Fungal Diversity (2019) 99:451–660
https://doi.org/10.1007/s13225-019-00438-1
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Freshwater Sordariomycetes
Zong-Long Luo1,2 • Kevin D. Hyde2 • Jian-Kui (Jack) Liu3 • Sajeewa S. N. Maharachchikumbura3,4 •
Rajesh Jeewon5 • Dan-Feng Bao1,6 • Darbhe Jayarama Bhat7 • Chuan-Gen Lin2 • Wen-Li Li1 •
Jing Yang2 • Ning-Guo Liu2 • Yong-Zhong Lu8 • Ruvishika S. Jayawardena2 • Jun-Fu Li2 • Hong-Yan Su1
Received: 30 July 2019 / Accepted: 3 October 2019 / Published online: 11 November 2019
Ó The Author(s) 2019
Abstract
Sordariomycetes is one of the largest classes of Ascomycota that comprises a highly diverse range of fungi mainly
characterized by perithecial ascomata and inoperculate unitunicate asci. Freshwater Sordariomycetes play an important
role in ecosystems and some of them have the potential to produce bioactive compounds. This study documents and
reviews the freshwater Sordariomycetes, which is one of the largest and important groups of fungi in aquatic habitats.
Based on evidence from DNA sequence data and morphology, we introduce a new order Distoseptisporales, two new
families, viz. Ceratosphaeriaceae and Triadelphiaceae, three new genera, viz. Aquafiliformis, Dematiosporium and
Neospadicoides, 47 new species, viz. Acrodictys fluminicola, Aquafiliformis lignicola, Aquapteridospora fusiformis,
Arthrinium aquaticum, Ascosacculus fusiformis, Atractospora aquatica, Barbatosphaeria lignicola, Ceratosphaeria
aquatica, C. lignicola, Chaetosphaeria aquatica, Ch. catenulata, Ch. guttulata, Ch. submersa, Codinaea yunnanensis,
Conioscypha aquatica, C. submersa, Cordana aquatica, C. lignicola, Cosmospora aquatica, Cylindrotrichum submersum,
Dematiosporium aquaticum, Dictyochaeta cangshanensis, D. ellipsoidea, D. lignicola, D. submersa, Distoseptispora
appendiculata, D. lignicola, D. neorostrata, D. obclavata, Hypoxylon lignicola, Lepteutypa aquatica, Myrmecridium
aquaticum, Neospadicoides aquatica, N. lignicola, N. yunnanensis, Ophioceras submersum, Peroneutypa lignicola,
Phaeoisaria filiformis, Pseudostanjehughesia lignicola, Rhodoveronaea aquatica, Seiridium aquaticum, Sporidesmiella
aquatica, Sporidesmium lageniforme, S. lignicola, Tainosphaeria lunata, T. obclavata, Wongia aquatica, two new combinations, viz. Acrodictys aquatica, Cylindrotrichum aquaticum, and 9 new records, viz. Chaetomium globosum, Chaetosphaeria cubensis, Ch. myriocarpa, Cordana abramovii, Co. terrestris, Cuspidatispora xiphiago, Sporidesmiella
hyalosperma, Stachybotrys chartarum, S. chlorohalonata. A comprehensive classification of the freshwater Sordariomycetes is presented based on updated literature. Phylogenetic inferences based on DNA sequence analyses of a combined LSU, SSU, RPB2 and TEF1a dataset comprising species of freshwater Sordariomycetes are provided. Detailed
information including their habitats distribution, diversity, holotype, specimens collected and classification are provided.
Keywords 47 new taxa Asexual morphs Phylogeny Sexual morphs Taxonomy
Introduction
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s13225-019-00438-1) contains supplementary material, which is available to authorized users.
& Hong-Yan Su
suhongyan16@163.com
Extended author information available on the last page of the article
Shearer (1993) defined freshwater ascomycetes in a broad
ecological sense as ‘‘all ascomycetes that occur on submerged or partially submerged substrates in aquatic habitats’’. Freshwater fungi have also been defined as ‘‘fungi
that the whole or part of their life cycle rely on freshwater’’
(Thomas 1996). The definition of Shearer (1993) has been
regarded as the best current working definition for the
freshwater ascomycetes (Cai et al. 2003a, 2014), and we
follow the definition of Shearer (1993) in this study.
Lignicolous freshwater fungi are those that grow on
submerged woody debris in freshwater streams, ponds,
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lakes, tree hollows, peat swamps and dams (Ho et al.
2002a; Vijaykrishna et al. 2005; Hyde et al. 2016a). They
play an important role in nutrient and carbon cycling,
biological diversity and ecosystem functioning of freshwater ecosystems with their ability to decompose lignocellulose in woody litter, softening the wood and releasing
nutrients (Hyde et al. 2016a). The lignicolous freshwater
fungi are highly diverse in the classes Dothideomycetes
and
Sordariomycetes
(Hyde
et
al.
