick-walled, flattened cells
of textura angularis. Pseudoparaphyses numerous, trabeculate, hyaline, distantly septate, branched, persistent, within
a gelatinous matrix and anastomosing above asci. Asci
8-spored, bitunicate, fissitunicate, clavate, short pedicellate, apically rounded, with a wide, shallow ocular chamber
and faint ring. Ascospores overlapping biseriate, ellipsoidal
with pointed ends, straight or curved, dark brown to black,
septate, polar cells lighter when young and apical cells
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Fungal Diversity (2020) 105:319–575
Fig. 103 Angustospora nilensis
(Material examined: EGYPT,
Sohag City, on decayed wood
submerged in the River Nile, 8
March 2005, F.A. Abdel-Aziz,
MFLU 15-1511, holotype).
a Vertical section of ascoma.
b Immature ascus. c Mature
ascus. d Ocular chamber in
ascus and faint apical ring. e, f
Ascospores at different stages of
maturity. Scale bars: a–c = 60
μm, d–f = 10 μm
with two walls, mostly symmetrical, surrounded by thin,
gelatinous, striate layer (Li et al. 2016a). Asexual morph:
Undetermined.
Type species: Angustospora nilensis Abdel-Aziz
Notes: Angustospora was introduced to accommodate
the freshwater species A. nilensis collected from Egypt (Li
et al. 2016a). Angustospora is reminiscent of Caryospora
in having large ascospores with a median septum and two
additional septa near poles (Ariyawansa et al. 2015; Li et al.
2016a; Jayasiri et al. 2019). However, Angustospora differs from the latter in having smaller ascomata, ellipsoidal
ascospores with paler end cells when young and two-walled
apical cells, and surrounded by gelatinous, striate layer (Li
et al. 2016a). Phylogenetic analyses placed Angustospora
and Caryospora in Testudinaceae and Caryosporaceae,
respectively (Li et al. 2016a; Wanasinghe et al. 2017).
Angustospora appears exclusively saprobic in freshwater
habitats.
List of freshwater Angustospora species
*Angustospora nilensis Abdel-Aziz, Fungal Diversity 78:
54 (2016); Fig. 103
Freshwater distribution: Egypt (Li et al. 2016a)
13
Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray., Stud.
Mycol. 64: 177 (2009)
Key to freshwater genera of Tetraplosphaeriaceae
1. Conidia cylindrical…………………………Tetraploa
1. Conidia not as above……………………………………2
2. Conidia variable in shape, but often obconical or pyriform………………………………………Ernakulamia
2. Conidia ovoid to obpyriform, or subglobose…………3
3. Conidia with apical appendages………Triplosphaeria
3. Conidia not as above……………………Shrungabeeja
Ernakulamia Subram., Kavaka 22/23: 67 (1996) [1994]
Saprobic on submerged wood in freshwater and rotten
leaves, branches in terrestrial habitats. Sexual morph:
Undetermined. Asexual morph: Colonies effuse, dark
brown to black. Mycelium superficial. Conidiophores semimacronematous. Conidiogenous cells monotretic, integrated,
determinate, terminal and intercalary, often clavate, globose
or subglobose, smooth or verruculose, subhyaline to brown,
cicatrized or non-cicatrized, with a well-defined pore in the
middle of each dark scar. Conidia variable in shape but
Fungal Diversity (2020) 105:319–575
often obconical or pyriform, dark brown to blackish brown,
muriform, often verrucose at the base where a pore is often
seen, internally filled with a mass of hyaline, septate hyphae
sometimes with swollen cells, appendiculate, with cylindrical, straight or flexuous, septate, brown, appendages (Ellis
1976; Delgado et al. 2018).
Type species: Ernakulamia cochinensis (Subram.)
Subram.
Notes: Ernakulamia is characterized by monotretic conidiogenous cells (Ellis 1976) and confirmed based on the
cultural method (Delgado et al. 2018). Ernakulamia was
placed in Tetraplosphaeriaceae based on two specimens
of E. cochinensis (PRC 3992 and IMI 114626) (Delgado
et al. 2018). They found an internal hyphal structure in E.
cochinensis which were also present in Polyplosphaeria
Kaz. Tanaka & K. Hiray. and Quadricrura Kaz. Tanaka et al.
(Tanaka et al. 2009; Delgado et al. 2018). Phylogeny also
confirmed the placement of Ernakulamia in Tetraplosphaeriaceae (Delgado et al. 2018; Hyde et al. 2020b; this study,
Fig. 110). Ernakulamia cochinensis was reported from terrestrial habitats (Delgado et al. 2018), while our collection
is from freshwater habitats.
List of freshwater Ernakulamia species
*Ernakulamia cochinensis (Subram.) Subram., Kavaka
22/23: 67 (1996) [1994]
Facesoffungi number: FoF09277; Fig. 104
Basionym: Petrakia cochinensis Subram., Beih. Sydowia
1: 15 (1957) [1956]
Synonymy: Piricauda cochinensis (Subram.) M.B. Ellis,
More Dematiaceous Hyphomycetes (Kew): 367 (1976)
Freshwater distribution: Thailand (this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, gregarious, black.
Mycelium composed of pale brown, septate hyphae. Conidiophores and conidiogenous cells not seen. Conidia 55–85 ×
35–55(–70) μm (x̄ = 70 × 14 μm, n = 15), variable in shape
but often obconical or pyriform, black, muriform which is
not clear when mature, thin-walled, easily cracked, often
verrucose at base, with numerous (up to 10), cylindrical,
septate, brown, smooth appendages, 6–105 × 3–4 μm.
Culture characteristics: On PDA, colony ellipsoidal,
reaching 35 mm in 25 days at 25 °C, grey from above,
brown from below, surface rough, with dense mycelium,
dry, raised, edge entire.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat161 (MFLU 18-1565), living culture MFLUCC 18-1237.
Notes: Our new collection MFLUCC 18-1237 is identified as Ernakulamia cochinensis based on identical LSU and
497
ITS sequence data with PRC 3992, and phylogenetic analysis (Fig. 110). MFLUCC 18-1237 has larger conidia than in
PRC 3992 from the natural substrate (55–85 × 35–55(–70)
μm vs. 24–60 × 18–53 μm), but resembles with those from
culture (36–81 × 19–56 μm) (Delgado et al. 2018). The
conidiogenous cells of E. cochinensis cannot be observed
from mature, blackish conidia in our collection. This is a
new habitat and geographical record for E. cochinensis from
freshwater in Thailand.
New combination of Ernakulamia
*Ernakulamia xishuangbannaensis (Tibpromma & K.D.
Hyde) W. Dong, H. Zhang & K.D. Hyde, comb. nov.
Index Fungorum number: IF557934; Facesoffungi number: FoF09278
Basionym: Polyplosphaeria xishuangbannaensis Tibpromma & K.D. Hyde, Fungal Diversity 93: 44 (2018)
Description of this species see Tibpromma et al. (2018)
Notes: Polyplosphaeria xishuangbannaensis was collected from dead leaves of Pandanus sp. (Tibpromma
et al. 2018). In the phylogenetic analysis of Tibpromma
et al. (2018), they did not include Ernakulamia so that P.
xishuangbannaensis clustered with the members of Polyplosphaeria, but without bootstrap support. In Hyde et al.
(2020b) and our study (Fig. 110), P. xishuangbannaensis
formed a well-supported and stable clade with the members
of Ernakulamia. Polyplosphaeria xishuangbannaensis has
black conidia which are features of Ernakulamia (Ellis 1976;
Delgado et al. 2018; Jayasiri et al. 2019), while Polyplosphaeria Kaz. Tanaka & K. Hiray. has brown conidia (Tanaka
et al. 2009; Tibpromma et al. 2018). Based on phylogenetic
analyses and morphology, we transfer P. xishuangbannaensis to Ernakulamia. However, Tibpromma et al. (2018)
described the monoblastic conidiogenous cells from P. xishuangbannaensis, which are different from the monotretic
one of Ernakulamia. More fresh collections are needed to
clarify the conidiogenesis of P. xishuangbannaensis.
Shrungabeeja V.G. Rao & K.A. Reddy, Indian J. Bot. 4(1):
109 (1981)
Saprobic on dead branch, culm or submerged wood.
Sexual morph: Colonies effuse, blackish brown. Mycelium
superficial or immersed. Conidiophores macronematous,
mononematous, cylindrical, erect, straight or flexuous,
unbranched, septate, pale brown to deep brown, smooth.
Conidiogenous cells monoblastic, determinate or percurrently proliferative, pale brown to brown, smooth. Conidia
acrogenous, subglobose or turbinate, aseptate, with appendages (Zhang et al. 2009a). Asexual morph: Undetermined.
13
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Fungal Diversity (2020) 105:319–575
Fig. 104 Ernakulamia
cochinensis (MFLU 18-1565,
new habitat and geographical record) a Colonies on
submerged wood. b–h Conidia.
i Germinated conidium. j, k
Colony on PDA (left-front,
right-reverse). Scale bars: b–i
= 50 μm
Type species: Shrungabeeja vadirajensis V.G. Rao &
K.A. Reddy
Notes: Shrungabeeja is characterized by macronematous
conidiophores, monoblastic, determinate or percurrent conidiogenous cells and subglobose or turbinate conidia with
appendages (Zhang et al. 2009a). Ariyawansa et al. (2015)
confirmed Shrungabeeja in Tetraplosphaeriaceae based on
molecular data of S. longiappendiculata Sommai et al. The
type species S. vadirajensis was initially collected from India
(Rao and Reddy 1981), later found from China (Zhang et al.
2009a). The sequence data of S. vadirajensis was lacking in
previous studies (Rao and Reddy 1981; Zhang et al. 2009a;
Ariyawansa et al. 2015). We describe another reference
specimen of S. vadirajensis with molecular data. Shrungabeeja species are often recorded from terrestrial habitats
(Zhang et al. 2009a; Ariyawansa et al. 2015; Kirschner et al.
2017), we report one new species and one new habitat and
geographical record from freshwater. The morphology of
13
Shrungabeeja species are very similar and thus sequence
data are of crucial importance to separate them.
List of freshwater Shrungabeeja species
*Shrungabeeja aquatica W. Dong, G.N. Wang & H. Zhang,
sp. nov.
Index Fungorum number: IF557935; Facesoffungi number: FoF09279; Fig. 105
Etymology: referring to aquatic habitat of this fungus
Holotype: MFLU 18-0978
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, brown to dark
brown. Mycelium partly superficial, partly immersed, composed of branched, septate, pale brown hyphae. Conidiophores 140–260 μm long, 6–8.5 μm wide, macronematous,
mononematous, erect, straight or flexuous, unbranched,
smooth, thick-walled, 4–10-septate, pale- to dark brown.
Conidiogenous cells 7–10 μm long, 5–5.5 μm wide at
base, up to 3.2 μm wide at the apex, monoblastic, terminal,
Fungal Diversity (2020) 105:319–575
499
Fig. 105 Shrungabeeja
aquatica (MFLU 18-0978,
holotype). a, b Colonies on
submerged wood. c–e Conidiophores with conidiogenous
cells. f–j Conidia. Scale bars: c,
f, i, j = 30 μm, d, e = 15 μm, g,
h = 50 μm
lageniform, determinate or percurrently proliferating, pale
brown to brown, smooth. Conidia 30–40 μm diam. ( x̄ = 35
μm, n = 15), solitary, dry, acrogenous, subglobose, aseptate,
pale brown to brown, internally filled with a mass of hyaline,
septate hyphae sometimes with swollen cells, with 4(–5)
appendages. Appendages 12–135 × 2.5–3.5 μm, filiform,
5–7-septate, smooth, pale brown to brown.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 15 days at 25 °C, grey from above and below,
surface rough, with dense mycelium, dry, raised, edge entire.
Material examined: THAILAND, Chiang Mai Province, on submerged wood in a stream, 29 March 2018, G.N.
Wang, L1 (MFLU 18-0978, holotype), ex-type living culture MFLUCC 18-0664.
Notes: Shrungabeeja aquatica is similar to S. vadirajensis in conidial size and length of appendages (Zhang et al.
2009a). However, the former has obvious percurrent proliferations of conidiogenous cells, which are not obvious
in S. vadirajensis. Phylogenetic analysis separates them as
distinct species (Fig. 110).
*Shrungabeeja vadirajensis V.G. Rao & K.A. Reddy,
Indian J. Bot. 4(1): 113 (1981)
Facesoffungi number: FoF09280; Fig. 106
Freshwater distribution: Thailand (this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, blackish brown.
Mycelium superficial or immersed. Conidiophores 135–220
13
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Fungal Diversity (2020) 105:319–575
Fig. 106 Shrungabeeja
vadirajensis (MFLU 17-1721,
new habitat and geographical record). a, b Colonies
on submerged wood. c–h
Conidiophores, conidogenous
cells and conidia. i–m Conidia.
n, o Colony on PDA (up-front,
down-reverse). Scale bars: c–k
= 30 μm, l, m = 50 μm
μm long, 6–7 μm wide, macronematous, mononematous,
erect, cylindrical, pale brown to deep brown, unbranched,
4–7-septate, straight or flexuous, smooth, thick-walled. Conidiogenous cells monoblastic, determinate or percurrently
proliferative, pale brown to brown, smooth. Conidia 28–45
μm diam. ( x̄ = 32.5 μm, n = 15), 3–4 μm wide at the base,
acrogenous, subglobose, ellipsoidal, or turbinate, aseptate,
internally filled with a mass of hyaline, septate hyphae sometimes with swollen cells, with (3–)4 horn-like appendages in
the anterior region, appendages 20–110 μm long, 3–5.5 μm
wide at the base and 1.5–2.5 μm wide at the apex.
Culture characteristics: On PDA, colony circular, reaching 20 mm in 15 days at 25 °C, grey from above, dark
brown from below, surface rough, with dense mycelium,
dry, raised, edge curved.
Material examined: THAILAND, Chiang Mai Province,
on submerged wood in a stream, 1 September 2017, X.D.
Yu, 4A (MFLU 17-1721), living culture MFLUCC 17-2362.
13
Notes: Our new collection MFLUCC 17-2362 is identified as Shrungabeeja vadirajensis based on the characters of conidia (shape and size), appendages (number and
length), conidiophores (length and width) and from tropical
area (Zhang et al. 2009a). We found several hyaline, septate
hyphae or sometimes swollen cells in the conidia which were
not described in HMAS189369 (Zhang et al. 2009a).
We further confirm placement of Shrungabeeja in Tetraplosphaeriaceae based on the fresh specimen of the type
species with molecular data (Fig. 110) and hyaline, septate
hyphae in conidia, which are also present in Ernakulamia, Polyplosphaeria and Quadricrura Kaz. Tanaka et al.
(Tanaka et al. 2009). This is a new habitat and geographical
record for S. vadirajensis from freshwater in Thailand.
Key to freshwater Shrungabeeja species
1. Conidiophores with obvious percurrent proliferations…
………………………………………………S. aquatica
1. Not as above……………………………S. vadirajensis
Fungal Diversity (2020) 105:319–575
Tetraploa Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 5:
459 (1850)
Description of this genus see Hyde et al. (2013)
Type species: Tetraploa aristata Berk. & Broome
Notes: Tetraploa was reinstated, suppressing Tetraplosphaeria Kaz. Tanaka & K. Hiray. by Hyde et al. (2013).
Hyde et al. (2020b) doubted that Dwibahubeeja N. Srivast.
et al. was a synonym of Tetraploa because of the similarities of T. dwibahubeeja Rajeshkumar et al. and D. indica N.
Srivast. et al. (type). However, this cannot be resolved until
sequence data of D. indica is available.
Tetraploa is similar to other genera in Tetraplosphaeriaceae in having conidia with several setose appendages,
but differs by its globose ascomata, narrowly fusiform
ascospores surrounded by a narrow mucilaginous appendage-like sheath and cylindrical, euseptate conidia with setose
appendages (Hyde et al. 2013).
The freshwater species Tetraploa scheueri Kaz. Tanaka
& K. Hiray. was proposed by Hyde et al. (2013) to replace
Tetraplosphaeria tetraploa (Scheuer) Kaz. Tanaka & K.
Hiray. The asexual morph of T. scheueri was reported as
T. aristata Berk. & Broome based on cultural methods
(Scheuer 1991). However, Tanaka et al. (2009) considered T.
aristata to be a species-complex composed of phylogenetically diverse lineages. Thus, the identification of T. aristata
should be re-evaluated (Tanaka et al. 2009). In our phylogenetic analysis (Fig. 110), two putative strains T. aristata
(CBS 996.70) and T. scheueri (CY112) cluster together and
there are two nucleotide differences in ITS sequence data,
which probably supports the conclusion of Scheuer (1991).
However, the real identities of two strains are unknown
because their morphology are unavailable (Tanaka et al.
2009; Rodrigues et al. 2011).
List of freshwater Tetraploa species
Tetraploa abortiva Aramb. & Cabello, Mycotaxon 30: 266
(1987)
Freshwater distribution: Argentina (Schoenlein-Crusius
and Grandi 2003)
*Tetraploa aristata Berk. & Broome, Ann. Mag. nat. Hist.,
Ser. 2 5: 459 (1850)
Freshwater distribution: Argentina and Brazil (Schoenlein-Crusius and Grandi 2003), China (Tsui et al. 2001c),
Hungary (Révay 1993), India (Karamchand and Sridhar
2008), Peru (Schoenlein-Crusius and Grandi 2003)
Tetraploa ellisii Cooke Grevillea 8(no. 45): 12 (1879)
Freshwater distribution: Peru (Schoenlein-Crusius and
Grandi 2003)
*Tetraploa puzheheiensis W. Dong, H. Yang & H. Zhang,
sp. nov.
501
Holotype: MFLU 20-0441
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Mycelium superficial. Conidiophores absent. Conidiogenous cells monoblastic. Conidia
24–35 × 15–24.5 μm (x̄ = 28.5 × 17.5 μm, n = 40), solitary,
short cylindrical, dark brown to black, verrucose, consisting of 2–3 columns of 7–9.5 µm wide, 4–5-celled, with 1–3
appendages. Appendages 3–27 µm long, 3–4.5 µm wide at
the base, 2–3.5 µm wide at the apex, 0–2-septate, dark brown
at the base and hyaline to pale brown at the apex, smooth,
unbranched, straight.
Cultural characteristics: Conidia germinated on PDA
within 24 hours, germtubes produced from apex of conidia.
Colonies on PDA reaching 18–20 mm diam. in 18 days at
25 °C, with dense, floccose, brown mycelium on the surface, reverse brown to dark brown, with undulate, pale
brown margin.
Material examined: CHINA, Yunnan Province, Puzhehei
Wetland, 24°8ʹ32ʺN, 104°6ʹ58ʺE, on submerged wood, 12
May 2018, H. Yang, P12 (MFLU 20-0441, holotype), extype living culture KUMCC 20-0151.
Notes: our collection KUMCC 20-0151 clusters with two
putative strains of Tetraploa aristata (CBS 996.70) and T.
scheueri (CY112) with high bootstrap support (Fig. 110).
Tetraploa scheueri CY112 was sequenced by Rodrigues
et al. (2011), without morphological description. However,
KUMCC 20-0151 differs from the holotype of T. scheueri
GZU 36-91 in having shorter appendages (3–27 µm vs.
263–350 µm) with less septa (0–2-septate vs. 17–22-septate) (Tanaka et al. 2009). The morphology of T. aristata
CBS 996.70 also cannot be compared as the conidia were
not produced in the culture (Tanaka et al. 2009). Ellis
(1949) reported two types of conidia of T. aristata, however,
KUMCC 20-0151 only has one type of conidia. Although
type A of T. aristata has similar conidial size with KUMCC
20-0151, T. aristata differs in having longer appendages
(12–80 µm vs. 3–27 µm) with more septa (1–6-septate vs.
0–2-septate). In addition, there are only two nucleotide differences in ITS sequence data between T. aristata (CBS
996.70) and T. scheueri (CY112), which indicates that
these two strains are very likely to be the same species and
at least one strain was misidentified. The real identities of
CBS 996.70 and CY112 cannot be resolved until their epitypification with sufficient DNA sequence data are obtained
in future. Given this, we introduce KUMCC 20-0151 as a
new species T. puzheheiensis. Two strains CBS 996.70 and
CY112 are named as Tetraploa spp. in our phylogenetic tree
(Fig. 110).
*Tetraploa scheueri Kaz. Tanaka & K. Hiray., Fungal
Diversity 63: 253 (2013); Fig. 108
Index Fungorum number: IF557936; Facesoffungi number: FoF09281; Fig. 107
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Fungal Diversity (2020) 105:319–575
Fig. 107 Tetraploa puzheheiensis (MFLU 20-0441, holotype). a, b Colonies on natural substrate. c–j Conidia. k Germinated conidium. l, m
Colony on PDA (up-front, down-reverse). Scale bars: c–k = 10 μm
Etymology: referring to aquatic habitat of this fungus
Basionym: Massarina tetraploa Scheuer, Mycol. Res. 95:
126 (1991)
Synonymy: Tetraplosphaeria tetraploa (Scheuer) Kaz.
Tanaka & K. Hiray., Stud. Mycol. 64: 182 (2009)
Freshwater distribution: UK (Scheuer 1991)
*Tetraploa yunnanensis W. Dong, H. Yang & H. Zhang,
sp. nov.
Index Fungorum number: IF557937; Facesoffungi number: FoF09282; Fig. 109
Etymology: referring to Yunnan province, where the holotype was collected
Holotype: MFLU 19-0524
13
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Mycelium superficial. Conidiophores absent. Conidiogenous cells monoblastic. Conidia
30–50 × 16–22 μm (x̄ = 43 × 19 μm, n = 10), solitary, short
to long cylindrical, brown to dark brown, verrucose, consisting of 2–3 columns of 7.5–10 µm wide, 4–6-celled, with 1–3
appendages. Appendages 25–83 µm long, 4–5.5 µm wide at
the base, 1.5–2.5 µm wide at the apex, 1–5-septate, brown,
smooth, unbranched, straight.
Cultural characteristics: Conidia germinated on PDA
within 24 hours. Colonies on PDA reaching 20–25 mm
diam. in 18 days at 25 °C, with dense, floccose, brown
mycelium on the surface, reverse brown to dark brown, with
undulate, pale brown margin.
Fungal Diversity (2020) 105:319–575
503
Fig. 108 Tetraploa scheueri (Material examined: UK, England, Exeter, Exminster marshes, on leaves of Carex acutiformis, 13 November 1988, Ch. Scheuer, GZU 36-91, holotype of Massarina tetraploa; Dried culture specimen of conidial state grown on MEA, derived
from ex-type culture, GZU 32-91). a–f from GZU 36-91. a Ascomata
on the host surface. b Section of ascoma. c Peridium. d Pseudoparaphyses. e Ascus. f Ascospores. g, h from GZU 32-91. g Conidia on
agar medium. h Conidium. Scale bars: a, g = 500 μm, b = 100 μm,
c–f, h = 10 μm
Material examined: CHINA, Yunnan Province, Puzhehei Wetland, 24°8ʹ32ʺN, 104°6ʹ58ʺE, on submerged wood,
12 May 2018, H. Yang, P13 (MFLU 19-0524, holotype),
ex-type living culture MFLUCC 19-0319; Thailand, Chiang Mai Province, on submerged wood, 9 February 2018,
G.N. Wang, A66 (MFLU 18-0959), living culture MFLUCC
18-0652.
Notes: Tetraploa yunnanensis clusters with a strain of T.
sasicola (NCYUCC 19-0371) with high bootstrap support
(Fig. 110). Tetraploa sasicola NCYUCC 19-0371 (MFLU
19-2698) is an ascomycetous species characterized by hyaline to yellowish brown ascospores without appendages
(Hyde et al. 2020b). Although asexual morph of NCYUCC
19-0371 was not formed in culture (Hyde et al. 2020b), phylogeny clearly separates them as distinct species (Fig. 110).
We found that NCYUCC 19-0371 clusters in a separated
clade between Tetraploa yunnanensis (MFLUCC 19-0319,
ex-type strain) and Tetraploa sasicola (KT 563, ex-type
strain) with high bootstrap support (Fig. 110). Comparison
of single LSU and ITS sequence data also shows 14 and 25
nucleotide differences between NCYUCC 19-0371 and KT
563, respectively. This indicates that NCYUCC 19-0371 and
KT 563 are phylogenetically different species (Jeewon and
Hyde 2016).
NCYUCC 19-0371 differs from KT 563 in having larger
asci (97–109 × 22–23 μm vs. 61–100 × 9–13 μm) and wider
ascospores (24–29 × 4–14 μm vs. 22.5–34 × 3–5 μm). Additionally, NCYUCC 19-0371 has hyaline to yellowish brown
ascospores without appendages, while KT 563 has hyaline
ascospores with a narrow sheath drawn out at both ends,
which is longer at the apex than the one at the base (Tanaka
et al. 2009; Hyde et al. 2020b). NCYUCC 19-0371 bearing
the name Tetraploa sasicola is probably a noval species, but
the introduction should be performed with re-examination of
the specimen MFLU 19-2698 or new collectionis.
Key to freshwater Tetraploa species
1. Conidia composed of 4 columns………………………2
1. Conidia composed of 2–3 columns……………………4
2. Conidia > 39 μm long on average, 30–51 × 15–26
μm………………………………………………T. ellisii
2. Conidia mostly < 39 μm long…………………………3
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Fungal Diversity (2020) 105:319–575
Fig. 109 Tetraploa yunnanensis (MFLU 19-0524, holotype). a, b Colonies on natural substrate. c–h Conidia. i Germinated conidium. j, k
Colony on PDA (up-front, down-reverse). Scale bars: c–i = 20 μm
3. Conidial appendages 263–350 µm long, 17–22-septate…
………………………………………………T. scheueri
3. Conidial appendages 12–80 µm long, 1–6-septate…
………………………………………………T. aristata
4. Conidia < 15 µm wide………………………T. abortiva
4. Conidia > 15 µm wide…………………………………5
5. Conidial appendages 25–83 µm long…T. yunnanensis
5. Conidial appendages 3–27 µm long…T. puzheheiensis
13
Triplosphaeria Kaz. Tanaka & K. Hiray., Stud. Mycol. 64:
185 (2009)
Saprobic on decaying or submerged bamboo in freshwater or terrestrial habitats. Sexual morph: Ascomata scattered
to gregarious, immersed under the epidermis, subglobose,
uniloculate, glabrous, with or without a short neck, periphysate. Peridium rim-like at sides, composed of vertically
orientated rectangular to cylindrical, hyaline, hyphoid cells,
poorly developed at the base. Pseudoparaphyses numerous,
trabeculate, narrow, hyaline, indistinctly septate, branched.
Fungal Diversity (2020) 105:319–575
505
Fig. 110 Phylogram generated from maximum likelihood analysis of combined
LSU, SSU, ITS, TUB and TEF
sequence data for species of
Tetraplosphaeriaceae. Bootstrap
values for maximum likelihood
equal to or greater than 75% and
Bayesian posterior probabilities
equal to or greater than 0.95
are placed near the branches as
ML/BYPP. Newly generated
sequences are in red and ex-type
strains are in bold. The new
species introduced in this study
are indicated with underline.