2013;
Maharachchikumbura et al. 2016; Liu et al. 2017; Lu et al.
2018a). There are a few freshwater taxa that belong to
Eurotiomycetes (Liu et al. 2015a, b), Orbiliomycetes (Swe
et al. 2009) and rarely Basidiomycetes (Hyde and Goh
1998a, 1999; Jones et al. 2014) collected from submerged
wood in freshwater habitats.
Sordariomycetes is the second largest class of
Ascomycota (Kirk et al. 2008; Hyde et al. 2013;
Maharachchikumbura et al. 2015, 2016). Species of Sordariomycetes are mainly characterized by non-lichenized,
perithecial ascomata and inoperculate unitunicate or nonfissitunicate asci (Zhang et al. 2006; Kirk et al. 2008). The
class Sordariomycetes has a cosmopolitan distribution and
accommodates mostly terrestrial taxa (Tang et al. 2007;
Maharachchikumbura et al. 2015, 2016; Hongsanan et al.
2017), but there are several sordariomycetous species that
have been reported from aquatic habitats (Hyde and Wong
2000; Cai and Hyde 2007; Raja et al. 2009a, b; Réblová
et al. 2010, 2016a; Shearer et al. 2010; Hu et al. 2013; Cai
et al. 2014; Jones et al. 2015; Lu et al. 2016; Su et al. 2016;
Yang et al. 2017, 2018a, b; Luo et al. 2018a, b).
Freshwater Sordariomycetes play an important role in
ecosystem functioning and many of them have shown
potential application value (Cai et al. 2014). Annulatascaceous, distoseptisporaceous, pleurotheciaceous and
halosphaeriaceous species are the most typical and common freshwater Sordariomycetes on submerged wood
(Hyde et al. 1999a; Cai et al. 2014; Li et al. 2016a; Réblová
et al. 2016a; Zhang et al. 2017a, b; Luo et al. 2018a; Yang
et al. 2018a). The modern classification system of the
overall Kingdom of Fungi, as well as Sordariomycetes,
have been significantly improved with the utilization of
molecular sequence data (Hibbett et al. 2007;
Maharachchikumbura et al. 2015; Spatafora et al. 2017).
Cai et al. (2014) provided a phylogenetic analysis of
freshwater Sordariomycetes based on LSU sequence data.
Their analysis showed that freshwater Sordariomycetes are
scattered in three subclasses, Sordariomycetidae,
Hypocreomycecetidae and Xylariomycetidae including 13
orders. However, during the last four years, several Sordariomycetes species have been introduced comprising
new genera, families, orders, subclasses and some of them
were collected from freshwater habitats (Réblová et al.
2015a, b, 2016a, b; Maharachchikumbura et al. 2015, 2016;
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Fungal Diversity (2019) 99:451–660
Su et al. 2016; Hongsanan et al. 2017; Wijayawardene
et al. 2017, 2018; Yang et al. 2017, 2018a, b; Zhang et al.
2017a, b; Song et al. 2018a, b; Wei et al. 2018).
In this study, 129 fresh isolates of sordariomyceteous
taxa were collected from freshwater habitats in China and
Thailand. One new order, two new families, three new
genera, 47 new species, two new combinations and 9 new
records are introduced. A modified backbone tree based on
phylogenetic analysis of combined LSU, SSU, RPB2 and
TEF1a sequence data of the freshwater Sordariomycetes is
provided. Phylogenetic trees under genus or family level
are also constructed in order to confirm the classification of
the new isolates.
Materials and methods
Sample collection and specimen examination
Submerged decaying wood samples were collected from
freshwater habitats in Yunnan Province (China) and Chiang Rai, Chiang Mai (Thailand) from 2013 to 2018. The
process of morphological studies follows Luo et al.
(2018a). Single spore isolates were obtained following the
method of Chomnunti et al. (2014) and cultured according
to Liu et al. (2010). Herbarium materials are deposited in
the Herbarium of Mae Fah Luang University (Herb.
MFLU), Chiang Rai, Thailand, Herbarium of Cryptogams
Kunming Institute of Botany Academia Sinica (Herb.
HKAS) and Dali University (Herb. DLU), Yunnan, China.
Living cultures are deposited at Mae Fah Luang University
Culture Collection (MFLUCC), Thailand, Kunming Institute of Botany culture collection(KUMCC) and culture
collection at Dali University (DLUCC) China. Facesoffungi and Index Fungorum numbers are provided (Jayasiri
et al. 2015; Index Fungorum 2019). New taxa are established based on guidelines outlined by Jeewon and Hyde
(2016). Taxa deposited in HKU(M) mostly lack sequence
data and were moved to IFRD in Kunming, China. These
need to be recollected so that reference specimens (sensu
Ariyawansa et al. 2014) can be designated.