Freshwater strains are indicated
with a red letter “F”. The tree
is rooted to Muritestudina chiangraiensis MFLUCC 17-2551
(Testudinaceae)
Ernakulamia tanakaii NFCCI 4615
Ernakulamia tanakaii NFCCI 4617
Ernakulamia tanakaii NFCCI 4616
Ernakulamia xishuangbannaensis KUMCC 17-0187
Ernakulamia cochinensis PRC 3992
Ernakulamia cochinensis MFLUCC 18-1237 F
Ernakulamia krabiensis MFLUCC 18-0237
99/1.00
87/0.97
100/1.00
Ernakulamia
100/1.00
Shrungabeeja longiappendiculata BCC 76463
Shrungabeeja longiappendiculata BCC 76464
Shrungabeeja aquatica MFLUCC 18-0664 F
Shrungabeeja vadirajensis MFLUCC17-2362 F
Polyplosphaeria thailandica MFLUCC 15-0840
Polyplosphaeria fusca KT 1686
86/-Polyplosphaeria fusca KT 1043
100/1.00
Polyplosphaeria fusca KT 1640
Polyplosphaeria fusca KT 2124
81/0.99
96/1.00 Polyplosphaeria fusca KT 1616
100/1.00
Polyplosphaeria pandanicola MFLUCC 17-2266
Polyplosphaeria nabanheensis KUMCC 16-0151
94/1.00
Quadricrura septentrionalis yone 176
Quadricrura septentrionalis yone 179
Quadricrura septentrionalis yone 44
Quadricrura
septentrionalis KT 920
100/1.00
Quadricrura septentrionalis HC 4983
Quadricrura septentrionalis HC 4984
84/0.96
Quadricrura bicornis yong 153
100/1.00
Quadricrura meridionalis KT 2607
Triplosphaeria cylindrica KT 1256
81/0.95
Triplosphaeria cylindrica KT 1800
Triplosphaeria cylindrica KT 2550
100/0.98
Triplosphaeria
yezoensis KT 1715
100/-Triplosphaeria yezoensis KT 1732
Triplosphaeria acuta KT 1170 F
Triplosphaeria maxima KT 870
99/0.99
Triplosphaeria sp. HC 4665
100/1.00 Triplosphaeria sp. KT 2546
Pseudotetraploa curviappendiculata KT 2558
Pseudotetraploa curviappendiculata HC 4932
Pseudotetraploa curviappendiculata HC 4930
Pseudotetraploa rajmachiensis NFCCI 4619
100/1.00
86/1.00
Pseudotetraploa rajmachiensis NFCCI 4618
Pseudotetraploa
rajmachiensis NFCCI 4620
79/0.99
Pseudotetraploa javanica HC 4934
Pseudotetraploa longissima HC 4933
100/0.99 Tetraploa yunnanensis MFLUCC 18-0652 F
90/0.99
Tetraploa yunnanensis MFLUCC 19-0319 F
85/-Tetraploa sasicola NCYUCC 19-0371
81/0.99
Tetraploa sasicola KT 563
Tetraploa sp. 1 KT 1684
91/1.00
Shrungabeeja
Polyplosphaeria
Quadricrura
Triplosphaeria
95/1.00
Tetraploa dwibahubeeja NFCCI 4623
Tetraploa dwibahubeeja NFCCI 4621
Tetraploa dwibahubeeja NFCCI 4622
Tetraploa pseudoaristata NFCCI 4625
100/1.00
Tetraploa pseudoaristata NFCCI 4624
Tetraploa pseudoaristata NFCCI 4626
Tetraploa sp. CY112
100/1.00
Tetraploa
sp. CBS 996.70
100/1.00
Tetraploa puzheheiensis KUMCC 20–0151 F
Tetraploa yakushimensis KT 1906
Tetraploa thrayabahubeeja NFCCI 4628
100/1.00
Tetraploa thrayabahubeeja NFCCI 4627
Tetraploa thrayabahubeeja NFCCI 4629
Tetraploa sp. 2 KT 2578
Tetraploa nagasakiensis KT 1682
Muritestudina chiangraiensis MFLUCC 17-2551
Pseudotetraploa
100/1.00
Tetraploa
0.03
Asci bitunicate, fissitunicate, cylindrical to clavate, with
rounded apex and short pedicellate. Ascospores 8-spored,
fusiform, with rounded ends, 1-septate, hyaline, smooth,
with an entire sheath (Tanaka et al. 2009). Asexual morph:
Hyphomycetous. Conidiophores absent. Conidiogenous cells
monoblastic. Conidia ovoid to obpyriform, brown, verrucose
at the base, composed of three columns, with three setose
appendages at the apex (Tanaka et al. 2009).
Type species: Triplosphaeria maxima Kaz. Tanaka & K.
Hiray.
Notes: Triplosphaeria was introduced for massarina-like
ascomycetes and tetraploa-like asexual morphs (Tanaka
et al. 2009). Triplosphaeria is similar to Tetraploa in overall morphology of conidia, but differs by its hemispherical ascomata with flattened base (Tanaka et al. 2009; Hyde
et al. 2013). Phylogeny clearly separates them as two distinct
genera (Tanaka et al. 2009; Hyde et al. 2020b; this study,
Fig. 110). Triplosphaeria acuta Kaz. Tanaka & K. Hiray. is
the only species collected from freshwater habitats (Tanaka
et al. 2009).
13
506
List of freshwater Triplosphaeria species
*Triplosphaeria acuta Kaz. Tanaka & K. Hiray., Stud.
Mycol. 64: 186 (2009)
Freshwater distribution: Japan (Tanaka et al. 2009)
Torulaceae Corda, Deutschl. Fl., 3 Abt. (Pilze Deutschl.)
2: 71 (1829)
Key to freshwater genera of Torulaceae
1. Conidiogenous cells holoblastic………………Torula
1. Conidiogenous cells enteroblastic……………………2
2. Conidia pyriform to rostrate, narrowly cylindrical and
obtuse at the apex………………………Rostriconidium
2. Conidia without above characters………………………3
3. Conidia cylindrical or obclavate, solitary or in short
chains……………………………………Dendryphion
3. Conidia clavate to subcylindrical, branch chains………
………………………………………………Neotorula
Dendryphion Wallr., Fl. crypt. Germ. (Norimbergae) 2: 300
(1833)
Saprobic on dead stems of herbaceous plants and decaying wood or submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, hairy or velvety, dark grey, olive, reddish brown or black. Mycelium
immersed. Stroma when present immersed or partly superficial. Conidiophores macronematous, mononematous, usually branched at the apex forming a stipe and head; stipe
straight or flexuous, usually stout, erect, brown to black,
smooth or with the upper part verruculose; branches usually paler, smooth or verruculose. Conidiogenous cells
monotretic or polytretic, sympodial, integrated or discrete,
terminal and intercalary on branches, clavate, cylindrical or
doliiform, cicatrized; scars usually large and dark. Conidia
acropleurogenous, catenate or solitary, dry, cylindrical with
rounded ends or obclavate, sometimes cheiroid, straight or
somewhat curved, pale to mid brown or olivaceous brown,
multi-septate, simple or branched, smooth or verrucose,
thin-walled (Ellis 1971; Su et al. 2016b).
Type species: Dendryphion comosum Wallr.
Notes: The hyphomycetous genus Dendryphion was
placed in Torulaceae based on D. europaeum Crous & R.K.
Schumach. (Crous et al. 2015a). Dendryphion formed a sister clade to Neotorula Ariyaw. et al., but differs in having
cylindrical or obclavate conidia without branched chains
(Hyde et al. 2016b; Su et al. 2016b, 2018). Five Dendryphion species were confirmed in Torulaceae with molecular
data, four of which were collected from freshwater habitats
in China (see list below). They formed a well-supported
monophyletic clade in Su et al. (2018). However, the
sequence data of the type species D. comosum is lacking.
13
Fungal Diversity (2020) 105:319–575
All freshwater Dendryphion species have similar size of conidiophores and conidia, except D. nanum (Nees) S. Hughes
which has the shortest conidiophores and longest conidia (Su
et al. 2016b, 2018). The length and width of conidiophores
can distinguish the other three species.
List of freshwater Dendryphion species
*Dendryphion aquaticum H.Y. Su & K.D. Hyde, Fungal
Diversity (2016); Fig. 111p–v
Freshwater distribution: China (Su et al. 2016b)
*Dendryphion fluminicola Z.L. Luo, D.J. Bhat & K.D.
Hyde, Mycol. Progr. 17(5): 533 (2018)
Freshwater distribution: China (Su et al. 2018)
*Dendryphion nanum (Nees) S. Hughes, Can. J. Bot. 36:
761 (1958); Fig. 111a–g
Basionym: Helminthosporium nanum Nees [as ‘Helmisporium’], Syst. Pilze (Würzburg): 67 (1816) [1816–17]
Freshwater distribution: China (Su et al. 2016b)
*Dendryphion submersum H.Y. Su & K.D. Hyde, Fungal
Diversity (2016); Fig. 111h–o
Freshwater distribution: China (Su et al. 2016b)
Key to freshwater Dendryphion species
1.
1.
2.
2.
3.
3.
Conidiophores < 100 μm long………………D. nanum
Conidiophores > 100 μm long………………………2
Conidiophores < 200 μm long…………D. fluminicola
Conidiophores > 200 μm long…………………………3
Conidiophores 7.5–11.5 μm wide………D. aquaticum
Conidiophores 3.5–4.5 μm wide………D. submersum
Neotorula Ariyaw., Z.L. Luo & K.D. Hyde, Fungal Diversity 80: 393 (2016)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies on the substratum
superficial, effuse, gregarious, hairy, dark brown. Mycelium
immersed, composed of septate, branched, thin-walled,
smooth, pale brown hyphae. Conidiophores macronematous, mononematous, erect, cylindrical, usually with short
branches near the apex, straight or flexuous, dark brown,
septate, smooth. Conidiogenous cells tretic, with a distinct
pore, integrated, terminal, doliiform or lageniform, subhyaline to pale brown. Conidia acrogenous, in branched chains,
dry, clavate to subcylindrical, straight, pale green to brown,
septate, verruculose, thin-walled (Su et al. 2016b).
Type species: Neotorula aquatica Z.L. Luo & K.D. Hyde
Notes: Neotorula was introduced to accommodate a
torula-like species N. aquatica collected from freshwater
habitats in China (Su et al. 2016b). Neotorula shares similar
Fungal Diversity (2020) 105:319–575
507
Fig. 111 Dendryphion spp. (Material examined: CHINA, Yunnan
Province, Dali, Nanjian, saprobic on decaying wood submerged in
stream, April 2014, H.Y. Su, S-059, HKAS 83997, holotype; WanHua stream, saprobic on submerged decaying wood, May 2014, H.Y.
Su, S-125, HKAS 84012). a–g D. nanum (HKAS 84012). a Conidiophores with conidia. b Conidiogenous cells and conidia. c–g Conidia.
h–o D. submersum (HKAS 83998). h, i Conidiophores. j Conidiogenous cells. k–o Conidia. p–v D. aquaticum (HKAS 83997). p
Conidiophores with conidia. q Conidiogenous cells and conidia. r–v
Conidia. Scale bars: a, p = 100 μm, b, c, i, j, q = 50 μm, d–g = 30
μm, h = 150 μm, k–o, r, s = 10 μm, t–v = 15 μm
characters with Torula Pers. in having clavate to subcylindrical, brown conidia forming in branched chains. However, the
former has macronematous and septate conidiophores, while
the latter has micronematous conidiophores reduced to conidiogenous cells or with one brown supporting cell (Su et al.
2016b). The placement of Neotorula was confirmed in Torulaceae based on sequence data of two freshwater species, N.
aquatica and N. submersa Z.L. Luo et al. (Hyde et al. 2016b;
Su et al. 2016b). Both species can be distinguished by the
length of conidia (see key below).
List of freshwater Neotorula species
*Neotorula aquatica Z.L. Luo & K.D. Hyde, Fungal Diversity (2016); Fig. 112a–d
Freshwater distribution: China (Su et al. 2016b)
13
508
Fungal Diversity (2020) 105:319–575
Fig. 112 Neotorula spp. (Material examined: CHINA, Yunnan Province, Dali, Cangshan Mountain, Lingquan stream, on submerged
decaying wood, March 2014, H.Y. Su, S-087, HKAS 84001, holotype; ibid., Dulong River, on submerged decaying wood, May 2015,
X.C. Tao, HD 1–10–7, HKAS 92660). a–d N. aquatica (HKAS
84001). a Conidiophores with conidia. b–d Conidia. e–k N. submersa (HKAS 92660). e Colonies on the substratum. f, g Conidiophores with conidia. h Conidiogenous cells with conidia. i–k
Conidia. Scale bars: a, c, f, g = 20 μm, b = 30 μm, d, k = 10 μm, h–j
= 15 μm
*Neotorula submersa Z.L. Luo, H.Y. Su & K.D. Hyde, Fungal Diversity (2016); Fig. 112e–k
Freshwater distribution: China (Hyde et al. 2016b)
subcylindrical, straight or flexuous, dark brown to black,
paler at the apex, septate, smooth. Conidiogenous cells
monotretic or polytretic, integrated, terminal, cylindrical,
dark brown. Conidia solitary, dry, pyriform to rostrate,
narrowly cylindrical and obtuse at the apex, dark brown to
black, with a thick, black, truncate basal scar and a pale cell
above the scar (Su et al. 2018).
Type species: Rostriconidium aquaticum Z.L. Luo, K.D.
Hyde & H.Y. Su
Notes: Rostriconidium was introduced for a freshwater
species R. aquaticum collected from China (Su et al. 2018).
Rostriconidium nested in Torulaceae with high bootstrap
support (Su et al. 2018; Tibpromma et al. 2018). Rostriconidium is similar to Sporidesmioides Jun F. Li et al. in
having macronematous, dark brown to black conidiophores
and dark brown to black, similar shape of ascospores with
cylindrical, narrower apex (Li et al. 2016b; Su et al. 2018).
Key to freshwater Neotorula species
1. Conidia 11–16 μm long……………………N. aquatica
1. Conidia 16–22 μm long……………………N. submersa
Rostriconidium Z.L. Luo, K.D. Hyde & H.Y. Su, Mycol.
Progr. 17(5): 536 (2018)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, scattered,
hairy, brown. Mycelium mostly immersed, composed
of pale brown, septate, unbranched hyphae. Conidiophores macronematous, mononematous, single, erect,
13
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509
However, the former has tretic conidiogenous cells, the latter
having blastic. Phylogeny clearly separates them as distinct
genera (Tibpromma et al. 2018).
3. Conidia in branched chains…………………T. masonii
3. Conidia not in chains…………………T. gaodangensis
List of freshwater Rostriconidium species
*Rostriconidium aquaticum Z.L. Luo, K.D. Hyde & H.Y.
Su, Mycol. Progr. 17(5): 536 (2018)
Freshwater distribution: China (Su et al. 2018)
Trematosphaeriaceae K.D. Hyde, Y. Zhang ter, Suetrong
& E.B.G. Jones, Fungal Diversity 63: 254 (2013)
Torula Pers., Ann. Bot. (Usteri) 15: 25 (1795)
Saprobic in freshwater or terrestrial habitats. Sexual
morph: Undetermined. Asexual morph: Colonies effuse,
velvety, dark brown to black. Mycelium mostly immersed.
Conidiophores micronematous, reduced to conidiogenous
cells, or with one supporting cell. Conidiogenous cells
mono- to polyblastic, solitary on mycelium, doliiform to
ellipsoid or clavate, cupulate, brown, smooth to verruculose. Conidia acrogenous, dry, brown, branched chains,
phragmosporous, strongly constricted at the septa, smooth
to verrucose; conidiogenous cell and fertile cell in conidial
chains (where branching occurs) darker brown than other
cells (Crous et al. 2015a).
Type species: Torula herbarum (Pers.) Link
Notes: Crous et al. (2015a) designated a neotype (CPC
24114) for the type species Torula herbarum. Torula has
over 500 epithets in Index Fungorum (2020) and recently
introduced species were manily based on molecular data (Su
et al. 2016b, 2018; Hyde et al. 2017, 2019). Four freshwater
species have been confirmed in Torula based on sequence
data (see list below). They can be distinguished by conidial
size and chains.
1.
1.
2.
2.
List of freshwater Torula species
*Torula aquatica Z.L. Luo, K.D. Hyde, X.J. Su & H.Y. Su,
Mycol. Progr. 17(5): 538 (2018)
Freshwater distribution: China (Su et al. 2018)
*Torula fici Crous [as ‘ficus’], IMA Fungus 6(1): 192 (2015)
Freshwater distribution: China (Su et al. 2018)
*Torula gaodangensis J. Yang & K.D. Hyde, Fungal Diversity 87: 113 (2017)
Freshwater distribution: China (Hyde et al. 2017)
*Torula masonii Crous, IMA Fungus 6(1): 195 (2015)
Freshwater distribution: China (Su et al. 2018)
Key to freshwater Torula species
1.
1.
2.
2.
Conidia < 20 μm long…………………………………2
Conidia > 20 μm long…………………………………3
Conidia 8.5–13.5 × 4.5–5.5 μm……………T. aquatica
Conidia 11–23 × 5.5–6.5 μm……………………T. fici
Key to freshwater genera of Trematosphaeriaceae
Ascospores hyaline……………………Falciformispora
Ascospores brown………………………………………2
Ascospores vermiform……………………Hadrospora
Ascospores fusiform…………………Trematosphaeria
Falciformispora K.D. Hyde, Mycol. Res. 96(1): 26 (1992)
Saprobic on submerged wood, intertidal mangrove wood,
dead wood or isolated from mycetoma of humans. Sexual
morph: Ascomata solitary or gregarious, erumpent becoming superficial, subglobose or ovoid, glabrous, black, coriaceous, ostiolate. Peridium two-layered, outer layer composed
of thick-walled angular or rounded brown cells, inner layer
composed of elongate, hyaline cells. Pseudoparaphyses
numerous, cellular, narrowly cylindrical, septate, anastomosing. Asci 8-spored, bitunicate, fissitunicate, clavate, pedicellate, with an ocular chamber. Ascospores bi- to tri-seriate,
fusiform, hyaline, multiseptate, surrounded by a thin mucilage sheath with or without a single scythe-like appendage
(Hyde 1992a). Asexual morph: Undetermined.
Type species: Falciformispora lignatilis K.D. Hyde
Notes: Falciformispora was introduced to accommodate
F. lignatilis collected from intertidal mangrove wood (Hyde
1992a). Falciformispora is characterized by fusiform, hyaline,
multiseptate ascospores surrounded by a mucilaginous sheath
and appendage (Hyde 1992a). Falciformispora lignatilis was
later collected from freshwater habitats in USA (Raja et al.
2009b). It differs from another freshwater species F. aquatica
D.F. Bao et al. in having smaller asci (110–136 × 20–32 µm vs.
115–167 × 28–47 µm) and ascospores (42–50 × 7.5–10 µm vs.
48–59 × 10–16 µm) (Hyde et al. 2019). Both F. aquatica and F.
lignatilis have a single scythe-like appendage, while F. uttaraditense Boonmee et al. lacks this character (Hyde et al. 2020b).
Falciformispora uttaraditense was introduced based solely on
ITS sequence data and morphology (Hyde et al. 2020b).
List of freshwater Falciformispora species
*Falciformispora aquatica D.F. Bao, K.D. Hyde & H.Y. Su,
Fungal Diversity 96: 73 (2019)
Freshwater distribution: Thailand (Hyde et al. 2019)
*Falciformispora lignatilis K.D. Hyde, Mycol. Res. 96(1):
27 (1992)
Freshwater distribution: USA (Raja et al. 2009b)
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Fungal Diversity (2020) 105:319–575
*Falciformispora uttaraditensis Boonmee & Huanraluek,
Fungal Diversity 100: 127 (2020)
Freshwater distribution: Thailand (Hyde et al. 2020b)
Key to freshwater Falciformispora species
1. Ascospores without a single scythe-like appendage……
…………………………………………F. uttaraditense
1. Ascospores with a single scythe-like appendage………2
2. Ascospores 48–59 × 10–16 µm……………F. aquatica
2. Ascospores 42–50 × 7.5–10 µm……………F. lignatilis
Hadrospora Boise, Mem. N. Y. bot. Gdn 49: 310 (1989)
Saprobic on woody or monocotyledonous hosts in terrestrial or submerged wood in freshwater habitats. Sexual
morph: Ascomata solitary to scattered, immersed to semiimmersed, erumpent, with black dots on the host surface,
uniloculate, globose to subglobose, dark brown to black,
glabrous, ostiolate, with a minute papilla. Peridium thinwalled, composed of black, broad pseudoparenchymatous
cells, arranged in a textura angularis. Pseudoparaphyses
numerous, trabeculate, filamentous, distinctly septate, frequently anastomosing. Asci 8-spored, bitunicate, fissitunicate, ovoid to ampulliform, sessile to minutely pedicellate,
apically rounded, with indistinct ocular chamber. Ascospores
uni- to tri-seriate, broadly fusiform, vermiform, brown to
reddish-brown, paler at the end cells, phragmosporous, with
or without sheath (Tanaka and Harada 2003a; Phookamsak
et al. 2014). Asexual morph: zalerion-like. Hyphomycetous. Conidia holoblastic, helicoid, multiseptate, constricted
at septa, brown to dark brown, smooth (Tanaka and Harada
2003a).
Type species: Hadrospora fallax (Mouton) Boise
Notes: Hadrospora was introduced to accommodate
two species H. clarkii (Sivan.) Boise and H. fallax, which
were transferred from Trematosphaetria (Boise 1989). The
familial placement of Hadrospora is ambiguous due to the
dearth of sequences. Hadrospora was accepted in Phaeosphaeriaceae (Hyde et al. 2013), but later suggested in
Trematosphaeriaceae based on morphology (Phookamsak
et al. 2014). Hadrospora has ovoid to ampulliform asci and
hyphomycetous asexual morph (Tanaka and Harada 2003a;
Phookamsak et al. 2014). Phaeosphaeriaceae members
mostly have cylindrical to cylindri-clavate asci and coelomycetous asexual morphs (Hyde et al. 2013). We place
Hadrospora in Trematosphaeriaceae following Phookamsak et al. (2014) and Wijayawardene et al. (2018, 2020),
until sequence data of H. fallax are obtained to confirm its
familial placement.
The ascospore size of Hadrospora fallax in the holotype (BR5 020009617141) are 55–80 × 16–22 µm (Zhang
et al. 2012b; Phookamsak et al. 2014). Tanaka and Harada
13
Fig. 113 Hadrospora fallax (redrawn from Phookamsak et al.
(2014), BPI 878936, BR 5020009617141, holotype). a–c from BPI
878936. d from BR 5020009617141, holotype. a Section through
ascoma. b Ascus. c, d Ascospores. Scale bars: a = 100 μm, b = 25
μm, c, d = 30 μm
(2003a) described the ascospores as larger, 63.5–104.5 ×
17.5–29 µm, with a conspicuous sharply delimited sheath
which was not mentioned in the holotype. Hadrospora fallax produced a zalerion-like asexual morph in PDA culture
(Tanaka and Harada 2003a).
List of freshwater Hadrospora species
Hadrospora fallax (Mouton) Boise, Mem. N. Y. bot. Gdn
49: 310 (1989); Fig. 113
Basionym: Trematosphaeria fallax Mouton, Bull. Soc. R.
Bot. Belg. 25(no. 1): 155 (1886)
Freshwater distribution: China (Luo et al. 2004)
Trematosphaeria Fuckel, Jb. nassau. Ver. Naturk. 23–24:
161 (1870) [1869–70]
Saorobic on lignocellulosic material in mangroves and
terrestrial or submerged wood in freshwater habitats. Sexual
morph: Ascomata solitary or clustered in groups, immersed,
becoming erumpent to semi-immersed, unilocular, subglobose, black, coriaceous, with ostiolate papilla. Peridium
uneven in thickness, a single layer, composed of small,
heavily pigmented, thick-walled cells of textura angularis.
Pseudoparaphyses dense, cellular, septate, embedded in
mucilage, branching and anastomosing between and above
the asci. Asci 8-spored, bitunicate, fissitunicate, clavate,
with a short, thick, furcate pedicellate, with a truncate ocular chamber. Ascospores uni- to bi-seriate, fusiform, dark
brown, trans-septate, secondary septum forming late or
Fungal Diversity (2020) 105:319–575
often absent, deeply constricted at the median septum, variously ornamented (Zhang et al. 2008b; Hyde et al. 2013).
Asexual morph: Undetermined. Hyphopodia-like structures
were produced in culture of Trematosphaeria pertusa (CBS
122368, ex-epitype) (Zhang et al. 2008b).
Type species: Trematosphaeria pertusa Fuckel
Notes: Trematosphaeria is an old genus and comprises
over 200 epithets in Index Fungorum (2020). The type species T. pertusa formed a robust phylogenetic clade with Falciformispora lignatilis in Trematosphaeriaceae (Suetrong
et al. 2009, 2011b; Zhang et al. 2009c; Hyde et al. 2013).
Two strains of T. hydrela (Rehm) Sacc. (CBS 880.70 and
HKUCC 10666) clustered in Lindgomycetaceae, which was
phylogenetically distant from Trematosphaeriaceae (Tsang
et al. 2014; Hyde et al. 2017; this study, Fig. 2). Tsang et al.
(2014) thought that these strains were likely to be misnamed.
The two strains need verification to confirm if they are misnamed or should be transferred T. hydrela to Lindgomycetaceae. We place T. hydrela in Trematosphaeriaceae for
now. Four freshwater Trematosphaeria species can be distinguished by septation and dimension of ascospores.
List of freshwater Trematosphaeria species
Trematosphaeria confusa (Garov.) Boise & D. Hawksw.,
Mycologia 77(2): 232 (1985)
Basionym: Verrucaria confusa Garov., Tentam. Dispos.
Lich. Langob.: 77 (1865)
Synonymy: Didymosphaeria confusa (Garov.) Rehm,
Hedwigia 18: 165 (1879)
Freshwater distribution: Australia (Hyde 1995c), Malaysia (Ho et al. 2001)
*Trematosphaeria hydrela (Rehm) Sacc., Syll. fung. (Abellini) 2: 117 (1883)
Basionym: Melanomma hydrelum Rehm, Hedwigia 21(6):
82 (1882)
Synonymy: Caryospora callicarpa Sacc., Michelia 1(no.
3): 331 (1878)
Melanomma callicarpum Berl., Icon. fung. (Abellini)
1(1): 37 (1890)
Trematosphaeria callicarpa (Sacc.) Sacc., Syll. fung.
(Abellini) 2: 116 (1883)
Freshwater distribution: Union of Soviet Socialist Republics (Petrak 1925)
Trematosphaeria lineolatispora K.D. Hyde, Mycol. Res.
96(1): 28 (1992)
Freshwater distribution: USA (Raja et al. 2009b)
*Trematosphaeria pertusa (Pers.:Fr.) Fuckel, Jb. nassau.
Ver. Naturk. 23–24: 161 (1870) [1869–70]
511
Basionym: Sphaeria pertusa Pers., Syn. meth. fung. (Göttingen) 1: 83 (1801)
Synonymy: see Index Fungorum (2020)
Freshwater distribution: England (Eaton and Jones 1971;
Eaton 1972; Kane et al. 2002), Hungary (Révay and Gönczöl
1990), Japan (Tanaka et al. 2015)
Key to freshwater Trematosphaeria species (updated from
(Tanaka et al. 2005a))
1. Ascospores 1–3-septate………………………………2
1. Ascospores 3–7-septate………………………………3
2. Ascospores (30–)34–45(–52) × (8–)8.5–11 μm………
………………………………………………T. confusa
2. Ascospores (21–)22–32(–34) × 6–10 μm………………
………………………………………………T. pertusa
3. Ascospores > 70 μm long……………………T. hydrela
3. Ascospores < 50 μm long……………T. lineolatispora
Wicklowiaceae Ariyaw. & K.D. Hyde, Fungal Diversity 75:
126 (2015)
Wicklowia Raja, A. Ferrer & Shearer, Mycoscience 51(3):
210 (2010)
Saprobic on submerged wood. Sexual morph: Ascomata
solitary to gregarious, immersed to erumpent, uniloculate,
subglobose, flattened dorsiventrally, glabrous, dark brown
to black, ostiolate, visible as a black oval to circular, shallow, crater-like depression on the substrate. Peridium thin,
composed of several layers of small pseudoparenchymatic
cells. Pseudoparaphyses sparse or numerous, cellular, hyaline, septate, embedded in a gel matrix. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate to broadly clavate,
sessile or short pedicellate, rounded at the apex, with an
ocular chamber when immature. Ascospores bi- to tri-seriate, ellipsoidal-oblong, rounded at the apices, hyaline, 1-septate, asymmetrical, smooth, thin-walled, surrounded by a
distinctly mucilaginous sheath (Raja et al. 2010a). Asexual
morph: Undetermined.
Type species: Wicklowia aquatica Raja, A. Ferrer &
Shearer
Notes: Wicklowia was introduced to accommodate the
freshwater species W. aquatica isolated from submerged
wood in Costa Rica and Florida (Raja et al. 2010a). The
second species W. submersa Boonmee et al. was collected
from submerged wood in Thailand (Boonmee et al. 2019).
Both species share similar morphological characters in the
shape and size of ascomata, asci and ascospores (Raja et al.
2010a; Boonmee et al. 2019). They can be distinguished by
the appendages and sheaths of the ascospores.
List of freshwater Wicklowia species
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512
Fungal Diversity (2020) 105:319–575
Fig. 114 Wicklowia aquatica
(Material examined: USA,
Florida, Apalachicola National
Forest, on submerged decorticated woody debris, 14 January
2006, H.A. Raja & J.L. Crane,
F76-2, ILL 40790, holotype;
Heredia, La Selva stream, on
submerged wood, 9 January
2006, M. Salazar, AF289-4).
a–d ILL 40790. a Ascomata on
wood. b Longitudinal section
through ascoma. c, d Bitunicate
asci. e, f AF289-4. Ascospores
showing gelatinous sheath and
basal appendages. Scale bars: a
= 100 µm, b, e, f = 10 µm, c, d
= 20 µm
*Wicklowia aquatica Raja, A. Ferrer & Shearer, Mycoscience 51: 211 (2010); Fig. 114
Freshwater distribution: USA (Raja et al. 2010a)
*Wicklowia submersa Boonmee, Sorvongxay & K.D. Hyde,
Phytotaxa 411: 76 (2019)
Freshwater distribution: Thailand (Boonmee et al. 2019)
Key to freshwater Wicklowia species
1. Ascospores surrounded by a gelatinous sheath, with filamentous appendages………………………W. aquatica
1. Ascospores surrounded by a distinctly mucilaginous
sheath, without any appendages…………W. submersa
Pleosporales genera incertae sedis
In this section, we list freshwater genera belonging to
Pleosporales genera incertae sedis (Wijayawardene et al.
2018, 2020). The familial placements need to be clarified
with more fresh collections and sequence data.
13
Ascorhombispora L. Cai & K.D. Hyde, Cryptog. Mycol.
28: 294 (2007)
Saprobic on submerged bamboo. Sexual morph: Ascomata solitary or gregarious, superficial, uniloculate, globose
to subglobose, dark brown to black, with ostiolate papilla.
Peridium thin, comprising two layers of relatively thinwalled cells of textura angularis. Pseudoparaphyses absent.
Asci 8-spored, bitunicate, obpyriform, broadly clavate to
saccate, pedicellate, with rounded apex, deliquescent at
maturity. Ascospores overlapping bi- to tri-seriate, broadly
fusiform to rhombic, straight, 3-euseptate, not constricted
at septa, median septum wide, forming a darker band, central cells large, trapezoid, dark brown to black, verruculose,
thick-walled, with bipolar, small, hemispherical, subhyaline to light brown cells, surrounded by gelatinous mucilaginous sheath (Cai and Hyde 2007b). Asexual morph:
Undetermined.
Type species: Ascorhombispora aquatica L. Cai & K. D.
Hyde
Notes: The monotypic genus Ascorhombispora is typified
by A. aquatica L. Cai & K. D. Hyde (Cai and Hyde 2007b).
Fungal Diversity (2020) 105:319–575
The genus is characterized by globose to subglobose ascomata, obpyriform, broadly clavate to saccate, deliquescent
asci without a pseudoparaphyses, and rhombic, 3-euseptate,
dematiaceous ascospores with smaller, subhyaline, bipolar
cells (Cai and Hyde 2007b). Ascorhombispora resembles
Caryospora in having superficial ascomata, saccate asci
and large dematiaceous ascospores with sheath, whereas
the latter has larger, carbonaceous ascomata, numerous
pseudoparaphyses, fusiform to broad-ellipsoid or biconic
ascospores without subhyaline, bipolar cells (Ariyawansa
et al. 2015). Phylogenetic analyses clearly separated them
(Cai and Hyde 2007b; Shearer et al. 2009). Ascorhombispora is morphologically and phylogenetically not conclusively associated with any families in Pleosporales (Shearer
et al. 2009). We re-examined the holotype of Ascorhombispora aquatica and did not find the verrucose ascospores
mentioned in Cai and Hyde (2007b), a feature probably obvious only in fresh specimen.
List of freshwater Ascorhombispora species
*Ascorhombispora aquatica L. Cai & K. D. Hyde, Cryptog.
Mycol. 28: 295 (2007); Fig. 115
Freshwater distribution: China (Cai and Hyde 2007b)
Fusiformiseptata W. Dong, H. Zhang & K.D. Hyde, gen.
nov.
Index Fungorum number: IF557938; Facesoffungi number: FoF09283
Etymology: referring to fusiform spores of this genus
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata black, scattered, immersed with a
neck erumpent through host surface, ellipsoidal or obovoid,
coriaceous, ostiolate. Neck subcylindrical or irregular, with
flattened top, thick-walled, composed of two-layered peridium, with orange pigment. Peridium comprising several layers of dark brown, thick-walled, compressed cells of textura
epidermoidea or irregular cells. Pseudoparaphyses numerous, cellular, broadly filiform, hyaline, septate, branched.
Asci 8-spored, bitunicate, clavate, short pedicellate, apically
rounded with well-developed ocular chamber. Ascospores
biseriate, fusiform, hyaline, septate, asymmetrical, thinwalled. Asexual morph: Undetermined.
Notes: Multigene phylogenetic analysis shows that our
new collection MFLU 18-1721 forms an independent lineage in Pleosporales (Fig. 2). Fusiformiseptata is therefore
introduced for this freshwater taxon. Fusiformiseptata has a
prominent neck with orange pigment on the inner wall and
forms an arrow-shaped base to the ascomata. These characters are special in Pleosporales and warrant a new genus.
List of freshwater Fusiformiseptata species
513
*Fusiformiseptata crocea W. Dong, H. Zhang & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557939; Facesoffungi number: FoF09284; Fig. 116
Etymology: “crocea” in reference to the orange pigment
in the neck
Holotype: MFLU 18-1721
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Ascomata 255–280 μm high, 190–220 μm
wide, black, scattered, immersed with a neck erumpent
through host surface, uniloculate, ellipsoidal or obovoid,
arrow-shaped at the basal region, coriaceous, ostiolate.
Neck 230–260 μm high, 270–290 μm wide, 75–95 μm thick,
prominent, subcylindrical or irregular, with flattened top,
thick-walled, composed of two layers, with outer wall comprising several layers of thick-walled, dark brown to black
cells of textura epidermoidea, inner wall comprising thinwalled, hyaline cells of textura epidermoidea with orange
pigment. Peridium 30–70 μm thick, comprising several layers of dark brown, thick-walled, compressed cells of textura
epidermoidea or irregular cells. Pseudoparaphyses 2.5 μm
diam., numerous, cellular, broadly filiform, hyaline, septate,
branched. Asci (85–)95–135(–150) × 16–20(–23) μm (x̄ =
112 × 18.5 μm, n = 15), 8-spored, bitunicate, clavate, short
pedicellate, apically rounded with well-developed ocular
chamber. Ascospores 36–42 × 6–9 μm ( x̄ = 40 × 8 μm,
n = 10), biseriate, fusiform, straight or curved, hyaline,
1-septate when young, 4(–5)-septate when mature, deeply
constricted at the central septum, asymmetrical, upper part
shorter than lower part, guttulate, thin-walled, smooth, with
a mucilaginous sheath, 15–25 μm wide. Asexual morph:
Undetermined.
Culture characteristics: On PDA, colony irregular, reaching 25 mm in 10 days at 25 °C, grey from above, brown
from below, surface rough, mucoid, raised, umbonate, edge
undulate.
Material examined: THAILAND, Songkhla Province,
on submerged wood in a stream, 10 May 2018, W. Dong,
hat1111-1 (MFLU 18-1721, holotype), ex-type living culture MFLUCC 18-1415; ibid., hat1111-2 (HKAS 105059,
isotype), ex-type living culture KUMCC 19-0080.
Isthmosporella Shearer & Crane, Mycologia, 91: 141 (1999)
Saprobic on submerged wood. Sexual morph: Ascomata scattered on natural substrate, immersed to superficial,
globose, oblique, black, glabrous, membranous, rostrate.
Neck short, cylindrical, periphysate. Peridium two-layered,
outer layer composed of brown, fusoid-cylindric pseudoparenchyma cells, inner layer composed of compressed,
subhyaline to light brown, fusoid cells. Pseudoparaphyses
sparse, cellular, broad, hyaline, septate. Asci 8-spored, bitunicate, fissitunicate, ventricose to clavate, broadly rounded
at the apex, without an apical apparatus or oculus, short
13
514
Fungal Diversity (2020) 105:319–575
Fig. 115 Ascorhombispora
aquatica (Material examined:
CHINA, Yunnan Province, Jinghong, on submerged bamboo
in a small forest stream, 26 January 2003, L. Cai, CAI-1H31,
HKU(M) 10859, holotype). a
Ascomata on the host surface. b
Section of ascoma. c Peridium.
d Ascus. e, f Ascospores. Scale
bars: a = 300 µm, b, d = 50 µm,
c, e, f = 30 µm
pedicellate. Ascospores tri- to tetra-seriate, cylindrical to
fusoid, isthmoid at center, tapering at both ends, straight
to slightly curved, hyaline, phragmoseptate, constricted at
septa, surrounded by a gelatinous sheath (Shearer and Crane
1999). Asexual morph: Undetermined.
Type species: Isthmosporella pulchra Shearer & Crane
Notes: The monotypic genus Isthmosporella, typified by
I. pulchra Shearer & Crane, was collected from submerged,
decorticated wood in USA (Shearer and Crane 1999). It was
initially placed in Phaeosphaeriaceae based on its small,
globose ascomata, relatively undifferentiated peridium of
small pseudoparenchymatic cells, and sparse pseudoparaphyses (Shearer and Crane 1999). However, the ventricose to
clavate asci, cylindrical to fusoid ascospores with isthmoid
13
at center do not fit with the familial concept of Phaeosphaeriaceae (Hyde et al. 2013). It is placed in Pleosporales genera incertae sedis (Wijayawardene et al. 2018, 2020), until
its DNA data are obtained. One collection of Isthmosporella
pulchra (A254-2), collected from the type locality by its
author (Dr. Lee Crane), is illustrated here (Fig. 117).
List of freshwater Isthmosporella species
Isthmosporella pulchra Shearer & Crane, 91: 142 (1999);
Fig. 117
Freshwater distribution: USA (Shearer and Crane 1999)
Rebentischia P. Karst., Fungi Fenniae Exsiccati, Fasc. 9:
881 (1869)
Fungal Diversity (2020) 105:319–575
Fig. 116 Fusiformiseptata crocea (MFLU 18-1721, holotype). a
Appearance of ascomata on host surface. b, c Ascomata cut through
horizontal showing the orange filler. d, g Vertical section of necks.
e, f, h Vertical section of ascomata. i–k Bitunicate asci. l Pseudo-
515
paraphyses. m Apex of ascus. n Pedicel of ascus. o–q Ascospores.
r Ascospore in Indian Ink. s Germinated ascospore. t, u Colony on
PDA (left-front, right-reverse). Scale bars: d–f = 100 μm, g = 50 μm,
h, m = 10 μm, i–l, n–s = 20 μm
13
516
Fungal Diversity (2020) 105:319–575
Fig. 117 Isthmosporella pulchra (Material examined: USA,
New York, Adirondack Park,
Piercefield, Tupper Lake at
public boat launch from Rt. 30,
UTM Zone 18, on submerged,
decorticated wood, A254-2).
a Squash mount of ascoma. b
Bitunicate ascus. c, d Ectotunica rupturing at the apex and
becoming strongly wrinkled
and curled inwards. e Pseudoparaphyses. f, h Ascospores.
Note gelatinous sheath with a
dense region near the isthmus
(arrowed in f). g Ascospore in
glycerin. Scale bars: a = 100
µm, b–h = 20 µm
Saprobic on decaying wood in terrestrial or submerged
wood in freshwater habitats. Sexual morph: Ascomata
solitary or clustered, immersed or erumpent to superficial,
uniloculate, globose to subglobose, somewhat depressed, or
ovoid, sometimes collapsed in age, apex blunt, black, coriaceous, ostiolate. Peridium soft, fleshy, often roughened by
protruding cells or short hyphae, composed of pseudoparenchymatous cells, light to dark vinaceous brown externally,
compressed, hyaline or vinaceous-tinged cells inwardly,
pigmented encrust on the surface of outermost cells. Pseudoparaphyses numerous, cellular, flexuous, narrow, hyaline,
septate, branched, anastomosing, persistent. Asci 8-spored,
bitunicate, fissitunicate, cylindric-clavate, short pedicellate,
apically thickened. Ascospores biseriate, elongate-obovoid
or narrowly clavate, rounded at the apex, base tapering to
a hyaline appendage, uaually slightly curved, brownish to
dark brown, transeptate, smooth, thin-walled (Barr 1980;
Boonmee et al. 2014). Asexual morph: Coelomycetous.
asteromella-like (Barr 1980).
Type species: Rebentischia pomiformis P. Karst.
13
Notes: Rebentischia, typified by R. pomiformis, is characterized by darkly pigmented ascomata, elongate-obovoid
ascospores with a base tapering to a hyaline appendage
(Boonmee et al. 2014). Barr (1980) placed Rebentischia in
Tubeufiaceae based on these characters, which was accepted
by Rossman (1987), Lumbsch and Huhndorf (2010) and
Boonmee et al. (2011). However, Boonmee et al. (2014)
placed it in Pleosporales incertae sedis based on the darkened ascospores with a setiform basal appendage after reexamination of the holotype of R. pomiformis. The asexual
morph is unknown with certainty, but probably is asteromella-like fungi that grow near the base of ascomata (Barr
1980).
Fourteen species are listed in Index Fungorum (2020),
all of which are old species without any sequence data. Barr
(1980) thought that there are only two valid entities, viz.
Rebentischia massalongoi (Mont.) Sacc. and R. unicaudata
(Berk. & Broome) Sacc. von Arx and Muller (1975) synonymized the type species R. pomiformis with R. massalongi, which was accepted by Boonmee et al. (2014).
Fungal Diversity (2020) 105:319–575
The freshwater species Rebentischia unicaudata fits well
with the generic concept of Rebentischia by dark ascomata,
clavate asci and narrowly obovoid ascospores with a hyaline basal cell (Barr 1980). It is similar to the type species,
but differs in the habitats (freshwater vs. terrestrial) and
ascospores size (17–25(–30) × 4–6(–7.5) µm vs. 28–40 ×
6–10 µm) (Barr 1980).
517
9.
9.
10.
10.
List of freshwater Rebentischia species
Rebentischia unicaudata (Berk. & Broome) Sacc., Syll.
fung. (Abellini) 2: 12 (1883)
Basionym: Sphaeria unicaudata Berk. & Broome, Ann.
Mag. nat. Hist., Ser. 33: 373 (1859)
Freshwater distribution: USA (Shearer and Crane 1986)
11.
11.
12.
12.
Tubeufiales Boonmee & K.D. Hyde, Fungal Diversity
68(1): 245 (2014)
Tubeufiaceae was well-studied by Lu et al. (2018b) based
on morphology and multigene phylogenetic analyses. We
provide brief generic notes, list and key to freshwater Tubeufiales species. The generic descriptions mostly refers to Lu
et al. (2018b).
13.
13.
14.
14.
Tubeufiaceae M.E. Barr, Mycologia 71(5): 948 (1979)
Key to freshwater helicosporous genera of Tubeufiaceae
(found on natural substrate) (updated from Lu et al.
(2018b))
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
8.
8.
Colonies dematiaceous………………………………2
Colonies hyaline………………………………………7
Conidia subglobose or broadly elliptical………………
………………………………………Pseudohelicoon
Conidia not subglobose or broadly elliptical
body…………………………………………………3
Colonies pale yellow to yellow-green…………………
………………………………………Helicosporium
Colonies pink, pale brown to dark brown……………4
Conidiophores arising from a pseudoparenchymatous
stromata………………………………Helicodochium
Conidiophores arising from creeping hyphae………5
Conidiophores < 50 μm long…………Berkleasmium
Conidiophores > 50 μm long…………………………6
Conidia with truncate basal cell and thickened lateral
walls, dilute fuliginous……………Helicotruncatum
Conidia without truncate basal cell and thickened lateral walls, hyaline to brown………………Helicoma
Conidiophores 0–3-septate……………………………8
Conidiophores multi-septate………………………10
Conidiophores pale brown, conidiogenous cells hyaline
to pale brown……………………Dematiohelicomyces
Conidiophores and conidiogenous cells hyaline……………………………………………………9
Conidiogenous cells tapering towards the apex,
inverted funnel at the apex, conidia not loose in
water…………………………………Helicohyalinum
Conidiogenous cells cylindrical, truncate at the apex,
conidia loosely coiled in water…………Helicomyces
Conidiophores setiferous, brown to dark
brown…………………………………Helicoarctatus
Conidiophores not setiferous, hyaline to
brown………………………………………………11
Conidiogenous cells cylindrical, with a truncate
apex…………………………………………………12
Conidiogenous cells cylindrical, denticulate………13
Conidiophores hyaline, conidia helicoid……
…………………………………………Helicotubeufia
Conidiophores pale brown to brown, conidia variable
in shape………………………………………Tubeufia
Conidiophores hyaline to brown, mostly unbranched,
> 110 μm long………………………Neohelicomyces
Conidiophores pale brown to dark brown, irregular,
20–450 μm long……………………………………14
Conidiophores branched, conidia mostly loosely
coiled……………………………Pseudohelicomyces
Conidiophores occasionally branched, conidia tightly
coiled………………………………Neohelicosporium
Aquaphila Goh, K.D. Hyde & W.H. Ho, Mycol. Res. 102(5):
588 (1998)
Saprobic on submerged wood or fallen leaves. Description of this genus see Goh et al. (1998a), Boonmee et al.
(2014) and Hyde et al. (2016b).
Type species: Aquaphila albicans Goh, K.D. Hyde &
W.H. Ho
Notes: Aquaphila is characterized by translucent or pale
yellowish, chalky white colonies, hyaline conidiophores
arising singly from the procumbent hyphae, denticulate conidiogenous cells with sympodially proliferating and fusoid
to sickle-shaped conidia (Goh et al. 1998a). Boonmee et al.
(2014) linked Tubeufia asiana Sivichai & C.K.M. Tsui to
the sexual morph of the type species A. albicans based on
molecular data. Aquaphila formed a sister clade to Chlamydotubeufia Boonmee & K.D. Hyde (Lu et al. 2018b; this study,
Fig. 129). Aquaphila albicans was only found from freshwater
habitats (Goh et al. 1998a; Hyde et al. 2016b) and confirmed
with molecular data by Hyde et al. (2016b) and this study.
List of freshwater Aquaphila species
*Aquaphila albicans Goh, K.D. Hyde & W.H. Ho, Mycol.
Res. 102(5): 588 (1998); Fig. 118
Freshwater distribution: Australia (Goh et al. 1998a),
Thailand (Hyde et al. 2016b; this study)
Saprobic on decaying wood submerged in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies on
13
518
Fungal Diversity (2020) 105:319–575
Fig. 118 Aquaphila albicans
(MFLU 15-2713). a, b Colonies on natural substrate. c–g
Conidiophores with attached
conidia. h–k Conidia. l Germinated conidium. m, n Colony on
PDA (up-front, down-reverse).
Scale bars: a = 100 μm, b = 50
μm, c, d, f–k = 10 μm, e = 5
μm, l = 20 μm
natural substrate effuse, translucent, chalky white. Mycelium
partly immersed in substrate, consisting of septate, branched,
hyaline, smooth, thin-walled hyphae. Conidiophores up to 20
μm long, 3–6 μm wide, arising singly from procumbent hyphae,
semi-macronematous, mononematous, flexuous, simple or
branched, septate, hyaline, thin-walled, smooth. Conidiogenous
13
cells mono- or poly-blastic, sympodial with cylindrical denticles with truncate ends, hyaline, with numerous integrated
tiny pegs. Conidia 65–75 × 8–10.5 μm (x̄ = 68 × 9.5 μm, n =
15), acrogenous, solitary, mostly fusoid to sickle-shaped, sometimes sigmoid, slightly curved and acute at both ends, basal cell
Fungal Diversity (2020) 105:319–575
obconical, 10–13-euseptate, constricted at the septa, hyaline to
pale yellowish, densely guttulate, smooth-walled.
Culture characteristics: On PDA, colony irregular, reaching 5 mm in 25 days at 25 °C, brown from above, dark
brown from below, surface dry, rough, with sparse mycelium, mostly immersed in culture, with an irregular edge.
Material examined: THAILAND, Prachuap Khiri
Khan Province, on submerged wood in a stream, 30 July
2015, K.D. Hyde, 147A (MFLU 15-2713), living culture
MFLUCC 15-0995; Nakhon Si Thammarat Province, on
submerged wood in a stream, 10 May 2018, W. Dong,
hat6109 (MFLU 18-1720), living culture KUMCC 19-0078.
Notes: Our two new collections MFLUCC 15-0995 and
KUMCC 19-0078 were collected from submerged wood
in Thailand, and are morphologically and phylogenetically
identical to Aquaphila albicans (Goh et al. 1998a; Hyde
et al. 2016b). Phylogenetic analysis supports our two collections to be A. albicans (Fig. 129).
Berkleasmium Zobel, Icon. fung. (Prague) 6: 4 (1854)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Berkleasmium concinnum (Berk.) S.
Hughes
Notes: Berkleasmium is characterized by superficial ascomata seated on a subiculum, with multi-celled setae covering
the whole ascomata, fusiform, multi-septate ascospores, and
dictyoconidia or helicoid conidia (Lu et al. 2018b). The type
species B. concinnum was linked to the asexual morph of
Neoacanthostigma septoconstrictum (Promp. & A.N. Mill.)
Boonmee & K.D. Hyde, based on phylogenetic evidence
and co-occurrence of ascomata of N. septoconstrictum and
sporodochia of B. concinnum (Tanney and Miller 2017).
Berkleasmium comprises over 45 epithets in Index Fungorum (2020). However, Lu et al. (2018b) accepted six species based on phylogenetic evidence and their similar sexual
morph features. Five species were reported from freshwater
habitats (see list below) and four of them were transferred
from Neoacanthostigma Boonmee et al. The sexual morph
of B. aquaticum (Y.Z. Lu et al.) Y.Z. Lu has the typical
ascomata of Berkleasmium (Lu et al. 2017b).
List of freshwater Berkleasmium species
*Berkleasmium aquaticum (Y.Z. Lu, Boonmee & K.D.
Hyde) Y.Z. Lu, Fungal Diversity 92: 150 (2018)
Basionym: Neoacanthostigma aquaticum Y.Z. Lu, Boonmee & K.D. Hyde, Cryptog. Mycol. 38(2): 176 (2017)
Freshwater distribution: China (Lu et al. 2017b)
*Berkleasmium guangxiense (Y.Z. Lu, Boonmee & K.D.
Hyde) Y.Z. Lu, Fungal Diversity 92: 150 (2018)
519
Basionym: Neoacanthostigma guangxiense Y.Z. Lu,
Boonmee & K.D. Hyde, Cryptog. Mycol. 38(2): 182 (2017)
Freshwater distribution: China (Lu et al. 2017b)
*Berkleasmium latisporum (Y.Z. Lu, Boonmee & K.D.
Hyde) Y.Z. Lu, Fungal Diversity 92: 152 (2018); Fig. 119
Basionym: Neoacanthostigma latisporum Y.Z. Lu, Boonmee & K.D. Hyde, Cryptog. Mycol. 38(2): 184 (2017)
Freshwater distribution: Thailand (Lu et al. 2017b; this
study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies effuse, brown. Mycelium mostly immersed, composed of branched, septate,
brown hyphae. Conidiophores and Conidiogenous cells not
observed. Conidia 85–150 μm diam. and conidial filament
5.5–21.5 μm wide ( x̄ = 123 × 14 μm, n = 15), 545–865
μm long, coiled 2–3.5 times when tightly coiled, becoming loosely coiled in water, basal cell elongate, rounded at
the tip, distinctly multi-septate, pale brown to dark brown,
smooth-walled.
Culture characteristics: On PDA, colony irregular, reaching 7 mm in 20 days at 25 °C, dark brown to black from
above, black from below, surface dry, rough, umbonate,
veined and wrinkled, mycelium mostly immersed in culture,
with an irregular edge.
Material examined: THAILAND, Prachuap Khiri Khan
Province, on submerged wood in a stream, 30 July 2015, K.D.
Hyde, 34C (MFLU 15-2686), living culture MFLUCC 15-0987.
Notes: Our collection MFLUCC 15-0987 clusters with
Berkleasmium latisporum with high bootstrap support
(Fig. 129). They have identical LSU and TEF sequence data,
which indicates that they are conspecific (Jeewon and Hyde
2016). MFLUCC 15-0987 has larger conidial diam. (85–150
μm vs. 95–120 μm) and wider conidial filament (5.5–21.5
μm vs. 12–14.5 μm) than the holotype of B. latisporum (Lu
et al. 2017b).
*Berkleasmium longisporum Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 150 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Berkleasmium thailandicum (Tanney & A.N. Mill.) Y.Z.
Lu & K.D. Hyde, Fungal Diversity 92: 152 (2018)
Basionym: Neoacanthostigma thailandicum Tanney &
A.N. Mill., IMA Fungus 8(1): 103 (2017)
Synonymy: Neoacanthostigma brunneisporum Y.Z. Lu,
Boonmee & K.D. Hyde, Cryptog. Mycol. 38(2): 179 (2017)
Freshwater distribution: Thailand (Lu et al. 2017b)
Key to freshwater Berkleasmium species
1. Conidiogenous cells mono- to poly-blastic……………
……………………………………………B. aquaticum
13
520
Fungal Diversity (2020) 105:319–575
Fig. 119 Berkleasmium
latisporum (MFLU 15-2686). a
Colonies on submerged wood.
b–e Conidia. f–i Conidial filament. j Germinated conidium.
k, l Colony on PDA (up-front,
down-reverse). Scale bars: a =
100 μm, b = 20 μm, c–e = 40
μm, f = 5 μm, g = 2 μm, h, i =
10 μm, j = 50 μm
1. Conidiogenous cells only monoblastic or polyblastic………………………………………………………2
2. Conidiogenous cells monoblastic………………………3
2. Conidiogenous cells polyblastic………………………4
3. Conidial filament 11–15 μm wide……B. longisporum
3. Conidial filament 9–11 μm wide………B. thailandicum
13
4. Conidial filament 12–14.5 μm wide……B. latisporum
4. Conidial filament 8–10 μm wide………B. guangxiense
Boerlagiomyces Butzin, Willdenowia 8(1): 39 (1977)
Fungal Diversity (2020) 105:319–575
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Boonmee et al.
(2014).
Type species: Boerlagiomyces velutinus (Penz. & Sacc.)
Butzin
Notes: Boerlagiomyces is characterized by superficial
ascomata with sparse, hyphal appendages and muriform
ascospores (Boonmee et al. 2014). Boerlagiomyces was
placed in Tubeufiaceae based on a reference sequence of
B. macrosporus V.G. Rao & Varghese (Doilom et al. 2017)
and accepted by Lu et al. (2018b). The type species B. velutinus has eight ascospores (Boonmee et al. 2014), while three
freshwater species, viz. B. grandisporus S.J. Stanley & K.D.
Hyde, B. lacunosisporus (K.D. Hyde) S.J. Stanley & K.D.
Hyde and B. websteri Shearer & J.L. Crane, mostly have two
ascospores (Hyde 1992c; Shearer and Crane 1995; Stanley
and Hyde 1997). The sequence data of above freshwater
species are needed to confirm their phylogenetic placement.
List of freshwater Boerlagiomyces species
Boerlagiomyces grandisporus S.J. Stanley & K.D. Hyde,
Mycol. Res. 101(5): 635 (1997)
Freshwater distribution: Philippines (Stanley and Hyde
1997)
Boerlagiomyces lacunosisporus (K.D. Hyde) S.J. Stanley
& K.D. Hyde [as ‘lacunosispora’], Mycol. Res. 101(5): 640
(1997)
Basionym: Garethjonesia lacunosispora K.D. Hyde,
Aust. Syst. Bot. 5(4): 411 (1992)
Freshwater distribution: Queensland, Australia (Hyde
1992c)
Boerlagiomyces websteri Shearer & J.L. Crane, Mycologia
87(6): 876 (1996) [1995]
Freshwater distribution: Illinois, USA (Shearer and Crane
1995)
Key to freshwater Boerlagiomyces species
1.
1.
2.
2.
Asci 2(–3–4)-spored…………………B. lacunosisporus
Asci 2-spored…………………………………………2
Ascospores 78–149 × 21–66 μm………B. grandisporus
Ascospores 66–107 × 26–35 μm……………B. websteri
Chlamydotubeufia Boonmee & K.D. Hyde, Fungal Diversity 51(1): 78 (2011)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Chlamydotubeufia huaikangplaensis Boonmee & K.D. Hyde
Notes: Chlamydotubeufia has globose ascomata, a
helicosporous asexual morph and dictyochlamydospores
521
(Boonmee et al. 2011). Eight epithets are listed in Index
Fungorum (2020). Lu et al. (2018b) excluded three and
accepted five species in Chlamydotubeufia based on multigene phylogenetic analysis. Four species are reported from
freshwater habitats and all have a holomorph (Hyde et al.
2017; Lu et al. 2018b), except C. chlamydospora (Shearer)
Boonmee & K.D. Hyde (Shearer 1987).
List of freshwater Chlamydotubeufia species
*Chlamydotubeufia chlamydospora (Shearer) Boonmee &
K.D. Hyde [as ‘chlamydosporum’], Fungal Diversity 51(1):
83 (2011)
Basionym: Helicoma chlamydosporum Shearer, Mycologia 79(3): 468 (1987)
Freshwater distribution: Panama (Shearer 1987)
*Chlamydotubeufia cylindrica Y.Z. Lu, Boonmee & K.D.
Hyde, Fungal Diversity 92: 155 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Chlamydotubeufia huaikangplaensis Boonmee & K.D.
Hyde, Fungal Diversity 51(1): 78 (2011)
Freshwater distribution: Thailand (Hyde et al. 2017)
*Chlamydotubeufia krabiensis Y.Z. Lu, Boonmee & K.D.
Hyde, Fungal Diversity 87: 116 (2017)
Freshwater distribution: Thailand (Hyde et al. 2017)
Key to freshwater Chlamydotubeufia species
1.
1.
2.
2.
3.
3.
Asexual morph………………………C. chlamydospora
Holomorph……………………………………………2
Asci < 90 μm long………………C. huaikangplaensis
Asci > 90 μm long………………………………………3
Asci 90–145 × 14–16 μm…………………C. krabiensis
Asci 90–110 × 15–18 μm…………………C. cylindrica
Dematiohelicomyces Y.Z. Lu, Boonmee & K.D. Hyde, Fungal Diversity 92: 159 (2018)
Saprobic on submerged wood. Description of this genus
see Lu et al. (2018b).
Type species: Dematiohelicomyces helicosporus (Boonmee, Y.Z. Lu & K.D. Hyde) Y.Z. Lu
Notes: The monotypic genus Dematiohelicomyces was
introduced for a known species Chlamydotubeufia helicospora Boonmee et al. based on morphology and phylogeny
(Lu et al. 2018b). Dematiohelicomyces is similar to Helicomyces Link by short conidiophores and helicoid conidia
with a spathulate basal end cell, but differs in having brown
conidiophores (Lu et al. 2018b). Dematiohelicomyces helicosporus is exclusively recorded from freshwater habitats
(Hyde et al. 2016b; Lu et al. 2018b).
13
522
List of freshwater Dematiohelicomyces species
*Dematiohelicomyces helicosporus (Boonmee, Y.Z. Lu &
K.D. Hyde) Y.Z. Lu, Fungal Diversity 92: 159 (2018)
Basionym: Chlamydotubeufia helicospora Boonmee, Y.Z.
Lu & K.D. Hyde, Fungal Diversity 80: 123 (2016)
Freshwater distribution: Thailand (Hyde et al. 2016b; Lu
et al. 2018b)
Dictyospora Brahaman., Y.Z. Lu, Boonmee & K.D. Hyde,
Mycosphere 8(7): 924 (2017)
Saprobic on submerged wood. Description of this genus
see Brahmanage et al. (2017).
Type species: Dictyospora thailandica Brahaman., Y.Z.
Lu, Boonmee & K.D. Hyde
Notes: The monotypic genus Dictyospora was introduced
for a freshwater species D. thailandica (Brahmanage et al.
2017), which was later recollected by Lu et al. (2018b) and
characterized by larger ascomata, smaller asci and shorter
ascospores than the holotype. Dictyospora has dictyochlamydospores which are similar to Chlamydotubeufia, but
phylogeny separates them as distinct genera (Brahmanage
et al. 2017; Lu et al. 2018b).
List of freshwater Dictyospora species
*Dictyospora thailandica Brahaman., Y.Z. Lu, Boonmee &
K.D. Hyde, Mycosphere 8(7): 924 (2017)
Freshwater distribution: Thailand (Brahmanage et al.
2017; Lu et al. 2018b)
Helicoarctatus Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 165 (2018)
Saprobic on submerged wood. Description of this genus
see Lu et al. (2018b).
Type species: Helicoarctatus aquaticus Y.Z. Lu, J.C.
Kang & K.D. Hyde
Notes: The monotypic genus Helicoarctatus is characterized by setiform conidiophores, discrete conidiogenous cells
arising laterally from lower portion of the conidiophores,
with each bearing 1–2 tiny conidiogenous loci, which are
similar to Helicosporium Nees (Lu et al. 2018b). Helicoarctatus differs from Helicosporium by larger conidia and
molecular characters (Lu et al. 2018b). The type species H.
aquaticus formed a sister clade with a sexual morphic species Neohelicoma fagacearum (Boonmee & K.D. Hyde) Y.Z.
Lu with high bootstrap support (Lu et al. 2018b). Helicoarctatus aquaticus and N. fagacearum are separated as distinct
genera based on phylogenetic analysis (Lu et al. 2018b).
List of freshwater Helicoarctatus species
*Helicoarctatus aquaticus Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 166 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
13
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Helicodochium J.S. Monteiro, R.F. Castañeda, A.C. Cruz &
Gusmão, Mycotaxon 127: 6 (2014)
Saprobic on submerged wood. Description of this genus
see Lu et al. (2018b).
Type species: Helicodochium amazonicum J.S. Monteiro
et al.
Notes: Helicodochium was introduced by Monteiro et al.
(2014) and characterized by sporodochial, apothecium-like,
pinkish to pink conidiomata with macronematous conidiophores arising from a thick-walled, pseudoparenchymatous stromata, and hyaline to pink conidia (Monteiro et al.
2014; Lu et al. 2018b). Lu et al. (2018b) transferred Helicodochium from Ascomycota genera incertae sedis to Tubeufiaceae based on morphology and phylogeny, together with
a new species H. aquaticum Y.Z. Lu et al. The type species
H. amazonicum lacks sequence data in Genbank.
List of freshwater Helicodochium species
Helicodochium amazonicum J.S. Monteiro, R.F. Castañeda,
A.C. Cruz & Gusmão, Mycotaxon 127: 6 (2014)
Freshwater distribution: Brazil (Monteiro et al. 2014)
*Helicodochium aquaticum Y.Z. Lu, Boonmee & K.D.
Hyde, Fungal Diversity 92: 168 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Helicodochium species
1. Conidia 70–82.5 μm diam.……………H. amazonicum
1. Conidia 42–60 μm diam.…………………H. aquaticum
Helicohyalinum Y.Z. Lu, J.K. Liu & K.D. Hyde, Fungal
Diversity 92: 168 (2018)
Saprobic on submerged wood. Description of this genus
see Lu et al. (2018b).
Type species: Helicohyalinum infundibulum Y.Z. Lu, J.K.
Liu & K.D. Hyde
Notes: Helicohyalinum differs from other helicosporous
genera by its flexuous, hyaline conidiophores with the terminal, monoblastic conidiogenous cells tapering towards the
apex (Lu et al. 2018b). Two freshwater species, H. aquaticum (Brahaman. et al.) J.K. Liu and H. infundibulum Y.Z.
Lu et al., were collected from Thailand (Brahmanage et al.
2017; Lu et al. 2018b). They can be distinguished by conidial size and coiling characters (conidia tightly coiled which
are not loose in water in H. infundibulum vs. conidia loosely
coiled or uncoiled in water in H. aquaticum).
List of freshwater Helicohyalinum species
*Helicohyalinum aquaticum (Brahaman., Y.Z. Lu, Boonmee & K.D. Hyde) J.K. Liu, Fungal Diversity 92: 170 (2018)
Fungal Diversity (2020) 105:319–575
Basionym: Chlamydotubeufia aquatica Brahaman., Y.Z.
Lu, Boonmee & K.D. Hyde, Mycosphere 8(7): 922 (2017)
Freshwater distribution: Thailand (Brahmanage et al.
2017)
*Helicohyalinum infundibulum Y.Z. Lu, J.K. Liu & K.D.
Hyde, Fungal Diversity 92: 170 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Helicohyalinum species
523
*Helicoma hongkongense (C.K.M. Tsui, Goh, K.D. Hyde &
Hodgkiss) Y.Z. Lu, Fungal Diversity 92: 187 (2018)
Basionym: Helicosporium hongkongense C.K.M. Tsui,
Goh, K.D. Hyde & Hodgkiss, Mycologia 90(2): 392 (1998)
Freshwater distribution: Hong Kong, China (Tsui et al.
2001a), Thailand (Lu et al. 2018b)
*Helicoma longisporum Y.Z. Lu, J.K. Liu & K.D. Hyde,
Fungal Diversity 92: 178 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
1. Conidia 62–89 μm diam.………………H. aquaticum
1. Conidia 40–50 μm diam.……………H. infundibulum
*Helicoma multiseptatum Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 180 (2018)
Freshwater distribution: China (Lu et al. 2018b)
Helicoma Corda, Icon. fung. (Prague) 1: 15 (1837)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Helicoma muelleri Corda
Notes: Helicoma is an old genus comprising over 90 epithets in Index Fungorum (2020). Most Helicoma species
lack molecular data and thus were considered to be probably
misidentified (Lu et al. 2018b). The sexual morphs were
reported by Boonmee et al. (2014). Based on new collections and molecular data, Lu et al. (2018b) redefined generic
concept of Helicoma as conidiogenous cells that are intercalary, cylindrical, with denticles, arising laterally from the
lower portion of conidiophores as tooth-like protrusions,
and conidia are pleurogenous, helicoid, hygroscopic, tapering towards apex and rounded at tip, coiled 1.5–5 times,
becoming loosely coiled in water. Lu et al. (2018b) accepted
57 species in Helicoma and eight of them were found from
freshwater habitats. The molecular data of H. gigasporum
(C.K.M. Tsui et al.) Y.Z. Lu are needed to confirm its placement in Helicoma.
*Helicoma nematosporum (Linder) Y.Z. Lu & K.D. Hyde,
Fungal Diversity 92: 189 (2018)
Basionym: Helicosporium nematosporum Linder, Ann.
Mo. bot. Gdn 16: 288 (1929)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Helicoma septoconstrictum Y.Z. Lu, J.K. Liu & K.D.
Hyde, Fungal Diversity 92: 185 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Helicoma species
1.
1.
2.
2.
3.
3.
4.
4.
List of freshwater Helicoma species
*Helicoma aquaticum Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 174 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Helicoma gigasporum (C.K.M. Tsui, Goh, K.D. Hyde &
Hodgkiss) Y.Z. Lu, Fungal Diversity 92: 187 (2018)
Basionym: Helicosporium gigasporum C.K.M. Tsui, Goh,
K.D. Hyde & Hodgkiss, Mycologia 90(2): 392 (1998)
Freshwater distribution: Hong Kong, China (Tsui et al.
2001a)
*Helicoma guttulatum Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 80: 125 (2016)
Freshwater distribution: Thailand (Hyde et al. 2016b)
5.
5.
6.
6.
7.
7.
Conidiogenous cells monoblastic………………………2
Conidiogenous cells mono- to polyblastic……………5
Conidia > 80 μm diam.……………H. septoconstrictum
Conidia < 80 μm diam.…………………………………3
Conidia < 50 μm diam.………………H. hongkongense
Conidia > 50 μm diam.…………………………………4
Conidia hyaline to pale brown, coiled 1–2 times………
………………………………………H. multiseptatum
Conidia pale yellow or pale brown, coiled 2–2.5 times…
…………………………………………H. gigasporum
Conidia > 45 μm diam.………………H. longisporum
Conidia < 45 μm diam.…………………………………6
Conidia < 30 μm diam.…………………H. guttulatum
Conidia > 30 μm diam.…………………………………7
Conidia 310–490 μm long…………H. nematosporum
Conidia 220–320 μm long………………H. aquaticum
Helicomyces Link, Mag. Gesell. naturf. Freunde, Berlin 3(12): 21 (1809)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Helicomyces roseus Link
Notes: The type species Helicomyces roseus is phylogenetically confused, because two isolates (BCC 3381 and
BCC 8808) were transferred to Tubeufia Penz. & Sacc., five
13
524
isolates of Luo et al. (2017) were transferred to Pseudohelicomyces Y.Z. Lu et al., and one isolate (CBS 283.51) of Tsui
et al. (2006b) is misidentified (Lu et al. 2018b). Although the
sequence data of H. roseus is lacking, Lu et al. (2018b) named
the clade comprising four freshwater species (see list below)
as Helicomyces sensu stricto. The epitype or reference specimen of H. roseus is needed to confirm the conclusion of Lu
et al. (2018b).
List of freshwater Helicomyces species
*Helicomyces chiayiensis (C.H. Kuo & Goh) Y.Z. Lu &
K.D. Hyde, Fungal Diversity 92: 205 (2018)
Basionym: Helicoma chiayiense C.H. Kuo & Goh, Mycol.
Progr. 17(5): 561 (2018)
Freshwater distribution: Taiwan, China (Kuo and Goh
2018)
*Helicomyces colligatus R.T. Moore, Mycologia 46(1): 89
(1954)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Helicomyces hyalosporus Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 201 (2018)
Freshwater distribution: China (Lu et al. 2018b)
Helicomyces roseus Link, Mag. Gesell. naturf. Freunde,
Berlin 3(1-2): 21 (1809)
Freshwater distribution: Australia (Hyde and Goh 1997),
Brunei (Ho et al. 2001), Hong Kong, China (Ho et al. 2001,
2002), Malaysia (Ho et al. 2001)
*Helicomyces torquatus L.C. Lane & Shearer, Mycotaxon
19: 291 (1984)
Freshwater distribution: Malaysia (Ho et al. 2001), Panama (Lane and Shearer 1984), Thailand (Lu et al. 2018b)
Key to freshwater Helicomyces species
1.
1.
2.
2.
3.
3.
4.
Conidiogenous cells monoblastic…………H. torquatus
Conidiogenous cells mono- to poly-blastic……………2
Conidia > 70 μm diam.…………………H. hyalosporus
Conidia < 70 μm diam.…………………………………3
Conidia < 50 μm diam.…………………H. chiayiensis
Conidia > 50 μm diam.…………………………………4
Conidia 50–70 μm diam., conidial filament 7–11 μm
wide………………………………………H. colligatus
4. Conidia 25–60 μm diam., conidial filament 2.5–6 μm
wide…………………………………………H. roseus
Helicosporium Nees, Syst. Pilze (Würzburg): 68 (1816)
[1816-17]
13
Fungal Diversity (2020) 105:319–575
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Helicosporium vegetum Nees
Notes: Helicosporium is an old genus comprising over
100 epithets in Index Fungorum (2020). The generic concept
of Helicosporium was redefined by Linder (1929) and Goos
(1989) and recently reviewed by Lu et al. (2018b) who gave
the latest update with additional collections and multigene
phylogenetic analysis. Helicosporium setiferum Y.Z. Lu
et al. was introduced by Lu et al. (2018b) to accommodate
five freshwater isolates. The isolates BCC 3332 and BCC
8125 provided by Tsui et al. (2006b) lack morphological
data, but phylogeny identified them as H. setiferum (Lu et al.
2018b). Thirteen species were accepted in Helicosporium
(Lu et al. 2018b) and five of them were reported from freshwater habitats in Thailand. We report a new freshwater species H. thailandense which is collected from Thailand.
List of freshwater Helicosporium species
*Helicosporium aquaticum Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 208 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Helicosporium flavisporum Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 210 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Helicosporium flavum Brahaman., Y.Z. Lu, Boonmee &
K.D. Hyde, Mycosphere 8(7): 931 (2017)
Freshwater distribution: Thailand (Brahmanage et al.
2017)
*Helicosporium setiferum Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 213 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Helicosporium thailandense H. Zhang, W. Dong & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557940; Facesoffungi number: FoF09285; Fig. 120
Etymology: referring to Thailand, where the holotype was
collected
Holotype: MFLU 18-1561
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
superficial, effuse, gregarious, yellow-green. Mycelium
partly immersed in natural substratum, branched, hyaline
to pale brown. Conidiophores 450–550 × 2.8–4.5 µm ( x̄
= 500 × 3.5 µm, n = 5), macronematous, mononematous,
sparsely branched, slender, cylindrical, smooth-walled,
brown, septate, slightly constricted at septa, thin-walled.
Conidiogenous cells 17–25 × 2.5–4.7 µm ( x̄ = 20 × 3.5
µm, n = 15), denticulate, holoblastic, mono- to polyblastic,
Fungal Diversity (2020) 105:319–575
525
Fig. 120 Helicosporium
thailandense (MFLU 18-1561,
holotype). a Colonies on
submerged wood. b, c Conidiophores with attached conidia.
d–f Denticulate conidiogenous
cells. g–k Conidia. l, m Colony
on PDA (left-front, rightreverse). Scale bars: b = 30 μm,
c = 50 μm, d, e, g–k = 20 μm,
f = 10 μm
integrated, terminal or intercalary, sympodial, arising laterally from lower portion of the conidiophores as tiny toothlike protrusions, with each bearing 1–6 tiny conidiogenous
loci, 1.2–2.2 × 1–1.5 µm ( x̄ = 1.8 × 1 µm, n = 10), brown,
smooth-walled. Conidia solitary, dry, pleurogenous, helicoid, tapering at tip, 22–28 µm diam. and conidial filament
2.6–3.8 µm wide ( x̄ = 25 × 3.2 µm, n = 10), 110–125 µm
long, loosely coiled 1.75–2.25 times, indistinctly septate,
8–10-septate, guttulate, hyaline, sometimes slightly light
yellow, smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 10 days at 25 °C, brown from above, dark brown
from below, surface rough, with dense mycelium, dry, edge
entire.
Material examined: THAILAND, Satun Province, on
submerged wood in a stream, 10 May 2018, W. Dong,
hat282-1 (MFLU 18-1561, holotype), ex-type living culture MFLUCC 18-1407; ibid., hat282-2 (HKAS 105051,
isotype), ex-type living culture KUMCC 19-0071.
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526
Notes: Helicosporium thailandense clusters with a terrestrial species H. viridiflavum Y.Z. Lu et al. with high
bootstrap support (Fig. 129). Helicosporium thailandense
is similar to H. viridiflavum in having yellow-green colonies
and long conidiophores (Lu et al. 2018b). However, they
can be distinguished by the conidiophore length (450–550
× 2.8–4.5 µm vs. 250–425 × 3–4 µm), conidial diameter
(22–28 µm vs. 20–23 µm) and septation (8–10 vs. 13–16).
Phylogenetic analysis supports them to be different species
(Fig. 129).
*Helicosporium vesicarium Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 213 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Helicosporium species
1.
1.
2.
2.
3.
3.
4.
4.
5.
Conidiophores < 65 μm long………H. flavum
Conidiophores > 65 μm long……………………………2
Conidia < 90 μm long……………………H. aquaticum
Conidia > 90 μm long…………………………………3
Conidiophores < 120 μm long…………H. vesicarium
Conidiophores > 120 μm long…………………………4
Conidiophores > 450 μm long…………H. thailandense
Conidiophores < 450 μm long…………………………5
Conidiophores 125–320 μm, conidia 13–21 μm diam.…
……………………………………………H. setiferum
5. Conidiophores 130–180 μm, conidia 12–15 μm diam.…
…………………………………………H. flavisporum
Helicotruncatum Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 220 (2018)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Helicotruncatum palmigenum (Penz. &
Sacc.) Y.Z. Lu & K.D. Hyde
Notes: Helicotruncatum was introduced by Lu et al.
(2018b) based on ITS and LSU sequence data of Helicoma
palmigenum (Penz. & Sacc.) Linder NBRC 32663 (Tsui
et al. 2006b) and previous literatures (Linder 1929; Matsushima 1971; Goos 1986). The thickened lateral wall of the
conidiophores and conidia of Helicotruncatum are distinct
and represents a generic feature in Tubeufiaceae (Lu et al.
2018b). Although the molecular characters of NBRC 32663
warrant a distinct genus, the morphology was not provided
(Tsui et al. 2006b). Helicotruncatum palmigenum commonly
occurrs in terrestrial habitats, and herein we report it from
freshwater habitats.
List of freshwater Helicotruncatum species
*Helicotruncatum palmigenum (Penz. & Sacc.) Y.Z. Lu &
K.D. Hyde, Fungal Diversity 92: 220 (2018); Fig. 121
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Fungal Diversity (2020) 105:319–575
Basionym: Helicosporium intermedium var. palmigenum
Penz. & Sacc., Malpighia 15(7-9): 249 (1902) [1901]
Synonymy: Helicoma palmigenum (Penz. & Sacc.) Linder,
Ann. Mo. bot. Gdn 16: 306 (1929)
Freshwater distribution: Thailand (this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies superficial, effuse, erect,
gregarious, black. Mycelium composed of partly immersed,
pale brown to brown, septate hyphae. Conidiophores 180–
310 × 5–6 µm ( x̄ = 250 × 5 µm, n = 10), macronematous,
mononematous, unbranched, erect, cylindrical, robust at
the base, ca 10 µm diam., smooth-walled, pale brown to
brown at the upper part, dark brown to black near the base,
septate, slightly constricted at septa, uneven pigmented,
pale brown near the septa, thin-walled. Conidiogenous cells
17–25 × 5–6 µm ( x̄ = 22 × 6 µm, n = 10), holoblastic,
monoblastic, integrated, determinate, terminal, subcylindrical, pale brown, smooth-walled, with broad, flat point
of conidial attachment. Conidia solitary, dry, acrogenous,
helicoid, truncate at tip, 30–40 µm diam. and conidial filament 10–15 µm wide ( x̄ = 35 × 13 µm, n = 10), 50–60 µm
long, tightly coiled 1.5–1.75 times, distinctly 12–14-septate,
guttulate, pale brown when young, becoming dark brown or
grey-brown when old, the basal cell truncate with thickened
lateral walls, smooth-walled.
Culture characteristics: On PDA, colony irregular, reaching 20 mm in 18 days at 25 °C, dark brown from above and
below, surface rough, with sparse mycelium, dry, umbonate,
edge undulate to filiform.
Material examined: THAILAND, Prachuap Khiri
Khan Province, on submerged wood in a stream, 30 July
2015, K.D. Hyde, 143B (MFLU 15-2715), living culture
MFLUCC 15-0993.
Notes: Our new collection clusters with H. palmigenum
(NBRC 32663) with high bootstrap support (Fig. 129).
Although NBRC 32663 is named as Helicotruncatum palmigenum, the morphology has not been determined (Tsui
et al. 2006b). Information on the holotype of H. palmigenum
is unavailable.
Our collection MFLUCC 15-0993 has similar characters with specimens reported in Linder (1929), Matsushima
(1971) and Goos (1986). However, conidia of MFLUCC
15-0993 have more septa (12–14 vs. 10–12) and wider
conidial filaments (10–15 µm vs. 11–12 µm) than in Linder
(1929), and longer conidiophores (180–310 µm vs. 60–140
µm) than in Matsushima (1971). MFLUCC 15-0993 has
similar conidial size with Goos (1986), but has longer conidiophores (180–310 µm vs. 60–215 µm). The length of conidiophores in Linder (1929) is not given.
Comparing nucleotides between NBRC 32663 and
MFLUCC 15-0993, there are two nucleotide differences
crossing 549 base pairs in LSU sequence data, while nine
Fungal Diversity (2020) 105:319–575
527
Fig. 121 Helicotruncatum
palmigenum (MFLU 15-2715,
new habitat and geographical record). a, b Colonies on
submerged wood. c, d Conidiophores with attached conidia.
e–g Conidiogenous cells.
h–k Conidia. l Germinated
conidium. m, n Colony on PDA
(up-front, down-reverse). Scale
bars: a, b = 100 μm, c = 40 μm,
d, l = 20 μm, e–k = 10 μm
nucleotide differences (including three deletions and three
insertions) in ITS sequence data. We name MFLUCC
15-0993 as Helicotruncatum palmigenum mainly based
on the molecular evidence (Tsui et al. 2006b), coupled
with similar morphological characters in previous publications (Linder 1929; Matsushima 1971; Goos 1986).
Epitypification of H. palmigenum is needed using a collection from its type locality, Indonesia, to confirm the previous studies (Linder 1929; Matsushima 1971; Goos 1986;
Tsui et al. 2006b) and our study. This is a new habitat and
geographical record for H. palmigenum from freshwater in
Thailand.
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528
Helicotubeufia Y.Z. Lu & J.K. Liu, Mycosphere 9(3): 500
(2018)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see (Liu et al. 2018a).
Type species: Helicotubeufia guangxiensis Y.Z. Lu & J.K.
Liu
Notes: Helicotubeufia was introduced by Liu et al.
(2018a) to accommodate two freshwater species H. guangxiensis and H. jonesii Y.Z. Lu & J.K. Liu, and one terrestrial
species H. hydei Y.Z. Lu & J.K. Liu. Helicotubeufia has dark
brown to black ascomata seated on a subiculum, cylindrical
asci, fusiform, multiseptate ascospores and helicosporous
asexual morphs (Liu et al. 2018a). Both sexual and asexual
morphs of H. guangxiensis were found on submerged wood
in nature. The asexual morph of H. jonesii is unknown.
List of freshwater Helicotubeufia species
*Helicotubeufia guangxiensis Y.Z. Lu & J.K. Liu, Mycosphere 9(3): 500 (2018)
Freshwater distribution: China (Liu et al. 2018a)
*Helicotubeufia jonesii Y.Z. Lu & J.K. Liu, Mycosphere
9(3): 503 (2018)
Freshwater distribution: China (Liu et al. 2018a)
Fungal Diversity (2020) 105:319–575
Type species: Neochlamydotubeufia fusiformis Y.Z. Lu,
Boonmee & K.D. Hyde
Notes: Neochlamydotubeufia was introduced with sexual
and asexual morphs which are characterized by superficial
ascomata with tapering, unbranched, thick-walled setae and
fusiform, multi-septate, hyaline to pale brown ascospores
and broadly oval to ellipsoid chlamydospores (Lu et al.
2018b). These characters are similar to Chlamydotubeufia
and Dictyospora, but phylogeny separates them as different
genera (Boonmee et al. 2011; Brahmanage et al. 2017; Lu
et al. 2018b). The sexual and asexual morphs of N. fusiformis were separately collected from natural submerged
wood and linked with molecular evidence (Lu et al. 2018b).
Neochlamydotubeufia khunkornensis (Boonmee & K.D.
Hyde) Y.Z. Lu was collected from dead wood in terrestrial
habitats and formed asexual morph in MEA (Boonmee et al.
2011). Lu et al. (2018b) found chlamydospores of N. khunkornensis from submerged wood. Neochlamydotubeufia fusiformis has smaller ascomata than N. khunkornensis and they
are separated by molecular data (Lu et al. 2018b).
List of freshwater Neochlamydotubeufia species
*Neochlamydotubeufia fusiformis Y.Z. Lu, Boonmee &
K.D. Hyde, Fungal Diversity 92: 221 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Helicotubeufia species
1. Ascospores 55–70 × 5–6 μm…………H. guangxiensis
1. Ascospores 35–43 (–47) × 4.5–7 μm…………H. jonesii
Muripulchra Z.L. Luo, H.Y. Su & K.D. Hyde, Cryptog.
Mycol. 38(1): 36 (2017)
Saprobic on submerged wood. Description of this genus
see Luo et al. (2017).
Type species: Muripulchra aquatica Z.L. Luo, H.Y. Su&
K.D. Hyde
Notes: The monotypic genus Muripulchra was introduced
by Luo et al. (2017) for a freshwater species M. aquatica,
which is distinct in Tubeufiaceae in producing micronematous conidiophores, obpyriform, muriform, dark brown,
thick-walled, granulate conidia.
List of freshwater Muripulchra species
*Muripulchra aquatica Z.L. Luo, H.Y. Su & K.D. Hyde,
Cryptog. Mycol. 38(1): 39 (2017)
Freshwater distribution: China (Luo et al. 2017)
Neochlamydotubeufia Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 92: 221 (2018)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
13
*Neochlamydotubeufia khunkornensis (Boonmee & K.D.
Hyde) Y.Z. Lu, Fungal Diversity 92: 222 (2018)
Basionym: Chlamydotubeufia khunkornensis Boonmee &
K.D. Hyde, Fungal Diversity 51(1): 80 (2011)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Neochlamydotubeufia species
1. Ascomata 140–180 × 140–170 μm………N. fusiformis
1. Ascomata 180–250 × 200–250 μm……………………
………………………………………N. khunkornensis
Neohelicomyces Z.L. Luo, Bhat & K.D. Hyde, Cryptog.
Mycol. 38(1): 39 (2017)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Luo et al. (2017) and
and Lu et al. (2018b).
Type species: Neohelicomyces aquaticus Z.L. Luo, Bhat
& K.D. Hyde
Notes: Neohelicomyces was introduced by Luo et al.
(2017) for three freshwater species. Lu et al. (2018b) introduced the fourth freshwater species N. hyalosporus Y.Z. Lu
et al. which differs from the others in having longer conidiophores and hyaline conidia. Neohelicomyces is characterized by white colonies, macronematous, branched conidiophores, mono- to poly-blastic, integrated conidiogenous
Fungal Diversity (2020) 105:319–575
529
Fig. 122 Neohelicomyces
dehongensis (MFLU 18-1168,
holotype). a, b Colonies on
submerged wood. c, d Conidiophores with attached conidia.
e Apex of conidiophore without
conidiogenous cells. f, g Conidiogenous cells. h–j Conidia.
k, l Colony on PDA (left-front,
right-reverse). Scale bars: c =
40 μm, d–f, h–j = 20 μm, g =
5 μm
cells with lateral minute denticles and helicoid conidia
(Luo et al. 2017). Two strains Tubeufia amazonensis Samuels et al. (ATCC 42524) and T. helicomyces Höhn. (MUCL
15702) cluster in Neohelicomyces (Lu et al. 2018b; this
study, Fig. 129), but are not synonymized because they lack
morphology (Tsui et al. 2006b). We introduce another two
freshwater Neohelicomyces species collected from Thailand
and China.
List of freshwater Neohelicomyces species
*Neohelicomyces aquaticus Z.L. Luo, Bhat & K.D. Hyde,
Cryptog. Mycol. 38(1): 40 (2017)
13
530
Freshwater distribution: China (Luo et al. 2017)
*Neohelicomyces dehongensis H. Zhang, W. Dong & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557941; Facesoffungi number: FoF09286; Fig. 122
Etymology: referring to Dehong, where the holotype was
collected
Holotype: MFLU 18-1168
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
superficial, effuse, gregarious, white. Mycelium composed
of partly immersed, partly superficial, hyaline to pale brown,
septate, branched hyphae, with mass of crowded, glistening conidia. Conidiophores 120–250 × 3.3–3.9 µm ( x̄ =
180 × 3.5 µm, n = 10), macronematous, mononematous,
unbranched or loosely branched, branches often forked close
to the base, upper part frequently sterile, erect, cylindrical,
pale brown, septate, slightly constricted at septa, guttulate,
smooth-walled, thin-walled. Conidiogenous cells 12–17.5
× 3–4 µm ( x̄ = 14.5 × 3.5 µm, n = 20), holoblastic, monoto polyblastic, integrated, sympodial, intercalary, cylindrical, with denticles, hyaline to pale brown, smooth-walled.
Conidia solitary, dry, pleurogenous, helicoid, rounded at
tip, 20–25 µm diam. and conidial filament 2.2–4 µm wide
( x̄ = 22 × 3 µm, n = 15), 145–210 µm long, tightly coiled
2.75–3.75 times, becoming loosely coiled in water, indistinctly multi-septate, guttulate, hyaline, smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 25 days at 25 °C, brown from above and below,
surface rough, with dense mycelium, dry, edge entire.
Material examined: CHINA, Yunnan Province, Dehong,
on submerged wood in a stream, 25 November 2017, G.N.
Wang, H3C-1 (MFLU 18-1168, holotype), ex-type living
culture MFLUCC 18-1029; ibid., H3C-2 (HKAS 101728,
isotype), ex-type living culture KUMCC 18-0078.
Notes: Neohelicomyces dehongensis clusters with three
strains of N. pallidus (Ces.) Y.Z. Lu & K.D. Hyde (Fig. 129).
Neohelicomyces dehongensis differs from N. pallidus in having larger conidia (20–25 µm diam. vs. 10–16 μm diam.) and
wider conidial filament (2.2–4 µm vs. 1–2 µm) (Linder 1929;
Zhao et al. 2007). Neohelicomyces pallidus (UAMH 10535)
clusters in the same clade with N. pandanicola Tibpromma
& K.D. Hyde, which is apart from the other three strains of
N. pallidus. This indicates that N. pallidus UAMH 10535 is
a distinct species. However, we do not change its status as
there is a lack of morphology.
*Neohelicomyces grandisporus Z.L. Luo, Boonmee & K.D.
Hyde, Cryptog. Mycol. 38(1): 44 (2017)
Freshwater distribution: China (Luo et al. 2017)
Fungal Diversity (2020) 105:319–575
*Neohelicomyces hyalosporus Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 225 (2018)
Freshwater distribution: China (Lu et al. 2018b)
*Neohelicomyces submersus Z.L. Luo, H.Y. Su & K.D.
Hyde, Cryptog. Mycol. 38(1): 40 (2017)
Freshwater distribution: China (Luo et al. 2017)
*Neohelicomyces thailandicus H. Zhang, W. Dong & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557942; Facesoffungi number: FoF09287; Fig. 123
Etymology: referring to Thailand, where the holotype was
collected
Holotype: MFLU 11-1079
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
superficial, effuse, gregarious, white. Mycelium mostly
immersed in natural substratum, sparsely branched, hyaline
to pale brown. Conidiophores 300–400 × 2.8–4.6 µm ( x̄
= 345 × 3.6 µm, n = 5), macronematous, mononematous,
sparsely branched, erect, slender, uneven in width, cylindrical, smooth-walled, brown to pale brown, septate, slightly
constricted at septa, thin-walled. Conidiogenous cells 8–11
× 2–3.5 µm ( x̄ = 9.7 × 2.8 µm, n = 10), holoblastic, monoto polyblastic, discrete, determinate, arising in a whorl
from pale brown, ampulliform-like or subcylindrical cells
or directly from conidiophores near every septum, with each
bearing 1–4 tiny conidiogenous loci, hyaline to pale brown,
smooth-walled. Conidia solitary, dry, pleurogenous, helicoid, rounded at tip, 14–18 µm diam. and conidial filament
1.3–1.8 µm wide ( x̄ = 15.5 × 1.6 µm, n = 10), 120–145
µm long, tightly coiled 3.75–4.25 times, indistinctly multiseptate, guttulate, hyaline, smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 5 mm in 10 days at 25 °C, brown from above and below,
surface rough, with dense mycelium, dry, edge entire.
Material examined: THAILAND, Chiang Rai Province,
on submerged wood in a stream, 18 January 2010, H. Zhang,
a47 (MFLU 11-1079, holotype), ex-type living culture
MFLUCC 11-0005.
Notes: Neohelicomyces thailandicus clusters in Neohelicomyces with high bootstrap support (Fig. 129). It can be
distinguished from the other species by its conidiogenous
cells arising from ampulliform-like or subcylindrical cells
and conidiophores near the septa, with each bearing 1–4
tiny conidiogenous loci (Luo et al. 2017; Lu et al. 2018b;
Tibpromma et al. 2018; Crous et al. 2019).
Key to freshwater Neohelicomyces species
1. Conidiophores < 170 μm long………N. grandisporus
1. Conidiophores > 170 μm long…………………………2
13
Fungal Diversity (2020) 105:319–575
Fig. 123 Neohelicomyces thailandicus (MFLU 11-1079, holotype).
a Colonies on submerged wood. b Conidiophores with attached
conidia. c–e Conidiogenous cells. f–h Conidia. i Germinated conid-
531
ium. j, k Colony on PDA (up-front, down-reverse). Scale bars: b, i =
30 μm, c–e = 10 μm, f–h = 5 μm
13
532
2.
2.
3.
3.
4.
4.
5.
Conidia hyaline…………………………………………3
Conidia pale brown……………………………………5
Conidia > 20 μm diam.…………………N. dehongensis
Conidia < 20 μm diam.…………………………………4
Conidia pleurogenous…………………N. thailandicus
Conidia acropleurogenous………………N. hyalosporus
Conidiophores 240.5–335.5 μm, conidia coiled 2–2.5
times………………………………………N. aquaticus
5. Conidiophores 172–285 μm, conidia coiled 3–3.5
times……………………………………N. submersus
Neohelicosporium Y.Z. Lu, J.C. Kang & K.D. Hyde, Mycol.
Progr. 17(5): 637(2017)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018a).
Type species: Neohelicosporium parvisporum Y.Z. Lu,
J.C. Kang & K.D. Hyde
Notes: Neohelicosporium was introduced by Lu et al.
(2018a) for five freshwater species. Neohelicosporium differs from Helicosporium by its integrated, sympodial conidiogenous cells and acrogenous and/or acropleurogenous
conidia (Lu et al. 2018b). Based on phylogeny and morphology, Lu et al. (2018b) transferred some species of Helicoma,
Helicosporium and Tubeufia to Neohelicosporium. Two
strains HKUCC 10235 and CBS 189.95 were temporarily
named as Neohelicosporium sp. until new collections with
molecular data are obtained (Lu et al. 2018b). All freshwater species were found from China and Thailand (see list
below). Neohelicosporium ellipsoideum Y.Z. Lu et al. is
the only freshwater species producing holomorph and N.
irregulare Y.Z. Lu et al. is unique in producing two lengths
of conidiophores (Lu et al. 2018b). Neohelicosporium species are variable in types of conidiophores and the size of
conidia and conidiophores.
List of freshwater Neohelicosporium species
*Neohelicosporium acrogenisporum Y.Z. Lu, J.C. Kang &
K.D. Hyde, Fungal Diversity 92: 228 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Neohelicosporium aquaticum Y.Z. Lu, J.C. Kang & K.D.
Hyde, Mycol. Progr. 17(5): 639(2017), [9] (2017)
Freshwater distribution: China (Lu et al. 2018a)
*Neohelicosporium astrictum Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 230 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Neohelicosporium ellipsoideum Y.Z. Lu, J.C. Kang &
K.D. Hyde, Fungal Diversity 92: 230 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
13
Fungal Diversity (2020) 105:319–575
*Neohelicosporium guangxiense Y.Z. Lu, J.C. Kang &
K.D. Hyde, Mycol. Progr. 17(5): 639(2017), [9] (2017)
Freshwater distribution: China (Lu et al. 2018a)
*Neohelicosporium hyalosporum Y.Z. Lu, J.C. Kang &
K.D. Hyde, Mycol. Progr. 17(5): 639(2017), [11] (2017)
Freshwater distribution: China (Lu et al. 2018a)
*Neohelicosporium irregulare Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 235 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Neohelicosporium krabiense Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 236 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Neohelicosporium parvisporum Y.Z. Lu, J.C. Kang &
K.D. Hyde, Mycol. Progr. 17(5): 637(2017), [7] (2017)
Freshwater distribution: China (Lu et al. 2018a)
*Neohelicosporium submersum H. Zhang, W. Dong & K.D.
Hyde, sp. nov.
Index Fungorum number: IF557943; Facesoffungi number: FoF09288; Fig. 124
Etymology: in reference to the submerged habitats of the
fungus
Holotype: MFLU 17-1734
Saprobic on decaying, submerged wood in freshwater.
Sexual morph: Undetermined. Asexual morph: Colonies
superficial, effuse, gregarious, white. Mycelium composed
of partly immersed, partly superficial, hyaline to pale brown,
septate hyphae, with mass of crowded conidia. Conidiophores long or short, uneven in width, 50–260 × 4–5.5 µm (x̄
= 147 × 4.9 µm, n = 10), macronematous, mononematous,
unbranched, erect, cylindrical, tapering at tip, pale brown,
septate, slightly constricted at septa, smooth-walled, thinwalled. Conidiogenous cells 13–17 × 4–5.5 µm ( x̄ = 15 ×
4.8 µm, n = 5), holoblastic, mono- to polyblastic, integrated,
determinate or sympodial, mostly intercalary or rarely terminal, cylindrical, with tiny tooth-like protrusions, mostly
bearing one, rarely two tiny conidiogenous loci, 4 × 1.5 µm,
pale brown, smooth-walled, loosely arranged. Conidia solitary, dry, mostly pleurogenous or rarely acrogenous, helicoid, rounded at tip, 47–52 µm diam. when tightly coiled,
55–70 µm diam. when loosely coiled, and conidial filament
3–5 µm wide ( x̄ = 50 × 4 µm, n = 5), 340–400 µm long,
tightly coiled 3–3.25 times, multi-septate, guttulate, hyaline,
smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 60 mm in 60 days at 25 °C, black from above and below,
surface rough, with sparse mycelium, dry, edge entire.
Mycelium mostly immersed, dark brown, septate, smooth.
Fungal Diversity (2020) 105:319–575
533
Fig. 124 Neohelicosporium
submersum (MFLU 17-1734,
holotype). a Colonies on submerged wood. b Conidiophores.
c Conidiophores with conidium.
d–f Conidiogenous cells. g, h
Conidia. Scale bars: b, c, h = 40
μm, d, g = 20 μm, e, f = 10 μm
Material examined: THAILAND, Phang Nga Province,
on submerged wood in a stream, 1 September 2017, X.D.
Yu, v7 (MFLU 17-1734, holotype), ex-type living culture
MFLUCC 17-2376.
Notes: Neohelicosporium submersum clusters with two
unnamed Neohelicosporium spp. (HKUCC 10235 and CBS
189.95) (Fig. 129). Our isolate MFLUCC 17-2376 is identical to HKUCC 10235 and CBS 189.95 based on available
sequences (LSU of HKUCC 10235 and ITS, LSU, SSU
of CBS 189.95). We introduce N. submersum, but do not
include these two strains because their morphological data
and some molecular data are lacking. Neohelicosporium submersum differs from another phylogenetically related species
N. krabiense by longer conidiophores (50–260 × 4–5.5 µm
vs. 45–140 × 5–6.5 µm) and larger conidia (47–52 µm diam.,
340–400 µm long vs. 30–40 µm diam., 230–320 µm long).
*Neohelicosporium taiwanense (C.H. Kuo & Goh) Y.Z. Lu
& K.D. Hyde, Fungal Diversity 92: 246 (2018)
Basionym: Helicosporium taiwanense C.H. Kuo & Goh,
Mycol. Progr. 17(5): 563 (2018)
Freshwater distribution: Taiwan, China (Kuo and Goh
2018)
*Neohelicosporium thailandicum Y.Z. Lu, J.C. Kang &
K.D. Hyde, Mycol. Progr. 17(5): 641(2017), [11] (2017)
Freshwater distribution: Thailand (Lu et al. 2018a)
Key to freshwater Neohelicosporium species
1.
1.
2.
Conidiophores micronematous………………………2
Conidiophores macronematous………………………4
Conidia > 25 μm diam.………………N. hyalosporum
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2.
3.
3.
4.
combinations based on morphology and multigene phylogenetic analyses, and designed P. talbotii as the type. We
follow Jayasiri et al. (2019) and conserve Pseudohelicomyces Y.Z. Lu, J.K. Liu & K.D. Hyde (Tubeufiaceae) against
Pseudohelicomyces Garnica & Valenz. (Hymenogastraceae).
Three species of Pseudohelicomyces were found from
freshwater habitats, viz. P. aquaticus Y.Z. Lu et al., P. hyalosporus Y.Z. Lu et al. and P. talbotii. Pseudohelicomyces
hyalosporus was formerly named as Helicomyces roseus
that comprises six isolates, of which five (Luo et al. 2017)
are from freshwater habitats and the other (CBS 283.51)
from terrestrial habitats was wrongly identified (Tsui et al.
2006b). The specimen HKAS 83995 of Luo et al. (2017)
was designated as the holotype of P. hyalosporus by Lu et al.
(2018b). Pseudohelicomyces talbotii was found from freshwater habitats by Lu et al. (2018b), with LSU, ITS, TEF and
RPB2 sequence data.
Conidia < 25 μm diam.………………………………3
Conidiophores up to 420 μm long……N. parvisporum
Conidiophores 50–280 μm long………N. astrictum
Conidiophores two different ranges of length…………
…………………………………………N. irregulare
4. Conidiophores only one range of length………………5
5. Conidiophores up to 74 μm long……N. thailandicum
5. Conidiophores > 75 μm long…………………………6
6. Holomorph……………………………N. ellipsoideum
6. Asexual morph………………………………………7
7. Conidia < 32 μm diam.…………………N. aquaticum
7. Conidia > 32 μm diam.………………………………8
8. Conidial filament > 5.5 μm wide………………………
……………………………………N. acrogenisporum
8. Conidial filament < 5.5 μm wide……………………9
9. Maximum length of conidiophores < 150 μm long……
…………………………………………N. krabiense
9. Maximum length of conidiophores > 150 μm long……
………………………………………………………10
10. Conidiogenous cells 10.5–16.5 μm long………………
………………………………………N. guangxiense
10. Conidiogenous cells 2.2−3.7 μm long…………………
…………………………………………N. taiwanense
Neotubeufia Chaiwan, Boonmee, Y.Z. Lu & K.D. Hyde,
Mycosphere 8(9): 1149 (2017)
Saprobic on submerged wood. Description of this genus
see Chaiwan et al. (2017).
Type species: Neotubeufia krabiensis Chaiwan, Boonmee,
Y.Z. Lu & K.D. Hyde
Notes: The monotypic genus Neotubeufia was introduced
by Chaiwan et al. (2017) for a freshwater sexual species N.
krabiensis producing superficial, glabrous ascomata, cylindrical asci and fasciculate ascospores. Neotubeufia formed
a distinct clade in Tubeufiaceae (Chaiwan et al. 2017; Lu
et al. 2018b).
List of freshwater Neotubeufia species
*Neotubeufia krabiensis Chaiwan, Boonmee, Y.Z. Lu &
K.D. Hyde, Mycosphere 8(9): 1149 (2017)
Freshwater distribution: Thailand (Chaiwan et al. 2017)
Pseudohelicomyces Y.Z. Lu, J.K. Liu & K.D. Hyde, Fungal
Diversity 92: 248 (2018)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Pseudohelicomyces talbotii (Goos) Y.Z. Lu
& K.D. Hyde
Notes: Pseudohelicomyces was proposed by Valenzuela
and Garnica (2000) with P. albus Garnica & E. Valenz.
as the type species. Lu et al. (2018b) formally introduced
this genus to accommodate two new species and three new
13
List of freshwater Pseudohelicomyces species
*Pseudohelicomyces aquaticus Y.Z. Lu, Boonmee & K.D.
Hyde, Fungal Diversity 92: 250 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Pseudohelicomyces hyalosporus Y.Z. Lu, J.K. Liu & K.D.
Hyde, Fungal Diversity 92: 251 (2018)
Freshwater distribution: China (Luo et al. 2017)
*Pseudohelicomyces talbotii (Goos) Y.Z. Lu & K.D. Hyde,
Fungal Diversity 92: 252 (2018)
Basionym: Helicosporium talbotii Goos, Mycologia
81(3): 368 (1989)
Freshwater distribution: Thailand (Lu et al. 2018b)
Key to freshwater Pseudohelicomyces species
1. Conidiophores micronematous, conidiogenous cells
monoblastic……………………………P. hyalosporus
1. Conidiophores macronematous, conidiogenous cells
mono- to polyblastic, sympodial………………………2
2. Conidia coiled 2.5–3.5 times………………P. aquaticus
2. Conidia coiled 1.5–2.5 times………………P. talbotii
Pseudohelicoon Y.Z. Lu & K.D. Hyde, Fungal Diversity
92: 254 (2018)
Saprobic on submerged wood. Description of this genus
see Lu et al. (2018b).
Type species: Pseudohelicoon subglobosum (Goh & C.H.
Kuo) Y.Z. Lu & K.D. Hyde
Basionym: Helicoon subglobosum Goh & C.H. Kuo [as
‘Helicoön’], Phytotaxa 346(2): 145 (2018)
Fungal Diversity (2020) 105:319–575
Notes: Pseudohelicoon is characterized by dark brown,
glistening colonies, short branched conidiophores, determinate conidiogenous cells and helicoid conidia that are tightly
coiled 7–13 times to form a subglobose or broadly elliptical
body, usually incorporating with mucilaginous substances
(Lu et al. 2018b). The type species P. subglobosum was
transferred from Helicoon to avoid further taxonomic confusion as the polyphyletic nature of Helicoon (Tsui and Berbee
2006; Goh and Kuo 2018; Lu et al. 2018b). Two freshwater
species are accepted in Pseudohelicoon and formed a wellsupported clade in Tubeufiaceae (Lu et al. 2018b). They can
be separated by the conidial shape and length (Goh and Kuo
2018).
List of freshwater Pseudohelicoon species
*Pseudohelicoon gigantisporum (Goh & K.D. Hyde) Y.Z.
Lu & J.K. Liu, Fungal Diversity 92: 254 (2018)
Basionym: Helicoon gigantisporum Goh & K.D. Hyde,
Mycol. Res. 100(12): 1487 (1996)
Freshwater distribution: Australia (Goh and Hyde 1996)
*Pseudohelicoon subglobosum (Goh & C.H. Kuo) Y.Z. Lu
& K.D. Hyde, Fungal Diversity 92: 255 (2018)
Basionym: Helicoon subglobosum Goh & C.H. Kuo [as
‘Helicoön’], Phytotaxa 346(2): 145 (2018)
Freshwater distribution: Taiwan, China (Goh and Kuo
2018)
Key to freshwater Pseudohelicoon species
1. Conidia broadly ellipsoidal, 78−123 μm long…………
………………………………………P. gigantisporum
1. Conidia subglobose, 66−78 μm long……………………
…………………………………………P. subglobosum
Thaxteriellopsis Sivan., Panwar & S.J. Kaur, Kavaka 4: 39
(1977) [1976]
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Lu et al. (2018b).
Type species: Thaxteriellopsis lignicola Sivan., Panwar
& S.J. Kaur
Notes: Boonmee et al. (2011) resurrected Thaxteriellopsis
based on morphology and phylogeny, and designated an epitype for the type species, T. lignicola. The sexual and asexual morphs of T. lignicola were described by Boonmee et al.
(2011) based on an epitype specimen and ex-epitype culture.
Doilom et al. (2017) reported its asexual morph from decaying inner surface of bark of Tectona grandis based on molecular evidence. Lu et al. (2018b) found another collection of
this species from submerged wood in freshwater habitats and
gave an updated description of sexual and asexual morphs.
535
List of freshwater Thaxteriellopsis species
*Thaxteriellopsis lignicola Sivan., Panwar & S.J. Kaur,
Kavaka 4: 39 (1977) [1976]
Freshwater distribution: Thailand (Lu et al. 2018b)
Tubeufia Penz. & Sacc., Malpighia 11(11–12): 517 (1898)
[1897]
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see (Lu et al. 2018b).
Type species: Tubeufia javanica Penz. & Sacc.
Notes: Tubeufia is an old genus comprising over 80
epithets in Index Fungorum (2020). After reassessment
of Tubeufiales, Lu et al. (2018b) accepted 50 species and
excluded T. helicomyces Höhn. based on multigene phylogeny and morphology. Both sexual and asexual morphs of
Tubeufia have various morphology as detailed in Lu et al.
(2018b). Tubeufia species are rather difficult to identify
without molecular data (Lu et al. 2018b). Recently, 17 new
species and six new combinations were added to the genus
(Lu et al. 2018b). Tubeufia is one of the largest freshwater
dothideomycetous genera comprising 29 freshwater species
(see list below). Most species were confirmed in Tubeufia
with molecular data, except T. claspisphaeria Kodsueb and
T. sympodiophora (Matsush.) Y.Z. Lu & K.D. Hyde. We
provide a key to freshwater Tubeufia species, which have
been studied well with molecular data.
List of freshwater Tubeufia species
*Tubeufia abundata Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal Diversity 92: 259 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia aquatica Z.L. Luo, Bhat & K.D. Hyde, Cryptog.
Mycol. 38(1): 44 (2017); Fig. 125
Freshwater distribution: China (Luo et al. 2017), Thailand (Lu et al. 2018b; this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum
superficial, effuse, gregarious, white. Mycelium composed
of partly immersed, hyaline to pale brown, septate, abundantly branched hyphae, with masses of crowded, glistening
conidia. Conidiophores 30–60 × 5–6 μm ( x̄ = 48.5 × 5.5
µm, n = 10), macronematous, mononematous, straight or
slightly curved, cylindrical, unbranched, 2–5-septate, brown
at the base, gradually paler upwards, sometimes swollen at
the base, smooth-walled. Conidiogenous cells 9.5–16 ×
4–5 µm ( x̄ = 12.5 × 4 µm, n = 10), holoblastic, polyblastic,
integrated, sympodial, terminal, subcylindrical or irregular,
truncate at the apex after conidial secession, denticulate,
pale brown, smooth-walled. Conidia solitary, acrogenous
or acropleurogenous, helicoid, rounded at the tip, 23–26
μm diam. and conidial filament 3–5 μm wide ( x̄ = 25.5 ×
13
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Fig. 125 Tubeufia aquatica
(MFLU 15-2712). a Colonies
on submerged wood. b–d Conidiophores with conidia. e–h
Conidia. i, j Conidiophores with
conidiogenous cells. k Germinated conidium. l, m Colony on
PDA (up-front, down-reverse).
Scale bars: a = 100 μm, b = 20
μm, c, e–k = 10 μm, d = 5 μm
3.8 μm, n = 10), 100–165 μm long, coiled 2.25–3.5 times,
distinctly multi-septate, guttulate, hyaline, smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 10 mm in 22 days at 25 °C, dark brown from above
and below, surface rough, with sparse superficial mycelium,
most mycelium immersed in culture, dry, edge entire.
13
Material examined: THAILAND, Prachuap Khiri
Khan Province, on submerged wood in a stream, 30 July
2015, K.D. Hyde, 143A (MFLU 15-2712), living culture
MFLUCC 15-0990.
Notes: Our new collection MFLUCC 15-0990 clusters
in the same clade with three freshwater strains of Tubeufia
Fungal Diversity (2020) 105:319–575
aquatica with high bootstrap support, which indicates that
they are the same species (Fig. 129). Tubeufia aquatica have
been collected from submerged wood in China and Thailand
(Luo et al. 2017; Lu et al. 2018b). They have very similar
morphological characters, except that the collection of Lu
et al. (2018b) has less septa (0–1-septate) of conidiophores
than the holotype (> 4-septate) in Luo et al. (2017) and our
537
new collection (2–5-septate). We identify our new collection
MFLUCC 15-0990 as T. aquatica based on morphology and
phylogenetic analysis (Fig. 129).
*Tubeufia brevis Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 263 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
Fig. 126 Tubeufia cylindrothecia (MFLU 11-0969). a
Colonies on submerged wood.
b–e Conidiophores, conidiogenous cells and conidia. f–h
conidia. i Germinated conidium.
j, k Colony on PDA (left-front,
right-reverse). Scale bars: b, i =
20 μm, c–h = 10 μm
13
538
Fungal Diversity (2020) 105:319–575
Fig. 127 Tubeufia cylindrothecia (MFLU 11-1107). a
Colonies on submerged wood.
b–d Conidiophores, conidiogenous cells, conidia. e, f conidia.
g Germinated conidium. h,
i Colony on PDA (up-front,
down-reverse). Scale bars: b,
e–g = 20 μm, c = 5 μm, d =
10 μm
*Tubeufia brunnea Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal Diversity 92: 265 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia chlamydospora Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 92: 266 (2018)
13
Freshwater distribution: Thailand (Lu et al. 2018b)
Tubeufia claspisphaeria Kodsueb, Mycologia 96(3): 668
(2004)
Freshwater distribution: Hong Kong, China (Kodsueb
et al. 2004)
Fungal Diversity (2020) 105:319–575
*Tubeufia cylindrothecia (Seaver) Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 128(7–8): 562 (1919);
Figs. 126, 127
Basionym: Ophionectria cylindrothecia Seaver, Mycologia 1(2): 70 (1909)
539
Freshwater distribution: China (Luo et al. 2017), Thailand (Lu et al. 2018b; this study)
Saprobic on decaying wood in terrestrial and freshwater
habitats. Sexual morph: Undetermined. Asexual morph:
Colonies superficial, effuse, gregarious, white to pale brown.
Mycelium composed of partly immersed, hyaline to pale
Fig. 128 Tubeufia tectonae
(MFLU 15-2696). a Colonies
on submerged wood. b, c
Conidiophores with conidia. d
Conidiophore with conidiogenous cell. e Conidial filament.
f–i Conidia. j Germinated
conidium. k, l Colony on PDA
(up-front, down-reverse). Scale
bars: a = 100 μm, b, c = 20 μm,
d, e = 5 μm, f–j = 50 μm
13
540
brown, septate, sparsely branched hyphae, with masses of
conidia. Conidiophores 60–140 × 3–5 μm ( x̄ = 108 × 3.8
µm, n = 10), macronematous, mononematous, cylindrical,
septate, branched or unbranched, erect, flexuous, pale brown
to brown, smooth-walled. Conidiogenous cells 10.5–18 ×
3.5–4 μm ( x̄ = 15.5 × 3.8 µm, n = 5), holoblastic, monoto poly-blastic, integrated, terminal, smooth, truncate at the
apex after conidial secession, each with single or several
conidia. Conidia solitary, acrogenous, helicoid, rounded
at tip, 24.5–28 μm diam. and conidial filament 3–4.2 μm
wide ( x̄ = 26 × 3.6 μm, n = 10), 100–210 μm long, coiled
2.75–3.25 times, indistinctly multi-septate, guttulate, hyaline
to pale brown, smooth-walled.
Culture characteristics: On PDA, colony irregular, dark
brown to black from above and below, surface rough, with
dense superficial or immersed mycelium, most mycelium
immersed in medium, dry, edge undulate.
Material examined: THAILAND, Chiang Mai Province, on submerged wood in a stream, 29 November. 2010,
H. Zhang, d6 (MFLU 11-0969), living culture MFLUCC
10-0919; ibid., d29 (MFLU 11-1107), living culture
MFLUCC 11-0076.
Notes: Our two new collections MFLUCC 10-0919 and
MFLUCC 11-0076 are identified as Tubeufia cylindrothecia
based on the identical ITS and LSU sequence data to the extype strain of T. cylindrothecia, and phylogenetic analysis
(Fig. 128). The asexual morph of T. cylindrothecia was first
reported by Luo et al. (2017) and later encountered by Lu
et al. (2018b) from freshwater habitats. Our freshwater collections have longer conidiophores (60–140 μm vs. 50–81
μm) and shorter conidia (100–210 μm vs. 256–314 μm) than
the holotype (Luo et al. 2017). Additionally, we found polyblastic conidiogenous cells in MFLU 11-0969, while they
are only monoblastic in the holotype (Luo et al. 2017).
*Tubeufia dictyospora Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 92: 271 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia eccentrica Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal Diversity 92: 271 (2018)
Freshwater distribution: China (Lu et al. 2018b)
*Tubeufia fangchengensis Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 273 (2018)
Freshwater distribution: China (Lu et al. 2018b)
*Tubeufia filiformis Y.Z. Lu, Boonmee & K.D. Hyde,
Mycol. Progr. 16(4): 407 (2017)
Freshwater distribution: Thailand (Lu et al. 2017a,
2018b)
13
Fungal Diversity (2020) 105:319–575
*Tubeufia geniculata (C.H. Kuo & Goh) Y.Z. Lu & K.D.
Hyde, Fungal Diversity 92: 297 (2018)
Basionym: Helicomyces geniculatus C.H. Kuo & Goh,
Mycoscience 59(5): 436 (2018)
Freshwater distribution: Taiwan, China (Lu et al. 2018b)
*Tubeufia guangxiensis Chaiwan, Boonmee, Y.Z. Lu &
K.D. Hyde, Mycosphere 8(9): 1150 (2017)
Freshwater distribution: China (Chaiwan et al. 2017; Lu
et al. 2018b)
*Tubeufia hechiensis Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 274 (2018)
Freshwater distribution: China (Lu et al. 2018b)
*Tubeufia hyalospora Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 80: 126 (2016)
Freshwater distribution: Thailand (Hyde et al. 2016b)
*Tubeufia inaequalis Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 278 (2018)
Freshwater distribution: China (Lu et al. 2018b), Thailand (Lu et al. 2018b)
*Tubeufia krabiensis Y.Z. Lu, Boonmee & K.D. Hyde, Fungal Diversity 92: 279 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia latispora Y.Z. Lu, Boonmee & K.D. Hyde,
Mycol. Progr. 16(4): 409 (2017)
Freshwater distribution: Thailand (Lu et al. 2017a)
*Tubeufia laxispora Y.Z. Lu, Boonmee & K.D. Hyde,
Mycol. Progr. 16(4): 409 (2017)
Freshwater distribution: Thailand (Lu et al. 2017a,
2018b)
*Tubeufia mackenziei Y.Z. Lu, Boonmee & K.D. Hyde,
Mycol. Progr. 16(4): 411 (2017)
Freshwater distribution: Thailand (Lu et al. 2017a)
*Tubeufia parvispora Tibpromma & K.D. Hyde, Fungal
Diversity 93: 74 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia rubra Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 285 (2018)
Freshwater distribution: China (Lu et al. 2018b)
*Tubeufia sympodihylospora Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 289 (2018)
Freshwater distribution: China (Lu et al. 2018b)
Fungal Diversity (2020) 105:319–575
Fig. 129 Phylogram generated from maximum likelihood
analysis of combined ITS, LSU,
TEF and RPB2 sequence data
for species of Tubeufiales. Bootstrap values for maximum likelihood equal to or greater than
75% and Bayesian posterior
probabilities equal to or greater
than 0.95 are placed near the
branches as ML/BYPP. Newly
generated sequences are in red
and ex-type strains are in bold.
The new species introduced in
this study are indicated with
underline. Freshwater strains are
indicated with a red letter “F”.
The tree is rooted to Botryosphaeria agaves MFLUCC
10-0051 and B. dothidea CBS
115476 (Botryosphaeriaceae)
541
Tubeufia guangxiensis MFLUCC 17-0038 F
Tubeufia guangxiensis MFLUCC 17-0045 F
Tubeufia guangxiensis GZCC 16-0091 F
Tubeufia guangxiensis GZCC 16-0054 F
Tubeufia guangxiensis GZCC 16-0090 F
100/1.00
Tubeufia guangxiensis MFLUCC 17-0046 F
Tubeufia guangxiensis GZCC 16-0041 F
100/1.00
Tubeufia eccentrica MFLUCC 17-1524 F
100/1.00
Tubeufia eccentrica GZCC 16-0048 F
Tubeufia eccentrica GZCC 16-0084 F
98/0.99
Tubeufia eccentrica GZCC 16-0035 F
99/1.00
Tubeufia sympodihylospora MFLUCC 17-0044 F
Tubeufia sympodihylospora GZCC 16-0049 F
98/1.00
100/1.00
Tubeufia sympodihylospora GZCC 16-0051 F
100/1.00
Tubeufia xylophila MFLUCC 17-1520 F
Tubeufia xylophila GZCC 16-0038 F
Tubeufia tratensis MFLUCC 17-1993 F
100/1.00 88/-- Tubeufia inaequalis MFLUCC 17-0053 F
Tubeufia inaequalis GZCC 16-0079 F
Tubeufia inaequalis GZCC 16-0087 F
100/1.00
Tubeufia inaequalis MFLUCC 17-1998 F
Tubeufia inaequalis BCC 8808
Tubeufia inaequalis MFLUCC 17-1989 F
100/1.00
Tubeufia rubra GZCC 16-0081 F
100/1.00 Tubeufia rubra GZCC 16-0083 F
Tubeufia krabiensis MFLUCC 16-0228 F
Tubeufia filiformis MFLUCC 16-1128 F
Tubeufia filiformis MFLUCC 16-0236 F
100/1.00
Tubeufia filiformis MFLUCC 16-1135 F
Tubeufia hyalospora MFLUCC 15-1250 F
90/0.95
100/-- Tubeufia geniculata NCYU U2-1B F
Tubeufia geniculata BCRC FU30849 F
100/1.00 Tubeufia abundata MFLUCC 17-2024 F
Tubeufia tectonae MFLUCC 12-0392
99/1.00
Tubeufia tectonae MFLUCC 16-0235 F
Tubeufia tectonae MFLUCC 17-1985 F
Tubeufia tectonae MFLUCC 15-1247 F
100/1.00 Tubeufia tectonae MFLUCC 15-0974 F
Tubeufia tectonae MFLUCC 16-0230 F
Tubeufia tectonae MFLUCC 17-1797 F
Tubeufia sessilis MFLUCC 16-0021
Tubeufia bambusicola MFLUCC 17-1803
100/1.00
91/1.00
Tubeufia latispora MFLUCC 16-0027 F
Tubeufia longiseta MFLUCC 15-0188
Tubeufia javanica MFLUCC 12-0545
Tubeufia cylindrothecia MFLUCC 16-1283 F
Tubeufia cylindrothecia MFLUCC 16-1253 F
77/1.00
Tubeufia cylindrothecia DLUCC 0572 F
Tubeufia cylindrothecia MFLUCC 17-1792 F
90/1.00 Tubeufia cylindrothecia MFLUCC 11-0076 F
76/0.99 Tubeufia cylindrothecia BCC 3585 F
100/1.00
Tubeufia cylindrothecia MFLUCC 10-0919 F
Tubeufia cylindrothecia BCC 3559
Tubeufia mackenziei MFLUCC 16-0222 F
100/1.00
95/1.00 Tubeufia lilliputea NBRC 32664
Tubeufia brevis MFLUCC 17-1799 F
99/1.00
*Tubeufia sympodilaxispora Y.Z. Lu, J.C. Kang & K.D.
Hyde, Fungal Diversity 92: 289 (2018)
Freshwater distribution: China (Lu et al. 2018b)
Tubeufia
Basionym: Helicoma sympodiophorum Matsush. [as
‘sympodiophora’], Matsush. Mycol. Mem. 7: 52 (1993)
Freshwater distribution: Hong Kong, China (Ho et al.
2002)
Tubeufia sympodiophora (Matsush.) Y.Z. Lu & K.D. Hyde,
Fungal Diversity 92: 299 (2018)
13
542
Fungal Diversity (2020) 105:319–575
Fig. 129 (continued)
100/-97/0.99
Tubeufia sympodilaxispora MFLUCC 17-0048 F
Tubeufia sympodilaxispora GZCC 16-0058 F
89/1.00
Tubeufia sympodilaxispora BCC 3580
Tubeufia chiangmaiensis MFLUCC 17-1801
100/1.00 Tubeufia chiangmaiensis MFLUCC 11-0514
100/1.00 Tubeufia parvispora MFLUCC 17-2009
100/1.00
Tubeufia parvispora MFLUCC 17-2003 F
Tubeufia parvispora MFLUCC 17-1992 F
96/0.99 Tubeufia dictyospora MFLUCC 17-1805
85/1.00
Tubeufia dictyospora MFLUCC 16-0220 F
100/1.00
Tubeufia machaerinae MFLUCC 17-0055
Tubeufia chlamydospora MFLUCC 16-0223 F
99/1.00
Tubeufia fangchengensis MFLUCC 17-0047 F
Tubeufia laxispora MFLUCC 16-0219 F
Tubeufia laxispora MFLUCC 16-0013 F
100/1.00
Tubeufia laxispora MFLUCC 16-0232 F
Tubeufia laxispora MFLUCC 17-2023 F
80/1.00
Tubeufia aquatica MFLUCC 15-0990 F
Tubeufia aquatica MFLUCC 17-1794 F
100/1.00
Tubeufia aquatica DLUCC 0574 F
79/0.99
Tubeufia aquatica MFLUCC 16-1249 F
Tubeufia taiwanensis BCRC FU30844 F
100/1.00
Tubeufia hechiensis MFLUCC 17-0052 F
Tubeufia brunnea MFLUCC 17-2022 F
100/1.00
Helicotruncatum palmigenum MFLUCC 15-0993 F
92/0.98
Helicotruncatum palmigenum NBRC 32663
Pseudohelicomyces hyalosporus MFLUCC 15-0343 F
Pseudohelicomyces hyalosporus KUMCC 15-0322 F
Pseudohelicomyces hyalosporus KUMCC 15-0411 F
100/1.00
Pseudohelicomyces hyalosporus KUMCC 15-0430 F
Pseudohelicomyces hyalosporus CBS 283.51 F
81/1.00
100/1.00
Pseudohelicomyces hyalosporus KUMCC 15-0281 F
Pseudohelicomyces talbotii MFLUCC 17-2021 F
Pseudohelicomyces talbotii MUCL 33010
93/0.98
Pseudohelicomyces aquaticus MFLUCC 16-0234 F
93/1.00
99/1.00
Pseudohelicomyces paludosus CBS 120503 / AR4206
Pseudohelicomyces indicus CBS 374.93
96/1.00 Helicomyces hyalosporus GZCC 16-0075 F
95/-Helicomyces hyalosporus GZCC 16-0073 F
100/1.00
Helicomyces hyalosporus GZCC 16-0070 F
80/-Helicomyces hyalosporus MFLUCC 17-0051 F
100/1.00
Helicomyces chiayiensis BCRC FU30842 F
Helicomyces torquatus MFLUCC 16-0217 F
99/1.00
Helicomyces colligatus MFLUCC 16-1132 F
90/1.00 Neohelicomyces pallidus CBS 962.69
100/1.00
Neohelicomyces pallidus CBS 271.52
98/1.00
Neohelicomyces pallidus CBS 245.49
Neohelicomyces dehongensis MFLUCC 18-1029 F
100/1.00
Neohelicomyces hyalosporus GZCC 16-0086 F
--/0.99
Neohelicomyces aquaticus MFLUCC 16-0993 F
Neohelicomyces aquaticus KUMCC 15-0463 F
Neohelicomyces grandisporus KUMCC 15-0470 F
100/1.00
Neohelicomyces deschampsiae CBS 145029
Neohelicomyces pallidus UAMH 10535
Neohelicomyces pandanicola KUMCC 16-0143
94/0.97
Neohelicomyces thailandicus MFLUCC 11-0005 F
87/0.99
Neohelicomyces submersus MFLUCC 16-1106 F
85/1.00
Tubeufia helicomyces MUCL 15702
Tubeufia amazonensis ATCC 42524
95/0.99
13
Tubeufia
Helicotruncatum
Pseudohelicomyces
Helicomyces
Neohelicomyces
Fungal Diversity (2020) 105:319–575
Fig. 129 (continued)
543
Muripulchra aquatica MFLUCC 15-0249 F
Muripulchra aquatica DLUCC 0571 F
95/-- Muripulchra aquatica KUMCC 15-0245 F
Muripulchra aquatica KUMCC 15-0276 F
100/0.99 Neohelicosporium guangxiense MFLUCC 17-0050 F
79/-- Neohelicosporium guangxiense GZCC 16-0067 F
90/-- Neohelicosporium guangxiense GZCC 16-0077 F
Neohelicosporium guangxiense MFLUCC 17-1522 F
99/1.00
Neohelicosporium guangxiense GZCC 16-0068 F
83/0.97
Neohelicosporium guangxiense GZCC 16-0042 F
88/1.00
Neohelicosporium guangxiense GZCC 16-0089 F
Neohelicosporium guangxiense MFLUCC 17-0054 F
Neohelicosporium panacheum CBS 257.59
78/1.00
Neohelicosporium irregulare MFLUCC 17-1808
Neohelicosporium irregulare MFLUCC 17-1796 F
95/1.00
94/1.00
Neohelicosporium taiwanense BCRC FU30841 F
Neohelicosporium laxisporum MFLUCC 17-2027
97/1.00 Neohelicosporium thailandicum MFLUCC 16-0221 F
87/1.00
Neohelicosporium aquaticum MFLUCC 17-1519 F
Neohelicosporium ellipsoideum MFLUCC 16-0229 F
Neohelicosporium acrogenisporum MFLUCC 17-2019 F
Neohelicosporium abuense CBS 101688
Neohelicosporium sp. HKUCC 10235
--/0.96
Neohelicosporium sp. CBS 189.95
Neohelicosporium submersum MFLUCC 17-2376 F
Neohelicosporium krabiense MFLUCC 16-0224 F
97/1.00 Neohelicosporium parvisporum GZCC 16-0100 F
Neohelicosporium parvisporum GZCC 16-0078 F
--/0.99
Neohelicosporium parvisporum MFLUCC 17-1793 F
Neohelicosporium parvisporum MFLUCC 17-1521 F
75/-Neohelicosporium parvisporum MFLUCC 17-1523 F
Neohelicosporium parvisporum MFLUCC 17-2010
--/1.00
Neohelicosporium parvisporum MFLUCC 17-1995 F
Neohelicosporium parvisporum MFLUCC 16-0218 F
99/0.96
Neohelicosporium parvisporum MFLUCC 17-1804
Neohelicosporium parvisporum MFLUCC 17-1807
Neohelicosporium fusisporum MFUCC 16-0642
Neohelicosporium astrictum MFLUCC 17-2004 F
89/-- Neohelicosporium griseum CBS 961.69
81/-Neohelicosporium griseum UAMH 1694
100/1.00
Neohelicosporium griseum JCM 9265
Neohelicosporium griseum CBS 113542
97/1.00
Neohelicosporium aurantiellum ANM 718
100/1.00
Neohelicosporium morganii CBS 281.54
Neohelicosporium morganii CBS 222.58
Boerlagiomyces macrospora MFLUCC 12-0388
100/1.00
Neohelicosporium hyalosporum GZCC 16-0076 F
97/1.00
Neohelicosporium hyalosporum GZCC 16-0063
95/1.00
99/1.00
Neohelicosporium ovoideum GZCC 16-0066
Neohelicosporium ovoideum GZCC 16-0064
98/0.97
100/1.00
*Tubeufia taiwanensis Y.Z. Lu & K.D. Hyde, Fungal Diversity 92: 291 (2018)
Freshwater distribution: Taiwan, China (Lu et al. 2018b)
*Tubeufia tectonae Doilom & K.D. Hyde, Fungal Diversity
82: 150 (2017); Fig. 128
Synonymy: Tubeufia roseohelicospora Y.Z. Lu, Boonmee
& K.D. Hyde, Fungal Diversity 80: 128 (2016)
Freshwater distribution: Thailand (Hyde et al. 2016b; Lu
et al. 2018b; this study)
Saprobic on submerged wood. Sexual morph: Undetermined. Asexual morph: Colonies on natural substratum
Muripulchra
Neohelicosporium
superficial, effuse, gregarious, hyaline to white. Mycelium
composed of partly immersed, hyaline to pale brown, septate
hyphae, with masses of crowded, glistening conidia. Conidiophores 40–60 × 5–7 μm, macronematous, mononematous,
erect, cylindrical, unbranched, septate, brown, smoothwalled. Conidiogenous cells 15–20 × 4–5.5 µm, holoblastic,
polyblastic, integrated, sympodial, terminal or intercalary,
subcylindrical, denticulate, brown, smooth-walled. Conidia
solitary, acrogenous or acropleurogenous, helicoid, rounded
at tip, 80–130 μm diam. and conidial filament 3.5–6.5 μm
wide ( x̄ = 105 × 5 μm, n = 10), 370–450 μm long, coiled
13
544
Fungal Diversity (2020) 105:319–575
Fig. 129 (continued)
93/1.00
83/1.00
Neochlamydotubeufia khunkornensis MFLUCC 16-1129 F
Neochlamydotubeufia khunkornensis MFLUCC 10-0118
Neochlamydotubeufia khunkornensis MFLUCC 16-0025 F
Neochlamydotubeufia khunkornensis MFLUCC 16-1126 F
Neochlamydotubeufia fusiformis MFLUCC 16-0214 F
Neochlamydotubeufia fusiformis MFLUCC 16-0016 F
Neochlamydotubeufia khunkornensis MFLUCC 10-0117
Pseudohelicoon subglobosum BCRC FU30843 F
Pseudohelicoon subglobosum NCYU K3-2-3 F
Neochlamydotu
beufia
Pseudohelicoon subglobosum NCYU K3-2-2 F
Pseudohelicoon
Pseudohelicoon subglobosum NCYU K3-2-1 F
Pseudohelicoon gigantisporum BCC 3550
99/-Helicodochium aquaticum MFLUCC 16-0008 F
100/1.00
Helicodochium aquaticum MFLUCC 17-2016 F
Helicodochium
100/1.00
Helicodochium aquaticum MFLUCC 18-0490 F
100/0.99
Helicohyalinum aquaticum MFLUCC 16-0014 F
100/1.00
Helicohyalinum aquaticum MFLUCC 16-1131 F
Helicohyalinum
Helicohyalinum infundibulum MFLUCC 16-1133 F
79/-- Dematiohelicomyces helicosporus MFLUCC 16-0007 F
100/1.00 Dematiohelicomyces helicosporus MFLUCC 16-0213 F Dematiohelicomyc
es
Dematiohelicomyces helicosporus MFLUCC 16-0003 F
100/0.96 Helicotubeufia guangxiensis MFLUCC 17-0040 F
100/1.00 Helicotubeufia guangxiensis MFLUCC 17-0041 F
Helicotubeufia
100/1.00 Helicotubeufia hydei MFLUCC 17-1980
Helicotubeufia hydei MFLUCC 17-1986
100/1.00
Helicotubeufia jonesii MFLUCC 17-0043 F
Aquaphila albicans MFLUCC 15-0995 F
Aquaphila albicans BCC 3520
Aquaphila albicans BCC 3463
Aquaphila
Aquaphila albicans BCC 3543
Aquaphila
albicans
KUMCC
19-0078
F
99/1.00
100/1.00
Aquaphila albicans MFLUCC 16-0010 F
Aquaphila albicans MFLUCC 16-0020
Chlamydotubeufia huaikangplaensis MFLUCC 16-0227 F
Chlamydotubeufia huaikangplaensis MFLUCC 16-0023 F
100/1.00
Chlamydotubeufia huaikangplaensis MFLUCC 10-0926
100/1.00
Chlamydotubeufia
Chlamydotubeufia chlamydosporum CBS 160.69
100/1.00
Chlamydotubeufia krabiensis MFLUCC 16-1134 F
Chlamydotubeufia cylindrica MFLUCC 16-1130 F
Berkleasmium aquaticum GZCC 16-0056 F
94/-Berkleasmium aquaticum GZCC 16-0044 F
Berkleasmium aquaticum GZCC 16-0034 F
86/-- Berkleasmium aquaticum MFLUCC 17-0039 F
100/1.00
Berkleasmium aquaticum MFLUCC 17-0049 F
Berkleasmium aquaticum GZCC 16-0055 F
Berkleasmium aquaticum GZCC 16-0039 F
99/1.00
Berkleasmium fusiforme MFLUCC 17-1978
Berkleasmium fusiforme MFLUCC 17-1987
100/1.00
100/1.00
Berkleasmium fusiforme MFLUCC 17-1979
Berkleasmium
Berkleasmium fusiforme MFLUCC 17-1988
Berkleasmium guangxiense MFLUCC 17-0042 F
100/1.00
Berkleasmium concinnum ILLS 80805
Berkleasmium concinnum ILLS 59356
Berkleasmium concinnum ILLS 80803
95/1.00
Berkleasmium concinnum ILLS 80802
Berkleasmium concinnum ILLS 80804
100/1.00
Berkleasmium concinnum DAOM 696499
97/1.00
100/1.00
Berkleasmium concinnum NB 789
100/1.00
13
Fungal Diversity (2020) 105:319–575
545
Fig. 129 (continued)
Berkleasmium thailandicum MFLUCC 16-0012 F
Berkleasmium thailandicum MFLUCC 16-0015 F
Berkleasmium thailandicum MFLUCC 16-1127 F
98/1.00
Berkleasmium thailandicum MFLUCC 15-1248 F
100/1.00
Berkleasmium thailandicum MFLUCC 17-1984
Berkleasmium thailandicum MFLUCC 17-2000 F
100/1.00
Berkleasmium longisporum MFLUCC 17-2002 F
97/1.00
Berkleasmium longisporum MFLUCC 17-1999 F
100/1.00
Berkleasmium longisporum MFLUCC 17-1990 F
99/0.98 Berkleasmium latisporum MFLUCC 15-0987 F
Berkleasmium latisporum MFLUCC 16-0019
Dictyospora thailandica MFLUCC 11-0509
82/0.97
Dictyospora thailandica MFLUCC 16-0215 F
92/0.98
Dictyospora thailandica MFLUCC 11-0512
100/1.00
Dictyospora thailandica MFLUCC 18-0641 F
Dictyospora thailandica MFLUCC 16-0001 F
100/1.00
85/1.00
97/1.00
--/0.96
--/0.99
--/0.99
Helicoma longisporum MFLUCC 16-0002 F
Helicoma longisporum MFLUCC 16-0211 F
98/1.00
Helicoma longisporum MFLUCC 16-0005 F
Helicoma longisporum MFLUCC 17-1997 F
Helicoma khunkornensis MFLUCC 10-0119
--/0.99
96/-- Helicoma septoconstrictum MFLUCC 17-1991 F
Helicoma septoconstrictum MFLUCC 17-2001 F
100/1.00
Helicoma brunneisporum MFLUCC 17-1983
--/0.96
Helicoma rugosum ANM 1169
Helicoma rugosum JCM 2739
Helicoma rugosum ANM 953
86
Helicoma rugosum ANM 196
87/1.00
Helicoma sp. HKUCC 9118
Helicoma miscanthi MFLUCC 11-0375
78/0.99
Helicoma rubriappendiculatum MFLUCC 18-0491
Helicoma rufum MFLUCC 17-1806
100/1.00
97/1.00
Helicoma aquaticum MFLUCC 17-2025 F
Helicoma linderi NBRC 9207
Helicoma nematosporum MFLUCC 16-0011 F
100/1.00
Helicoma muelleri CBS 964.69
Helicoma muelleri UBC F13877
97/1.00
95/-Helicoma inthanonense MFLUCC 11-0003
100/1.00
Helicoma dennisii NBRC 30667
Helicoma guttulatum MFLUCC 16-0022 F
100/1.00 Helicoma ambiens UAMH 10534
Helicoma ambiens UAMH 10533
89/1.00
Helicoma fusiforme MFLUCC 17-1981
94/1.00
Helicoma multiseptatum GZCC 16-0080 F
Helicoma hongkongense MFLUCC 17-2005 F
100/1.00
Helicoma siamense MFLUCC 10-0120
Helicoma tectonae MFLUCC 12-0563
Helicoarctatus aquaticus MFLUCC 17-1996 F
Neotubeufia krabiensis MFLUCC 16-1125 F
1.25–2 times, becoming loosely coiled or uncoiled in water,
indistinctly multi-septate, guttulate, hyaline, smooth-walled.
Culture characteristics: On PDA, colony circular, reaching 30 mm in 20 days at 25 °C, grey to brown from above
and below, surface rough, with sparse superficial mycelium,
most mycelium immersed in culture, dry, edge undulate.
Material examined: THAILAND, Prachuap Khiri Khan
Province, on submerged wood in a stream, 30 July 2015,
K.D. Hyde, 34D (MFLU 15-2696), living culture MFLUCC
15-0974.
Notes: Tubeufia tectonae was initially collected from bark
of Tectona grandis and characterized by subhyaline to white
Berkleasmium
Dictyospora
Helicoma
Helicoarctatus
Neotubeufia
colonies, mostly loosely coiled conidia (Doilom et al. 2017).
Later, a new freshwater species T. roseohelicospora Y.Z. Lu
et al. was introduced based on light pink colonies, tightly
coiled conidia and wider conidial filaments (Hyde et al.
2016b). Lu et al. (2018b) found another two isolates of T.
roseohelicospora and two isolates of T. tectonae from freshwater habitats, with additional TEF sequence data. However,
all isolates of the two species, including our new collection
MFLU 15-2696, do not show phylogenetic difference in
the phylogenetic tree (Lu et al. 2018b; this study, Fig. 129).
Furthermore, they have nearly identical TEF sequence data
13
546
Fungal Diversity (2020) 105:319–575
f
Fig. 129 (continued)
Helicosporium vegetum CBS 269.52
Helicosporium vegetum CBS 254.75
Helicosporium vegetum NBRC 30345
99/1.00
Helicosporium vegetum CBS 941.72
76/0.99
Helicosporium sp. NBRC 9014
100/1.00 Helicosporium vesicarium MFLUCC 17-1795
Helicosporium flavisporum MFLUCC 17-2020 F
98/1.00 Helicosporium luteosporum MFLUCC 16-1233
Helicosporium luteosporum MFLUCC 16-0226
Helicosporium flavum MFLUCC 16-1230 F
Helicosporium thailandense MFLUCC 18-1407 F
90/0.99
85/-Helicosporium viridiflavum MFLUCC 17-2336
100/-- Helicosporium setiferum MFLUCC 17-1994 F
98/-Helicosporium setiferum MFLUCC 17-2006 F
100/1.00 Helicosporium setiferum BCC 8125
100/1.00
Helicosporium setiferum BCC 3332
100/1.00
Helicosporium setiferum MFLUCC 17-2007 F
Helicosporium aquaticum MFLUCC 17-2008 F
87/1.00
Helicoma vaccinii CBS 216.90
Thaxteriellopsis lignicola MFLUCC 10-0121
Thaxteriellopsis lignicola MFLUCC 16-0024 F
Thaxteriellopsis lignicola MFLUCC 10-0123
Thaxteriellopsis lignicola MFLUCC 10-0122
Thaxteriellopsis lignicola MFLUCC 16-0026 F
100/1.00
Thaxteriellopsis lignicola MFLUCC 10-0124
Thaxteriellopsis lignicola MFLUCC 15-0898
Botryosphaeria dothidea CBS 115476
Botryosphaeria agaves MFLUCC 10-0051
99/1.00
100/1.00
100/1.00
which indicates that they are conspecific. We, therefore, synonymize T. roseohelicospora with T. tectonae.
*Tubeufia tratensis Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal Diversity 92: 295 (2018)
Freshwater distribution: Thailand (Lu et al. 2018b)
*Tubeufia xylophila (P.N. Singh & S.K. Singh) Y.Z. Lu &
K.D. Hyde, Fungal Diversity 92: 299 (2018)
Basionym: Helicosporium xylophilum P.N. Singh & S.K.
Singh [as ‘xylophilous’], Curr. Res. Envir. & App. Myc.
6(4): 251 (2016)
Freshwater distribution: China (Lu et al. 2018b)
Key to freshwater Tubeufia species
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
Holomorph known……………………………………2
Holomorph unknown…………………………………4
Conidial length > 400 μm……………T. guangxiensis
Conidial length < 400 μm……………………………3
Conidia > 330 μm long…………………T. filiformis
Conidia < 330 μm long……………T. cylindrothecia
Sexual morph……………………………T. latispora
Asexual morph………………………………………5
Conidia non-helicoid…………………………………6
Conidia helicoid………………………………………7
Conidia produce chlamydospores, form small globose
to subglobose secondary conidia………………………
……………………………………T. chlamydospora
Conidia produce dictyospores, lack secondary
conidia…………………………………T. dictyospora
Conidiogenous cells of two types……………………8
Conidiogenous cells of one type………………………9
13
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
18.
18.
19.
19.
20.
20.
21.
Helicosporium
Thaxteriellopsis
Outgroup
Conidia < 40 μm diam.…………………T. abundata
Conidia > 40 μm diam.……………………T. tectonae
Colonies often with stalked sclerotia………………10
Colonies without stalked sclerotia…………………11
Setae absent……………………………T. taiwanensis
Setae present……………………………T. geniculata
Colonies red-brown when dried………………T. rubra
Colonies hyaline to white or pale brown……………12
Conidiophores unbranched…………………………13
Conidiophores branched……………………………17
Conidiogenous cells sympodial……………………14
Conidiogenous cells determinate……………………15
Conidiogenous cells truncate at the apex………………
……………………………………………T. xylophila
Conidiogenous cells denticulate…………T. aquatica
Conidiogenous cells polyblastic, denticulate…………
…………………………………………T. mackenziei
Conidiogenous cells monoblastic, each with single
conidium……………………………………………16
Conidia 17.5–40 μm diam., 111–182 μm long………
……………………………………………T. laxispora
Conidia 16–33 μm diam., 110–225 μm long…………
…………………………………………T. hyalospora
Colonies brown……………………………T. brunnea
Colonies hyaline to white or pale brown……………18
Conidiophores up to 210 μm long……………………
…………………………………………T. hechiensis
Conidiophores < 140 μm long………………………19
Conidia < 20 μm diam.…………………T. parvispora
Conidia > 20 μm diam.………………………………20
Conidiophores mostly > 300 μm long………………21
Conidiophores mostly < 300 μm long………………25
Conidiophores 400–570 μm long………T. eccentrica
Fungal Diversity (2020) 105:319–575
547
Fig. 130 Lepidopterella palustris (Material examined: USA,
Illinois, Johnson County, Elvira
Cypress Swamp, on submerged
twig, 3 August 1977, J.L.
Crane & J.D. Schoknecht, ILLS
42219, isotype). a Herbarium
label. b Section of ascoma. c
Pseudoparaphyses. d, e Asci.
f, g Ascospores. Scale bars: b
= 200 µm, c = 4 µm, d, e = 30
µm, f, g = 15 µm
21.
22.
22.
23.
23.
24.
24.
25.
25.
26.
Conidiophores < 480 μm long………………………22
Conidia 38–45 μm diam.……………………T. brevis
Conidia > 45 μm diam.………………………………23
Conidia 310–370 μm long………T. sympodihylospora
Conidia up to 480 μm long…………………………24
Conidiophores 33–86 μm long…………T. inaequalis
Conidiophores 25–45 μm long…T. sympodilaxispora
Conidia > 260 μm long, 260–305 μm……T. tratensis
Conidia < 260 μm long……………………………26
Conidiogenous cells intercalary or terminal, 7–13 μm
long………………………………T. fangchengensis
26. Conidiogenous cells terminal, 8.5–21 μm long………
…………………………………………T. krabiensis
Wiesneriomycetaceae Suetrong, Rungjind., Somrith. &
E.B.G. Jones, Phytotaxa 176(1): 288 (2014)
Wiesneriomyces Koord., Verh. K. Akad. Wet., tweede
sect. 13(4): 246 (1907)
Saprobic on decaying wood in freshwater or terrestrial
habitats. Description of this genus see Suetrong et al. (2014).
Type species: Wiesneriomyces laurinus (Tassi) P.M. Kirk
Notes: Wiesneriomyces has sporodochial conidiomata
on a dark pseudoparenchymatous stalk, with subulate setae
and cylindrical, hyaline, aseptate, catenate conidia (Suetrong
et al. 2014). Wiesneriomyces formed a well-supported clade
basal to Tubeufiaceae in Tubeufiales based on molecular
data (Suetrong et al. 2014). Three species are accepted
in Wiesneriomyces and W. conjunctosporus Kuthub. &
Nawawi is found from freshwater habitats (Kuthubutheen
and Nawawi 1988), but not confirmed with molecular data.
List of freshwater Wiesneriomyces species
*Wiesneriomyces conjunctosporus Kuthub. & Nawawi,
Trans. Br. mycol. Soc. 90(4): 619 (1988)
Freshwater distribution: Malaysia (Kuthubutheen and
Nawawi 1988)
Dothideomycetes families incertae sedis
Argynnaceae Shearer & J.L. Crane, Trans. Br. mycol. Soc.
75(2): 193 (1980)
13
548
Lepidopterella Shearer & J.L. Crane, Trans. Br. mycol. Soc.
75(2): 194 (1980)
Saprobic on submerged twigs. Sexual morph: Ascomata
slightly immersed to superficial, solitary, globose to subglobose, dark brown to black, lacking ostioles. Peridium pseudoparenchymatic, composed of several layers of brown to
black cells. Pseudoparaphyses numerous or sparse, cellular,
hyaline, septate, branched. Asci 8-spored, bitunicate, subglobose to globose, pedicellate, lacking an ocular chamber,
thick-walled. Ascospores overlapping, irregularly arranged,
papilionaceous, brown to orange-brown, 1-septate, constricted and darker at the septum, smooth-walled (Shearer
and Crane 1980). Asexual morph: Undetermined.
Type species: Lepidopterella palustris Shearer & J.L.
Crane
Notes: Lepidopterella, typified by L. palustris, was placed
in Argynnaceae together with the type genus Argynna Morgan based on the papilionaceous ascospores (Shearer and
Crane 1980). Lepidopterella probably does not belong to
Argynnaceae because of the different ascomata. Lepidopterella possesses globose to subglobose ascomata and brown
to orange-brown ascospores, while Argynna has discal apothecia and pale brown ascospores with a dark band at the
septum (Shearer and Crane 1980; Raja and Shearer 2008;
Hyde et al. 2013). Phylogenetic analyses showed L. palustris did not form strong relationships with any families and
orders in Dothideomycetes (Shearer et al. 2009; Peter et al.
2016). Lepidopterella is retained in Argynnaceae (Hyde
et al. 2013), which is accepted by Wijayawardene et al.
(2018, 2020). Two species are accepted in Lepidopterella
and both of them are from freshwater habitats. They can be
distinguished by ascospore morphology and size (see key
below).
List of freshwater Lepidopterella species
*Lepidopterella palustris Shearer & J.L. Crane, Trans. Br.
mycol. Soc. 75(2): 194 (1980); Fig. 130
Freshwater distribution: USA (Shearer and Crane 1980;
Raja et al. 2009b)
*Lepidopterella tangerina Raja & Shearer, Mycologia
100(3): 481 (2008)
Freshwater distribution: USA (Raja and Shearer 2008;
Raja et al. 2009b)
Key to freshwater Lepidopterella species
1. Ascospores papilionaceous, 15.6–22.8 µm × 25–30
µm…………………………………………L. palustris
1. Ascospores broadly reniform, 16–19 µm × 30–35
µm…………………………………………L. tangerina
13
Fungal Diversity (2020) 105:319–575
Pseudorobillardaceae Crous, Fungal Systematics and Evolution 3: 108 (2019)
Pseudorobillarda M. Morelet, Bull. Soc. Sci. nat. Arch.
Toulon et du Var 175: 5 (1968)
Saprobic on decaying plant materials in terrestrial and
freshwater habitats. Sexual morph: Undetermined. Asexual morph: Pycnidia solitary, scattered or gregarious,
immersed, uniloculate, ellipsoidal, glabrous, dark brown
to black, ostiolate. Periduim composed of several layers of
cells of textura angularis or textura prismatica, darker in
the outer layers, paler in the inner layers. Conidiophores
sparsely septate and branched at the base, or reduced to
conidiogenous cells. Pseudoparaphyses present or absent,
cellular. Conidiogenous cells discrete or integrated, cuboid,
short-cylindrical, subcylindrical, ampulliform, or lageniform, colourless, smooth. Conidia fusiform, subcylindrical, ellipsoid, naviculate, unicellular or euseptate, hyaline,
smooth, bearing attenuated, divergent appendages (Nag Raj
1993).
Type species: Pseudorobillarda phragmitis (Cunnell) M.
Morelet
Notes: Pseudorobillarda is a coelomycetous genus, characterized by hyaline conidia with divergent appendages (Nag
Raj 1993). The placement of Pseudorobillarda is unstable
as it either clustered with Pleosporales (Rungjindamai
et al. 2012), or nested between Tubeufiales and Mytilinidiales (Tangthirasunun et al. 2014). Recently, it was placed
in a newly established family Pseudorobillardaceae which
resides in Minutisphaerales (Crous et al. 2018a, 2019b).
However, Pseudorobillardaceae was shown to not belong
to Minutisphaerales in our phylogenetic tree (Fig. 2), and
therefore placed in Dothideomycetes families incertae sedis.
The freshwater species Pseudorobillarda sojae Uecker &
Kulik was transferred to Stauronematopsis Abbas et al. based
on re-examination of an isotype (IMI 1298796) (Abbas et al.
2002). However, P. sojae (BCC 20495, NTCL090) formed
a well-supported, monophyletic clade with other Pseudorobillarda species (Rungjindamai et al. 2012; Tangthirasunun
et al. 2014; Crous et al. 2018a; this study, Fig. 2). We place
P. sojae in Pseudorobillarda.
List of freshwater Pseudorobillarda species
*Pseudorobillarda sojae Uecker & Kulik, Mycologia 78:
450 (1986)
Synonymy: Stauronematopsis sojae (Uecker & Kulik)
Abbas, B. Sutton & Ghaffar, Pakist. J. Bot. 34: 123 (2002)
Freshwater distribution: Thailand (Plaingam et al. 2005)
Dothideomycetes genera incertae sedis
Alascospora Raja, Violi & Shearer, Mycologia 102(1): 33
(2010)
Fungal Diversity (2020) 105:319–575
Saprobic on submerged, decomposed petioles. Sexual
morph: Ascomata scattered, partly immersed to superficial, uniloculate, globose, subglobose or hemispherical
and flattened at the base, translucent to light brown when
young, darkening with age, membranous, ostiolate. Peridium
composed of hyaline to brown pseudoparenchymatic cells,
dark amorphous material irregularly arranged on the surface. Pseudoparaphyses sparse, celular, broadly filamentous,
sometimes irregular, hyaline, septate. Asci 8-spored, bitunicate, fissitunicate, globose to subglobose, thick-walled, short
pedicellate. Ascospores overlapping irregularly arranged,
ellipsoidal, straight, hyaline when young, brown at maturity, 1-septate, septum becoming thicker and darker in older
ascospores, asymmetrical, upper cell longer and wider than
lower cell, with a gelatinous sheath; sheath often condenses
on ascospores to give a verruculose appearance (Raja et al.
2010b). Asexual morph: Undetermined.
Type species: Alascospora evergladensis Raja, Violi &
Shearer
Notes: Alascospora evergladensis was isolated from
submerged, decomposed petioles of Nymphaea odorata in
Florida, USA (Raja et al. 2010b). A gelatinous wing-like
sheath surrounds the ascospores in A. evergladensis (Raja
et al. 2010b). Alascospora shares similar morphological
Fig. 131 Alascospora evergladensis (Material examined: USA,
Florida, Everglades Water Management District, Water Conservation
Area 2A, on submerged Nymphaea odorata petiole, 17 November
2008, H.A. Raja & H. Violi, SFWMD 1-1, holotype, ILL40789). a
Ascoma mounted in water. b Squash mount of ascoma. c Pseudoparaphyses. d Young, globose, thick-walled ascus. e Fissitunicate ascus. f
549
characters with Minutisphaera, such as subglobose, thickwalled asci and 1-septate, brown ascospores (Raja et al.
2010b, 2015; Ferrer et al. 2011). The sequence data of A.
evergladensis are unavailable and the relationship between
Alascospora and Minutisphaera is pending molecular data.
The holotype of the type species A. evergladensis was examined and described by Pem et al. (2019b). They placed Alascospora in Dothideomycetes genera incertae sedis based on
morphology and this was accepted by Wijayawardene et al.
(2020). We follow this placement and illustrate the holotype
specimen of A. evergladensis in this study (Fig. 131).
List of freshwater Alascospora species
Alascospora evergladensis Raja, Violi & Shearer, Mycologia 102(1): 36 (2010); Fig. 131
Freshwater distribution: USA (Raja et al. 2010b)
Ascominuta Ranghoo & K.D. Hyde, Mycoscience 41: 1
(2000)
Saprobic on submerged wood. Sexual morph: Ascomata solitary or aggregated, partly immersed to superficial, uniloculate, globose, dark brown to black, ostiolate,
periphysate. Peridium composed of several layers of hyaline to dark brown, compressed cells of textura angularis.
Young, multiguttulate ascospore with wing-shaped, gelatinous sheath
(arrows). g Verruculose, brown ascospores at maturity. Scale bars: a
= 100 µm, b = 50 µm, c–f = 10 µm, g = 15 µm. Some images were
taken from the Freshwater Ascomycetes Database: http://fungi.life.
illinois.edu (maintained by Carol A. Shearer and Huzefa A. Raja)
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Fungal Diversity (2020) 105:319–575
Fig. 132 Ascominuta spp. a–c
A. lignicola (redrawn from
Ranghoo and Hyde (2000),
HKUM 5246, holotype). a
Peridium of ascoma. b Ascus. c
Ascospore. d Ascospore of A.
ovalispora (redrawn from Hu
et al. (2010a), MFLU 08-1176,
holotype). Scale bars: a–d =
10 μm
Pseudoparaphyses sparse, cellular, comprising short chains
of globose cells, often breaking up into individual cells. Asci
4-spored, bitunicate, globose, short or non-pedicellate, without an apical apparatus. Ascospores overlapping multiseriate, broadly fusiform or ovoid, occasionally curved, hyaline,
1-septate, constricted at the septum, smooth-walled, with a
distinct sheath (Ranghoo and Hyde 2000; Hu et al. 2010a).
Asexual morph: Undetermined.
Type species: Ascominuta lignicola Ranghoo & K.D.
Hyde
Notes: Ascominuta comprises two freshwater species,
characterized by globose asci containing four spores and
broadly fusiform or ovoid, hyaline, 1-septate ascospores
with a distinct sheath (Ranghoo and Hyde 2000; Hu et al.
2010a). Ascominuta was probably an earlier name for Natipusilla based on morphology and habitat (see notes of Natipusilla) (Hyde et al. 2013), but needs to be established by
molecular data. Ascominuta was initially placed in Ascomycetes genera incertae sedis (Ranghoo and Hyde 2000), then
placed in Dothideomycetes genera incertae sedis (Lumbsch
and Huhndorf 2007), which was accepted by Wijayawardene
et al. (2018, 2020). Ascominuta lignicola cannot be aligned
with other Dothideomycetous species in our sequence
dataset, so we exclude this species from our phylogenetic
analyses.
Ascominuta lignicola Ranghoo & K.D. Hyde has broadly
fusiform ascospores surrounded by an irregular, mucilaginous sheath, while A. ovalispora D.M. Hu & K.D. Hyde has
ovoid ascospores surrounded by sac-like, persistent sheath.
Ascominuta lignicola has smaller ascospores than A. ovalispora (see key below) (Ranghoo and Hyde 2000; Hu et al.
2010a).
List of freshwater Ascominuta species
*Ascominuta lignicola Ranghoo & K.D. Hyde, Mycoscience 41: 1 (2000); Fig. 132a–c
13
Freshwater distribution: China (Ranghoo and Hyde
2000), Thailand (Hu et al. 2010b)
Ascominuta ovalispora D.M. Hu & K.D. Hyde, Sydowia
62: 192 (2010); Fig. 132d
Freshwater distribution: Thailand (Hu et al. 2010a)
Key to freshwater Ascominuta species
1. Ascospores 22–25 × 8–10 μm………A. lignicola
1. Ascospores 28–30 × 9–10 μm………A. ovalispora
Lucidascocarpa A. Ferrer, Raja & Shearer, Mycologia
100(4): 642 (2008)
Saprobic on submerged wood. Sexual morph: Ascomata
scattered to clustered, immersed or superficial, uniloculate,
globose to subglobose, white, membranous, with long,
cylindrical, white necks, periphysate. Peridium composed
of several layers of elongate, thin-walled, hyaline cells of
textura angularis. Pseudoparaphyses absent. Asci 8-spored,
bitunicate, fissitunicate, clavate, pedicellate, with gelatinous
material around the apex of the endoascus. Ascospores overlapping biseriate, ellipsoidal to fusiform, hyaline, septate,
verruculose, thin-walled, surrounded by a large gelatinous
sheath (Ferrer et al. 2008). Asexual morph: Undetermined.
Type species: Lucidascocarpa pulchella A. Ferrer, Raja
& Shearer
Notes: The monotypic genus Lucidascocarpa, typified by
L. pulchella, is unusual in Dothideomycetes. It has white ascomata with long neck, gelatinous material around the apex of
the endoascus, and phragmoseptate, rough-walled ascospores
surrounded by a large gelatinous sheath (Ferrer et al. 2008).
The light coloured ascomata and well-developed sheath of
Lucidascocarpa are similar to Aliquandostipite (Jahnulales)
(Inderbitzin et al. 2001), but no pseudoparaphyses and wide,
brown, septate hyphae in Lucidascocarpa. Lucidascocarpa was
suggested in Dothideales based on fissitunicate asci without
Fungal Diversity (2020) 105:319–575
551
Fig. 133 Lucidascocarpa pulchella (Material examined: ECUADOR, Yasuni National Park, Laguna-Tiputini River, A. Ferrer,
A566-1). a Ascomata on wood from a freshwater habitat; note: the
ascomata can be completely or partially submerged with hyaline
necks protruding from the surface of the wood. b Squash mount of
ascoma showing neck with hyaline periphysis. c Partially empty
ascus with gelatinous material surrounding the apex of the endoas-
cus. d Ascospore with arrows indicating expanded gelatinous sheaths
in water. e Ascospore with gelatinous sheath in aqueous nigrosin. f
Detached gelatinous sheath. Scale bars: a = 200 µm, b = 100 µm,
c–f = 20 µm. Some images were taken from the Freshwater Ascomycetes Database: http://fungi.life.illinois.edu (maintained by Carol A.
Shearer and Huzefa A. Raja)
pseudoparaphyses, but it cannot be assigned to any existing
families (Ferrer et al. 2008). It is now placed in Dothideomycetes genera incertae sedis (Wijayawardene et al. 2018, 2020).
Distributional studies of freshwater ascomycetes indicated that
L. pulchella might be restricted to the neotropics (Ferrer et al.
2008; Shearer et al. 2015). Sequence data of L. pulchella are
needed.
Saprobic on various hosts in freshwater, terrestrial and
marine habitats. Sexual morph: Undetermined. Asexual
morph: Colonies on natural substrate effuse or sporodochial,
green, greenish blue, lavender, blackish brown or black. Mycelium superficial or partly immersed, consisting of branched,
septate, thin-walled, smooth or verrucose, pale to medium
brown hyphae. Conidiophores micronematous or semimacronematous, mononematous, scattered or closely packed
together in the sporodochia, straight or flexuous, unbranched
or irregularly branched, hyaline to brown, septate or aseptate,
smooth, thin-walled, sometimes with swollen cells. Conidiogenous cells monoblastic, integrated, terminal, determinate,
single, cylindrical, doliiform or subspherical, hyaline or subhyaline. Conidia acrogenous, solitary, dry, simple, oblong, pyriform, clavate, ellipsoidal, subspherical or irregular, rounded at
the ends, straight, sometimes spirally twisted, brown to black,
List of freshwater Lucidascocarpa species
Lucidascocarpa pulchella A. Ferrer, Raja & Shearer, Mycologia 100: 644, 2008; Fig. 133
Freshwater distribution: Costa Rica and Ecuador (Ferrer
et al. 2008), Peru (Shearer et al. 2015)
Monodictys S. Hughes, Can. J. Bot. 36: 785 (1958)
13
552
muriform, smooth or verrucose, basal cell sometimes inflated,
paler and thinner-walled than the other cells. Conidial secession mostly schizolytic (Cai et al. 2006a).
Type species: Monodictys putredinis (Wallr.) S. Hughes
Notes: Monodictys is characterized by muriform, irregular-shaped conidia usually formed on micronematous conidiophores (Cai et al. 2006a). Monodictys species were isolated
from wood, leaves, stems, litter, lichens, soil and dung (Seifert
et al. 2011), and human skin (Singh and Barde 1985) in freshwater and marine habitats (Rao and De Hoog 1986; Goh and
Hyde 1999).
The sexual morphs of several monodictys-like species
were linked to Ascotaiwania Sivan. & H.S. Chang, Hyaloscypha Boud., Nereiospora E.B.G. Jones et al., Ohleria
Fuckel and Tubeufia based on cultural methods (Samuels
1980; Mouzouras and Jones 1985; Chang 2001; Hosoya
and Huhtinen 2002). Monodictys is similar to Berkleasmium
Zobel in conidial morphology, but differs by the micronematous conidiophores (Cai et al. 2006a). Monodictys is polyphyletic (Tanaka et al. 2015). The sequence data of the type
species M. putredinis are unavailable, thus Monodictys is
tentatively placed in Dothideomycetes genera incertae sedis
(Wijayawardene et al. 2018, 2020).
Ten species occur in freshwater habitats (see list below)
and none of them has sequence data. They can be distinguished by sporodochial conidiomata, conidial shape, size,
septa and ornamentation.
List of freshwater Monodictys species
Monodictys castaneae (Wallr.) S. Hughes, Can. J. Bot. 36:
785 (1958)
Basionym: Hyphelia castaneae Wallr. [as ‘castanea’], Fl.
crypt. Germ. (Norimbergae) 2: 244 (1833)
Freshwater distribution: USA (Shearer 1972)
Monodictys cruciseptata J.L. Crane & Schokn., in Schoknecht & Crane, Can. J. Bot. 61(8): 2244 (1983)
Freshwater distribution: USA (Schoknecht and Crane
1983)
Monodictys fluctuata (Tandon & Bilgrami) M.B. Ellis,
Mycol. Pap. 125: 5 (1971)
Basionym: Acrospeira fluctuata Tandon & Bilgrami, Sci.
Cult. 27: 408 (1961)
Freshwater distribution: Thailand (Hu et al. 2010b)
Monodictys melanocephaloides Goh & K.D. Hyde, Fungal
Diversity 3: 77 (1999)
Freshwater distribution: China (Goh and Hyde 1999),
Thailand (Hu et al. 2010b)
Monodictys pandani Matsush., Matsush. Mycol. Mem. 4:
12 (1985)
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Fungal Diversity (2020) 105:319–575
Freshwater distribution: Australia (Hyde and Goh
1998a), China (Tsui et al. 2001c)
Monodictys paradoxa (Corda) S. Hughes, Can. J. Bot. 36:
786 (1958)
Freshwater distribution: Thailand (Hu et al. 2010b)
Monodictys putredinis (Wallr.) S. Hughes, Can. J. Bot. 36:
785 (1958)
Basionym: Melanconium putredinis Wallr., Fl. crypt.
Germ. (Norimbergae) 2: 181 (1833)
Freshwater distribution: South Africa (Hyde et al. 1998),
China (Tsui et al. 2000, 2001c)
Monodictys sessilis Hol.-Jech., in Holubová-Jechová & Mercado Sierra, Česká Mykol. 40(3): 151 (1986)
Freshwater distribution: China (Tsui et al. 2000, 2001c)
Monodictys striata (Petch) V. Rao & de Hoog, Stud. Mycol.
28: 30 (1986)
Basionym: Sporidesmium striatum Petch, Ann. R. bot.
Gdns Peradeniya 6(3): 249 (1917)
Freshwater distribution: China (Tsui et al. 2000)
Monodictys trichocladiopsis Goh & K.D. Hyde, Fungal
Diversity 3: 79 (1999)
Freshwater distribution: China (Goh and Hyde 1999)
Key to freshwater Monodictys species
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
8.
8.
9.
9.
Sporodochial conidiomata present……………………2
Sporodochial conidiomata absent………………………4
Conidia dictyoseptate…………M. melanocephaloides
Conidia irregularly septate……………………………3
Conidia 16–22 × 27–35 μm…………………M. striata
Conidia 68–115 × 36–48 μm…………………M. sessilis
Conidia < 10 cells………………………………………5
Conidia > 10 cells………………………………………6
Conidia globose to subglobose………M. cruciseptata
Conidia pyriform to ellipsoidal……M. trichocladiopsis
Basal cell conspicuous……………………M. paradoxa
Basal cell inconspicuous if hyaline……………………7
Conidia subglobose to broadly obovoidal, brown to
black………………………………………M. castaneae
Conidia ellipsoidal to elongate, medium brown if obovoidal…………………………………………………8
Mature conidia rough-walled………………M. fluctuata
Mature conidia smooth-walled…………………………9
Conidia sessile……………………………M. pandani
Conidia with some hyaline or pale brown supporting
cells………………………………………M. putredinis
Fungal Diversity (2020) 105:319–575
553
Fig. 134 Ocala scalariformis
(Material examined: USA,
Florida, Ocala National Forest,
Mary Lake, on submerged
decorticated woody debris, 16
July 2006, H.A. Raja & J.L.
Crane, F121-1, ILL40594,
holotype). a Sqaush mount of
ascoma. b Pseudoparaphyses.
c, d Bitunicate asci. e Asci and
ascospores. f Ascospore showing appendages. g Ascospores.
Scale bars: a, c = 40 µm, b,
d, e, g = 20 µm, f = 15 µm.
Images were taken from the
Freshwater Ascomycetes
Database: http://fungi.life.illin
ois.edu (maintained by Carol A.
Shearer and Huzefa A. Raja)
Ocala Raja & Shearer, Fungal Diversity 34: 80 (2009)
Saprobic on submerged wood. Sexual morph: Ascomata
scattered, immersed or superficial, uniloculate, globose to
subglobose, brown to dark brown, or semi-transparent, membranous, with ostiolate papilla. Peridium thin-walled, composed of textura angularis to epidermoidea in surface view.
Pseudoparaphyses sparse, cellular, broad, hyaline, septate,
branched. Asci 8-spored, bitunicate, fissitunicate, obclavate
to fusoid, flattened at the apex, with an elongate, tapered
pedicel. Ascospores overlapping uni- to bi-seriate, narrowly
ellipsoidal to fusiform, straight or slightly curved, hyaline,
phragmoseptate, thick-walled, with or without amorphous
apical gelatinous appendages (Raja et al. 2009a). Asexual
morph: Undetermined.
Type species: Ocala scalariformis Raja & Shearer
Notes: The monotypic genus Ocala was introduced for a
freshwater species O. scalariformis collected from Florida
(Raja et al. 2009a). Ocala is characterized by obclavate to
fusoid asci, narrowly ellipsoidal to fusiform, hyaline, phragmoseptate, thick-walled ascospores with or without amorphous apical gelatinous appendages (Raja et al. 2009a). The
phylogenetic placement of Ocala is questionable because
the ex-type strain F121 of O. scalariformis clustered with
Natipusilla bellaspora in Natipusillales with high bootstrap
support (Raja et al. 2015). However, Ocala has brown to
dark brown ascomata, obclavate to fusoid asci and phragmoseptate ascospores contrasting with the hyaline to light
brown ascomata, subglobose asci and mostly 1-septate
ascospores of Natipusilla (Raja et al. 2009a, 2012; Ferrer
et al. 2011). We notice that only SSU sequence data of O.
scalariformis (NG_061098, F121, from holotype) is deposited in GenBank and included in the phylogenetic analysis of Raja et al. (2015). The phylogenetic placement of O.
scalariformis is pending awaiting more sequences. Ocala is
presently placed in Dothideomycetes genera incertae sedis
(Wijayawardene et al. 2018, 2020).
List of freshwater Ocala species
*Ocala scalariformis Raja & Shearer, Fungal Diversity 34:
80 (2009); Fig. 134
Freshwater distribution: USA (Raja et al. 2009a)
Quintaria Kohlm. & Volkm.-Kohlm., Bot. Mar. 34: 34
(1991)
Saprobic on submerged wood in freshwater or dead tip
of prop root of Rhizophora mangle in terrestrial habitats.
13
554
Sexual morph: Ascomata scattered or loosely gregarious,
immersed, with or without clypeus, uniloculate, subglobose
to globose, conical with flattened bases or obpyriform, glabrous, dark brown to black, coriaceous or carbonaceous,
with ostiolate papilla. Ostiole central or ecentric, circular,
short, brown to black, periphysate. Clypeus when present
composed of host cells, brown fungal hyphae and melanin
particles. Peridium thin to medium, comprising several layers of brown pseudoparenchymatous angular cells mixing
with host cells. Pseudoparaphyses numerous, trabeculate,
hyaline, branched, anastomosing above the asci, embedded
in a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, cylindrical to clavate, with conspicuous or inconspicuous IKI negative apical structures, short pedicellate,
persistent. Ascospores overlapping bi- to tri-seriate, ellipsoidal to cylindrical or fusiform, straight to slightly curved,
hyaline, multiseptate, constricted at the septa, wall minutely
verrucose or smooth, guttulate, with or without sheath
(Hyde and Goh 1999b; Cai et al. 2006a). Asexual morph:
Undetermined.
Type species: Quintaria lignatilis (Kohlm.) Kohlm. &
Volkm.-Kohlm.
Notes: Quintaria was initially described for a marine
species, Q. lignatilis, which is characterized by immersed
ascomata with fusoid, hyaline, multiseptate ascospores
(Kohlmeyer and Volkmann-Kohlmeyer 1991). Quintaria
is morphologically similar to Trematosphaeria, but can
be distinguished by completely immersed ascomata with
black incrustations lining the sides of the ostiolar canal, and
hyaline ascospores (Kohlmeyer and Volkmann-Kohlmeyer
1991; Zhang et al. 2012b). Quintaria resembles the members
of Lophiostomataceae in having cylindrical to clavate asci
and fusiform, hyaline ascospores, but they are phylogenetically distant (Suetrong et al. 2009; Zhang et al. 2012b).
The placement of Quintaria is unstable. Quintaria was
referred to Lophiostomataceae (Cai et al. 2006a; Jones et al.
2009), Pleosporales genera incertae sedis (Zhang et al.
2012b) and Dothideomycetes genera incertae sedis (Wijayawardene et al. 2017; Hongsanan et al. 2020a) in different
papers. Two strains of Q. lignatilis, BCC 17444 and CBS
117700, clustered with Lophiostomataceae and Astrosphaeriellaceae, respectively (Zhou et al. 2013). However, two strains
clustered together and close to Testudinaceae in Suetrong
et al. (2009) and Zhang et al. (2012b). We found that there
are over 100 nucleotide differences in LSU sequence data
between BCC 17444 and CBS 117700, which indicates that
they are different generic species and even probably belong to
different families. The morphology of BCC 17444 and CBS
117700 are unknown, so we can not compare their morphology with the holotype specimen No. 4365a of Q. lignatilis
examined by Zhang et al. (2012b). Based on the unstable
relationships of Quintaria, the difference of sequences of
two strains, we place Quintaria in Dothideomycetes genera
13
Fungal Diversity (2020) 105:319–575
incertae sedis, following Wijayawardene et al. (2017), until
the new collections of Q. lignatilis are obtained.
Two freshwater species Quintaria aquatica K.D. Hyde &
Goh and Q. microspora Yin. Zhang et al. lack sequence data.
Quintaria aquatica has larger ascospores than Q. microspora (42–52 × 11–14 μm vs. 26–31 × 5–5.5 μm) and posses
a sheath (Hyde and Goh 1999b; Zhang et al. 2008a). Zhang
et al. (2012b) suspected that all freshwater species are likely
to be excluded from this genus and only the generic type Q.
lignatilis retained, but this needs confirmation. The freshwater species Q. submersa K.D. Hyde & Goh now is placed
in Neolindgomyces Jayasiri et al. as N. submersus (K.D.
Hyde & Goh) Jayasiri & K.D. Hyde based on molecular
data (Jayasiri et al. 2019).
Fig. 135 Wettsteinina spp. a–e W. niesslii (redrawn from Scheuer
(1995), lectotype). a Longitudinal section of ascoma. b Ascus and
interascal filaments. c–e Young, mature and senescent ascospores. f–
h W. moniliformis (redrawn from (Van Ryckegem and Aptroot 2001),
holotype, GENT). f Ascus. g Ascospore. h Pseudoparaphyses filament. Scale bars: a = 50 μm, b–e = 20 μm, f, g = 10 μm, h = 20 μm
Fungal Diversity (2020) 105:319–575
555
List of freshwater Quintaria species
Quintaria aquatica K.D. Hyde & Goh, Nova Hedwigia 68:
258 (1999)
Freshwater distribution: Australia (Hyde and Goh 1999b)
Freshwater distribution: England (Ingold 1955; Apinis
et al. 1972; Taligoola et al. 1972), USA (Shearer 1972)
Quintaria microspora Yin. Zhang, K.D. Hyde & J. Fourn.,
Cryptogamie Mycologie 29: 180 (2008)
Freshwater distribution: France (Zhang et al. 2008a)
1. Ascospores with pointed ends, 55–70 × 9.5–12 μm……
………………………………………………W. niesslii
1. Ascospores with rounded ends, 19–23 × 7–9 μm………
…………………………………………W. moniliformis
Key to freshwater Wettsteinina species
Key to freshwater Quintaria species
1. Ascospores without sheath………………Q. microspora
1. Ascospores with sheath……………………Q. aquatica
Wettsteinina Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 116: 126 (1907)
Saprobic or hemibiotrophic on submerged wood in freshwater or plant leaves and stems in terrestrial habitats. Sexual
morph: Ascomata small-sized, scattered, immersed with a
protruding broad papilla. Peridium composed of several layers of large, polygonal, thin-walled cells. Pseudoparaphyses
deliquescing at maturity. Asci bitunicate, fissitunicate, subglobose to obpyriform, sessile, with a small ocular chamber,
thick-walled. Ascospores 2–4-seriate, broadly clavate, hyaline and become pale brown when mature, septate, smooth,
surrounded by a hyaline gelatinous sheath (Zhang et al.
2012b). Asexual morph: Undetermined.
Type species: Wettsteinina gigantospora (Rehm) Höhn.
Notes: Wettsteinina is characterized by thick-walled,
obpyriform asci, asymmetrical ascospores, and lack pseudoparaphyses at maturity (Zhang et al. 2012b). The placement
of Wettsteinina is confused because of lack of sequences of
the type species, W. gigantospora. Wettsteinina was placed
in Pleosporaceae based on its “pleospora-type” centrum
(Shoemaker and Babcock 1987), later excluded from Pleosporaceae based on molecular data of three species, but not
assigned to other families (Kodsueb et al. 2006). Wettsteinina is placed in Dothideomycetes genera incertae sedis by
Wijayawardene et al. (2018, 2020). The molecular data of
freshwater species W. moniliformis Van Ryck. & Aptroot
and W. niesslii E. Müll. are unavailable. They can be distinguished from each other by the size of ascospores (see
key below).
List of freshwater Wettsteinina species
Wettsteinina moniliformis Van Ryck. & Aptroot, Nova Hedwigia 73(1–2): 164 (2001); Fig. 135f–h
Freshwater distribution: Denmark (Van Ryckegem and
Aptroot 2001)
Wettsteinina niesslii E. Müll., Sydowia 4(1–6): 204 (1950);
Fig. 135a–e
Genus formerly placed in Dothideomycetes
Bahusutrabeeja Subram. & Bhat, Can. J. Bot. 55: 2204
(1977)
Saprobic on submerged wood in freshwater or rotten
leaf, wood in terrestrial habitats. Sexual morph: Undetermined. Asexual morph: Colonies hairy on wood. Mycelium mostly immersed, composed of septate, brown hyphae.
Conidiophores macronematous, erect, straight, cylindrical,
smooth, brown, paler towards the apex, lacking hyphopodia
at the base, septate, simple or branched, thick-walled, often
proliferating percurrently through the collarette. Conidiogenous cells monophialidic, with a distinct collarette, terminal, integrated. Phialides producing many conidia singly and
successively from the tip. Phialoconidia acrogenous, initial
Fig. 136 a–g Bahusutrabeeja dwaya (redrawn from Subramanian
and Bhat (1977), MUBL 2351, holotype). a, c, d Conidiophores
bearing conidia. b Conidiophore. e, f Conidia. Scale bars: a = 50 μm,
b = 30 μm, c, e, f = 10 μm, d = 15 μm
13
556
conidium pear-shaped, others globose to subglobose, hyaline
with granular cytoplasm, aggregated in slimy mass at the
tip of the phialide, aseptate, with some slender appendages,
with or without a large, lipid globule (Subramanian and Bhat
1977; Tsui et al. 2001a).
Type species: Bahusutrabeeja dwaya Subram. & Bhat
Notes: Bahusutrabeeja was established to accommodate
a taxon producing pear-shaped to subglobose, unicellular,
hyaline phialoconidia with some slender surface appendages
(Subramanian and Bhat 1977). The type species B. dwaya
was initially isolated from dead twigs of Coffea arabica in
India (Subramanian and Bhat 1977), subsequently collected
from freshwater habitats in Hong Kong, China (Tsui et al.
2000, 2001a, c) and Malaysia (Kuthubutheen et al. 1992).
The collection from Malaysia (IMI 344362) has longer
appendages than the holotype (MUBL 2351) (up to 12 μm
vs. 4.5–5.5 μm). The collection of Tsui et al. (2001a) has a
large lipid globule, which was not found in other collections.
Bahusutrabeeja dwaya clustered with Neodeightonia subglobosa C. Booth with strong bootstrap support and Bahusutrabeeja was placed in Botryosphaeriaceae (Shenoy et al.
2010). Wijayawardene et al. (2017) listed Bahusutrabeeja in
Botryosphaeriales genera incertae sedis. A strain GUFCC
4904 of B. dwaya collected from India (Shenoy et al. 2010),
is included in our phylogenetic tree. The result shows that
B. dwaya nests between Botryosphaeriaceae and Phyllostictaceae in Botryosphaeriales with strong bootstrap support
(Fig. 2). However, Pem et al. (2019b) thought that GUFCC
4904 is a wrong isolation and transferred Bahusutrabeeja
to Chaetosphaeriales in Sordariomycetes based on a ITS
sequence of CBS 261.77 derived from the ex-type culture
of B. dwaya. The hyphomycetous species B. dwaya also
can not be placed in Botryosphaeriales based on morphology (Hyde et al. 2013; Crous et al. 2015b; Linaldeddu et al.
2016; Tao et al. 2016). Only ITS sequence of B. dwaya (CBS
261.77) is deposited in GenBank. Additional sequences are
expected, so the placement of B. dwaya would be clear with
phylogenetic analyses. We follow Pem et al. (2019b) and
Wijayawardene et al. (2020) and place Bahusutrabeeja in
Chaetosphaeriales (Sordariomycetes).
List of freshwater Bahusutrabeeja species
*Bahusutrabeeja dwaya Subram. & Bhat, Can. J. Bot. 55:
2204 (1977); Fig. 136
Freshwater distribution: China (Tsui et al. 2000, 2001a,
2001c), Malaysia (Kuthubutheen et al. 1992)
Discussion
Studies of lignicolous freshwater fungi have greatly
advanced in the last decade, with molecular data allowing
a better resolution of species, the linking of asexual and
13
Fungal Diversity (2020) 105:319–575
sexual morphs, as well as placements at the family, ordinal and class levels (Zhang et al. 2012a, 2016, 2017a; Su
et al. 2016b; Dong et al. 2018; Luo et al. 2019). As a result,
many new species have been published (Lu et al. 2018b;
Luo et al. 2019; Dong et al. 2020; Hyde et al. 2020b). Many
other collections and new records have also been made, but
not published as it is hard to find a publishing outlet (Hyde
et al. 2020a). With the new AJOM notes (Hyde et al. 2020a),
this should become easier, while surveys of fungi in streams
with their percentage occurrence and ecology should also
be encouraged as in previous earlier papers (Goh and Hyde
1999; Ho et al. 2001; Cai et al. 2003a). Hyde et al. (2020a)
noted the need to publish new records and contributed 100
new records and collections of taxa, and found morphology and sequence variability for each taxon. It is, therefore,
important to define and understand a species using different
collections from different regions or habitats (Hyde et al.
2020d).
Adaptions of freshwater Dothideomycetes to dispersal
Adaptations of microfungi in freshwater were thought to
include a) disappearance of an ascus apical ring and release
of ascospores via deliquescent asci and b) the presence
of ascospore appendages (Shearer 1993a). Hyde and Goh
(2003) reviewed the adaptations of filamentous fungi for
dispersal in freshwater and should be consulted. The aquatic
hyphomycetes generally have spore shapes or trap air bubbles which are conductive for dispersal in water and efficient
at becoming trapped in debris or river foam. Ascomycetes
on the other hand, have passive of active dispersal of spores
from the asci and appendages for effective entrapment in
debris or foam. Thus, although release of ascospores is not
often seen, an ascus apical ring (often very large) can be
seen in many freshwater species (Luo et al. 2019; Hyde
et al. 2020a, b, d). We consider the key characteristics in
the adaptions of freshwater Dothideomycetes for dispersal
in freshwater are 1) the active ejection of ascospores (fissitunicate dehiscence), 2) the ornamentation and appendages
of ascospores through which they attach to debris, and 3)
some special adaptations of ascomata which hang on mycelial strands as in Jahnulales. In Table 2, the key characters
of freshwater sexual genera are summarized with species
numbers. Among 30 freshwater genera examined, 22 have
ascospores with sheaths, appendages or ornamentations,
and three have fasciculate or papilionaceous ascospores
(Table 2). These characters are likely to help taxa in dispersal and adapt to life in flowing freshwater. Genera which
have with most freshwater species, such as Ascagilis, Jahnula, Lindgomyces and Neohelicascus support our conclusion. Most ascospores in Ascagilis have pads at each end
(Figs. 6f, g, 10, 11, see key to freshwater Ascagilis species)
or other appendage types. The ascomata of Jahnula are special in having ascomata with a stalk-like strand (Figs. 6c, 7,
Freshwater genera (species number) Key characters
Ascomata
Alascospora (1)
Angustospora (1)
Aquatospora (1)
Aquihelicascus (3)
Ascagilis (7)
Ascominuta (2)
Ascorhombispora (1)
Crassiclypeus (1)
Dictyospora (1)
Fusiformiseptata (1)
Isthmosporella (1)
Jahnula (11)
Lepidopterella (2)
Lindgomyces (14)
Lolia (2)
Lucidascocarpa (1)
Mamillisphaeria (1)
Megalohypha (1)
Minutisphaera (5)
Natipusilla (4)
Neohelicascus (8)
Neotrematosphaeria (1)
With a gelatinous sheath; sheath often condenses on ascospores to give a
verruculose appearance
Immersed to erumpent
With a thin, gelatinous, striate layer
Superficial, with carbonized peridium which composed of black amorphous –
layer
Pseudostromata comprises brown to black fungal material growing in cor–
tex of host cells, visible on the host surface as blackened ostiolar dots
Semi-immersed or superficial, without appendages
With a pad at each end
Partly immersed to superficial
With a distinct sheath
Superficial
Verruculose, with bipolar, small, hemispherical, subhyaline to light brown
cells, surrounded by gelatinous mucilaginous sheath
Immersed, with a crest-like ostiolar neck, with a well-developed clypeus
With an internal chamber and a narrow sheath which drawn out at both ends
Superficial, seated on a subiculum, covered with brown to black setae
Fasciculate
Immersed with a neck erumpent through host surface, arrow-shaped at the –
basal region
Immersed to superficial
Cylindrical to fusoid, isthmoid at center, tapering at both ends, surrounded by
a gelatinous sheath
–
Semi-immersed to erumpent, become superficial with base remaining
immersed, with a stalk-like strand or stoloniferous hyphae attached to the
substratum, or covered by sparse hair-like projections
Immersed to superficial
Irregularly arranged in asci, papilionaceous
Semi-immersed to nearly superficial
Usually with a sheath or bipolar mucilaginous appendages
Immersed to erumpent, usually stain the substrate purple
With a prominent gelatinous sheath
Immersed or superficial, with long, cylindrical, white necks
Verruculose, with a large gelatinous sheath
Superficial, under a pseudostroma that forms a thin layer on the host surTwo types of ascospores. Type 1: symmetric, with pad-like, mucilaginous
face, with palisade-like cells at the sides of peridium
appendage at each end and with some mucilage associated around the
spore; Type 2: asymmetric, with a thin mucilaginous sheath
Superficial, sometimes stalked
Irregularly arranged in asci, rough-walled, longitudinally striate
Erumpent to superficial, covered with black hairs
Asymmetrical, smooth- or rough-walled when mature, with sheath and filamentous appendages
Erumpent to superficial, without appendages
Mostly with a complex gelatinous sheath
Pseudostromata comprising brown to black fungal material growing in
Sometimes with a sheath
cortex of host, visible on the host surface as blackened ostiolar dots
Superficial, covered with few setae, with one-layered peridium
–
Deeply immersed, stain substrate purple, visible as black, slit-like spot on
Minutely verruculose, usually with a thin gelatinous sheath
the substrate surface
Semi-immersed to erumpent, with a crest-like apex and a pore-like opening With bipolar appendages
557
13
Neojahnula (1)
Neomassariosphaeria (1)
Partly immersed to superficial
Ascospores
Fungal Diversity (2020) 105:319–575
Table 2 Key characters of freshwater sexual genera
Fungal Diversity (2020) 105:319–575
Usually with a large, expanding gelatinous sheath
With a distinctly mucilaginous sheath
Are freshwater species taxonomically unique?
Marine, freshwater and terrestrial fungi are treated as
different groups because they survive in different habitats
(Jones et al. 2009, 2019; Suetrong et al. 2009; Doilom et al.
2017; Luo et al. 2019; Dong et al. 2020). In phylogenetic
analyses, freshwater fungi mostly group in Dothideomycetes
and Sordariomycetes (Luo et al. 2019; Hyde et al. 2020a, b),
and some in Eurotiomycetes (Dong et al. 2018, 2020; Yu
et al. 2018; Wang et al. 2019). Lu et al. (2018b) showed that
many freshwater species clustered in Tubeufiaceae in Dothideomycetes, especially the helicosporous hyphomycetes.
Luo et al. (2019) showed that freshwater species mostly
clustered in Diaporthomycetidae in Sordariomycetes. In
our study, freshwater species mostly belong in Pleosporales
and Tubeufiales, some in Jahnulales, Kirschsteiniotheliales,
Minutisphaerales and Natipusillales (Fig. 2). These examples indicate that freshwater fungi are a distinctive group,
and habitat will affect classification. A good example from
our study is Helicascus. Freshwater Helicascus species differ
from marine Helicascus species in their asci and ascospores,
and our phylogenetic analyses clarifies that they belong in
different genera (see entry on Morosphaeriaceae). We suggest that as further genera and species are studied in more
detail, this conclusion will become more clear.
Note: “–” ascospores lack an obvious feature
Tingoldiago (3)
Wicklowia (2)
Purpureofaciens (1)
With a thin mucilaginous sheath
Fasciculate, elongated
Often with amorphous apical gelatinous appendages
With a mucilaginous sheath, which is wavy in outline
Superficial
Immersed or superficial
Erumpent, become superficial with base remaining immersed, covered with
sparse hyaline hairs
Semi-immersed to superficial, stain the substrate purple and produce reddish pigmentation at the apex
Immersed to erumpent
Immersed to erumpent, visible as a black, oval to circular, shallow, craterlike depression on the substrate
13
Neotubeufia (1)
Ocala (1)
Pseudojahnula (1)
Ascospores
9). This possibly enables the ascomata to trail in running
water which may aid in the dispersal of ascospores. The
ascospores of Lindgomyces usually have a sheath or bipolar
mucilaginous appendages (Fig. 57, see key to freshwater
Lindgomyces species). Neohelicascus has pseudostromata
that comprises brown to black fungal material growing in
the host cortex (Figs. 81, 82, 83).
Ascomata
Freshwater genera (species number) Key characters
Table 2 (continued)
558
Cellular and trabeculate pseudoparaphyses in the classification of freshwater Dothideomycetes
Pseudoparaphyses have always been regarded as an important criterion in the subdivision of Dothideomycetes (Liew
et al. 2000). In previous phylogenetic studies, species having
trabeculate pseudoparaphyses did not form a monophyletic
clade at the ordinal level (Liew et al. 2000). Also, our study
reveals that Pleosporales members have two types of pseudoparaphyses, e.g. Astrosphaeriellaceae, including our new
genus Aquatospora, have obviously trabeculate pseudoparaphyses (Figs. 28–35), while most members of Lentitheciaceae
have cellular pseudoparaphyses (Figs. 49, 52). Natipusillales
is distinctive in having sparse or lacking pseudoparaphyses.
Jahnulales mostly has cellular pseudoparaphyses, but Jahnula apiospora has obviously trabeculate pseudoparaphyses
and differs from other Jahnula species in having asymmetric
ascospores (Fig. 6i). The phylogenetic placement of J. apiospora is pending awaiting molecular data, but it clearly does
not belong in this order. To clearly describe pseudoparaphyses
is important in fungal classification (Liew et al. 2000), and
Fungal Diversity (2020) 105:319–575
also difficult as in some species they deliquesce at maturity.
Therefore, it is important to study the early stages of ascoma,
which was earlier proposed by Eriksson (1981).
Future work
The world map in Fig. 1 illustrates the world distribution
of freshwater Dothideomycetes. Among 35 countries from
where freshwater Dothideomycetes have been collected, China
has almost 150 species, followed by Thailand with almost 140
species (Fig. 1). This is undoubtedly because many researchers
reside in these areas, however, Thailand has been shown to have
an amazing diversity (Hyde et al. 2018b). Studies of freshwater
fungi in poorly studied areas (shown in Fig. 1) are encouraged
as they should reveal numerous undiscovered species.
It is now becoming more and more urgent to study freshwater fungi due to the melting of glaciers with climate
change (Vargo et al. 2020; Wang et al. 2020). This has had a
major impact on the global climate, such as storms, flooding,
heatwaves and drought. This may cause the disappearance
of freshwater in certain areas or periodic drying of streams
and rivers and result in the extinction of freshwater species,
including fungi (Tsui et al. 1998). Thus the diversity should
be studied before it disappears.
Several freshwater genera comprise only a single species
(Table 2), e.g. Alascospora evergladensis, Angustospora nilensis
and Aquatospora cylindrica. It is highly likely that other species
exist as has been shown in Acrogenospora, where nine species
were discovered in a relatively small region (Bao et al. 2020).
Additionally, some speciose genera (Index Fungorum 2020),
contain only a single freshwater species, e.g. Byssosphaeria
schiedermayeriana, Herpotrichia dalisayi, Keissleriella linearis,
Lophiotrema rubi, Melanomma australiense and Splanchnonema
britzelmayrianum. Whether those species are terrestrial immigrants from nearby forests, different genera or regular freshwater
inhabitants is unclear. Thus, a better understanding of the ecology
of freshwater fungi is needed.
Acknowledgements The authors would like to thank the Sichuan Science and Technology Program (Grant No. 2019YFH0175, Grant No.
2018SZDZX0020), the National Key R&D Program of China (Grant
No. 2019YFC1803504; Grant No. 2016YFC0502204). Huang Zhang
would like to thank National Natural Science Foundation of China (Project ID: NSF 31500017 to Huang Zhang) and State Scholarship fund
of China (201908530031). Kevin D. Hyde thanks the Foreign Experts
Bureau of Yunnan Province, Foreign Talents Program (2018; Grant No.
YNZ2018002), Thailand Research grant entitled Impact of climate change
on fungal diversity and biogeography in the Greater Mekong Subregion
(Grant No. RDG6130001). Kevin D. Hyde also thanks Chiang Mai University for the award of Visiting Professor. Mingkwan Doilom would like
to thank the 5th batch of Postdoctoral Orientation Training Personnel in
Yunnan Province (Grant No. Y934283261) and the 64th batch of China
Postdoctoral Science Foundation (Grant No. Y913082271). Rungtiwa
Phookamsak thanks CAS President’s International Fellowship Initiative
(PIFI) for young staff (Grant No. Y9215811Q1), the National Science
Foundation of China (NSFC) project code 31850410489 (Grant No.
Y81I982211) and Chiang Mai University for financial support. Sinang
Honsanan would like to thank National Natural Science Foundation of
559
China for supporting the project Biodiversity, Taxonomy, Phylogeny,
Evolution and Phytogeography of phytopathogens in Dothideomycetes
from Southern China (Grant No. 31950410548) for funding this research.
Dhanushka Wanasinghe would like to thank CAS President’s International
Fellowship Initiative (PIFI) for funding his postdoctoral research (number
2019PC0008), the 64th batch of China Postdoctoral Science Foundation (Grant No. Y913083271) and the National Science Foundation of
China and the Chinese Academy of Sciences (Grant No. 41761144055).
Saranyaphat Boonmee would like to thank the Thailand Research Fund
(No. TRG6180001) and Mae Fah Luang University Research Fund (No.
631C15001). Wei Dong would like to thank Dr. Chayanard Phukhamsakda
for her suggestions on the manuscript. Prof. D. Jayarama Bhat is sincerely
thanked for his review on hyphomycetes. Dr. Ying Zhang is thanked for
her help on the discussion of Neomassariosphaeria. Chenxue Liu, Mingjie
Wei, Chunxue Li, Melvina J. D’souza, Dr. Haixia Wu and Prof. Di Zhang
are thanked for their helps on the herbarium work. We would like to thank
Dr. Shaun Pennycook for his essential nomenclatural review. Kazuaki
Tanaka would like to thank the Japan Society for the Promotion of Science (JSPS, 19K06802).
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
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copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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Zhang Y, Zhang XD, Fournier J, Chen JJ, Hyde KD (2014c) Lindgomyces griseosporus, a new aquatic ascomycete from Europe
including new records. Mycoscience 55:43–48
Zhang JQ, Zhou YP, Dou ZP, Fournier J, Zhang Y (2015) Helicascus
unilocularis sp. nov., a new freshwater pleosporalean ascomycete
from the Caribbean area. Mycol Prog 14:1–8
Zhang H, Dong W, Hyde KD, Bahkali AH, Liu JK, Zhou DQ, Zhang D
(2016) Molecular data shows Didymella aptrootii is a new genus
in Bambusicolaceae. Phytotaxa 247:99–108
Zhang H, Dong W, Hyde KD, Maharachchikumbura SSN, Hongsanan
S, Jayarama Bhat D, Al-Sadi AM, Zhang D (2017a) Towards a
natural classification of Annulatascaceae-like taxa: introducing
Atractosporales ord. nov. and six new families. Fungal Divers
85:75–110
Zhang JF, Liu JK, Hyde KD, Yang W, Liu ZY (2017b) Fungi from
Asian Karst formations II. Two new species of Occultibambusa
(Occultibambusaceae, Dothideomycetes) from karst landforms
of China. Mycosphere 8:550–559
Zhang SN, Hyde KD, Jones EBG, Cheewangkoon R, Liu JK (2018)
Acuminatispora palmarum gen. et sp. nov. from mangrove habitats. Mycol Prog 17:1173–1188
Zhang SN, Hyde KD, Jones EBG, Jeewon R, Cheewangkoon R, Liu JK
(2019) Striatiguttulaceae, a new pleosporalean family to accommodate Longicorpus and Striatiguttula gen. nov. from palms.
MycoKeys 49:99–129
Zhang SN, Hyde KD, Jones EBG, Cheewangkoon R, Liu JK (2020)
Additions to Fissuroma and Neoastrosphaeriella (Aigialaceae,
Pleosporales) from palms. Mycosphere 11:269–284
Zhao GC, Zhao RL (2012) The higher microfungi from forests of Yunnan Province. Yunnan Science and Technology Press, Kunming
Zhao GZ, Liu XZ, Wu WP (2007) Helicosporous hyphomycetes from
China. Fungal Divers 26:313–524
Zhao YZ, Zhang ZF, Cai L, Peng WJ, Liu F (2018) Four new filamentous fungal species from newly-collected and hive-stored bee
pollen. Mycosphere 9:1089–1116
Zhou Y, Gong GS, Zhang SR, Liu N, Wang JJ, Li PL, Yu XM (2013)
A new species of the genus Trematosphaeria from China. Mycol
Prog 13:33–43
Zhu D, Luo ZL, Baht DJ, McKenzie EHC, Bahkali AH, Zhou DQ,
Su HY, Hyde KD (2016) Helminthosporium velutinum and H.
aquaticum sp. nov. from aquatic habitats in Yunnan Province.
China. Phytotaxa 253:179–190
Affiliations
Wei Dong1,2,3,4,5 · Bin Wang6 · Kevin D. Hyde4,5,7,8 · Eric H. C. McKenzie9 · Huzefa A. Raja10 · Kazuaki Tanaka11 ·
Mohamed A. Abdel‑Wahab12 · Faten A. Abdel‑Aziz12 · Mingkwan Doilom5,13,14,15 · Rungtiwa Phookamsak8,13,14,15 ·
Sinang Hongsanan16 · Dhanushka N. Wanasinghe13,14,15 · Xian‑Dong Yu1,17 · Gen‑Nuo Wang18 · Hao Yang1 ·
Jing Yang4,19 · Kasun M. Thambugala20 · Qing Tian4,7 · Zong‑Long Luo21 · Jian‑Bo Yang13,14,15 · Andrew N. Miller22 ·
Jacques Fournier23 · Saranyaphat Boonmee4,19 · Dian‑Ming Hu24 · Sarunya Nalumpang2,3 · Huang Zhang1,25,26
1
Faculty of Agriculture and Food, Kunming University
of Science and Technology, Kunming 650500, China
11
Faculty of Agriculture and Life Science, Hirosaki University,
Bunkyo-cho3, Hirosaki, Aomori 035-8561, Japan
2
Department of Entomology and Plant Pathology, Faculty
of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
12
Department of Botany and Microbiology, Faculty of Science,
Sohag University, Sohag 82524, Egypt
13
CAS Key Laboratory for Plant Diversity and Biogeography
of East Asia, Kunming Institute of Botany, Chinese Academy
of Sciences, Kunming 650201, Yunnan, China
3
Innovative Agriculture Research Center, Faculty
of Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
14
4
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Mai 57100, Thailand
Centre for Mountain Futures (CMF), Kunming Institute
of Botany, Kunming 650201, Yunnan, China
15
East and Central Asia Regional Office, World Agroforestry
Centre (ICRAF), Kunming 650201, Yunnan, China
16
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Sciences and Oceanography, Shenzhen
University, Shenzhen 518060, China
17
School of Life Science and Technology, University
of Electronic Science and Technology of China,
Chengdu 611731, China
5
6
Innovative Institute of Plant Health, Zhongkai University
of Agriculture and Engineering, Haizhu District,
Guangzhou 510225, China
School of Environment and Resource, Southwest University
of Science and Technology, Sichuan 621010, China
7
Mushroom Research Foundation, 128 M.3 Ban Pa Deng
T. Pa Pae, A. Mae Taeng, Chiang Mai 50150, Thailand
8
Research Center of Microbial Diversity and Sustainable
Utilization, Faculty of Sciences, Chiang Mai University,
Chiang Mai 50200, Thailand
18
Faculty of Environmental Science and Engineering, Kunming
University of Science & Technology, Kunming 650500,
China
9
Manaaki Whenua-Landcare Research, Private Bag 92170,
Auckland, New Zealand
19
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
10
Department of Chemistry and Biochemistry, University
of North Carolina, Greensboro 27413, USA
13
Fungal Diversity (2020) 105:319–575
575
20
Genetics and Molecular Biology Unit, Faculty of Applied
Sciences, University of Sri Jayewardenepura, Gangodawila,
Nugegoda 10250, Sri Lanka
24
Bioengineering and Technological Research Centre
for Edible and Medicinal Fungi, Jiangxi Agricultural
University, Nanchang 330045, China
21
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, China
25
Department of Botany, University of British Columbia,
Vancouver, BC V6T 1Z4, Canada
22
Illinois Natural History Survey, University of Illinois,
Champaign, IL 61820, USA
26
23
Las Muros, 09420 Rimont, France
Yunnan Key Lab of Soil Carbon Sequestration and Pollution
Control, Kunming University of Science and Technology,
Kunming 650500, China
13
Saranyaphat Boonmee