DNA extraction, PCR amplification
and sequencing
Genomic DNA was extracted from fungal mycelium grown
on PDA or MEA at room temperature by using a EZ gene
TM Fungal gDNA kit (GD2416) according to the manufacturer’s instructions. The gene regions of the large subunit of the nuclear ribosomal DNA (LSU), the internal
transcribed spacers (ITS), the small subunit of the nuclear
ribosomal DNA (SSU), the translation elongation factor
(TEF1a) and RNA polymerase II subunit 2 (RPB2) were
Fungal Diversity (2019) 99:451–660
amplified using the primer pairs LR0R/LR7 (Vilgalys and
Hester 1990), ITS5/ITS4, NS1/NS4 (White et al. 1990),
983F/2218R (Rehner and Buckley 2005), fRPB2-5F/
fRPB2-7cR (Liu et al. 1999) respectively. The ITS, LSU,
SSU, RPB2 and TEF1a amplification reactions were carried out using the method described by Luo et al. (2018a).
DNA sequencing were performed with the primers mentioned above at Tsingke Biological Engineering Technology and Services Co., Ltd (Yunnan, P.R. China).
Phylogenetic analysis
Sequences generated from different primers were analyzed
with other sequences obtained from GenBank. The
sequences were deposited in GenBank and the accession
numbers in the analyses were provided in Supplementary
material 1. The sequence data were aligned using MAFFT
v.7.110 online program (http://mafft.cbrc.jp/alignment/ser
ver/) (Katoh and Standley 2013) and manually adjusted via
BioEdit v.7.2.3 (Hall 1999).
‘‘ALTER’’ (Glez-Peña et al. 2010) was used to format
the aligned fasta file for RAxML analysis. Maximum
likelihood (ML) analysis was performed at the CIPRES
Science Gateway v.3.3 (http://www.phylo.org/portal2/;
Miller et al. 2010) using RAxML v.8.2.10 as part of the
‘‘RAxML-HPC2 on XSEDE’’ tool (Stamatakis et al. 2008;
Stamatakis 2014). All free model parameters were estimated by RAxML with ML estimates of 25 per site rate
categories. The final ML search was conducted using the
GTRGAMMA ? I model. The best scoring tree was
selected with a final likelihood value of - 194313.319877.
Phylogenetic tree was visualized using FigTree v1.4.0
(http://tree.bio.ed.ac.uk/software/figtree/, Rambaut 2012).
Results
Phylogenetic analysis of combined LSU, SSU,
RPB2 and TEF1a sequence data
The combined LSU, SSU, RPB2 and TEF1a sequence
dataset comprised 857 taxa with Dothidea sambuci
(AFTOL ID 274), Leotia lubrica (AFTOL ID 1) and Microglossum rufum (AFTOL ID 1292) as the outgroup taxa.
The combined aligned sequence matrix comprises LSU
(768 bp), SSU (884 bp), RPB2 (850 bp) and TEF1a
(815 bp) sequence data for 857 taxa with a total of 3317
characters (Including the gaps), of which 1046 characters
were constant, 391 variable characters were parsimonyuninformative and 1880 characters were parsimony informative. The RAxML tree is shown in Fig. 1.
In the phylogenetic tree (Fig. 1), the 854 strains (including 129 fresh isolates) of Sordariomycetes included in
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the analysis cluster into six subclasses: Sordariomycetidae,
Hypocreomycetidae and Xylariomycetidae as in the previous treatments of Lumbsch and Huhndorf (2010), Diaporthomycetidae and Lulworthiomycetidae as suggested by
Maharachchikumbura et al. (2015), as well as Savoryellomycetidae as suggested by Hongsanan et al. (2017). Of
these included strains, 356 were isolated from freshwater
habitats and distributed in 47 clades as follows:
Clade 1 represents the family Junewangiaceae with ten
species which belong to three genera, viz. Dictyosporella,
Junewangia and Sporidesmiella. In this study, we provide
sequence data for Sporidesmiella novae-zelandiae and S.
hyalosperma for the first time, and introduce a new species
S. aquatica.
Clade 2 represents the family Pseudoproboscisporaceae
with four species from four genera, viz. Aquaticola
hyalomura, Cateractispora recepticuli, Diluvicola aquatica, Pseudoproboscispora thailandensis.
Clade 3 represents the family Atractosporaceae. Presently, Atractosporales comprises the single family Atractosporaceae which includes two genera. All taxa of
Atractosporales are collected from freshwater habitats
(Réblová et al. 2016b; Zhang et al. 2017a, b). Two freshwater fungal strains obtained from this study cluster together with other Atractospora species and represent an
independent lineage with strong support (100% ML). We
therefore introduce a new species Atractospora aquatica
sp. nov.
Clade 4 represents the genus Cancellidium with two
strains of Cancellidium applanatum. These two strains
were collected from freshwater habitats (Zelski et al.
2014).
Clade 5 represents the family Conlariaceae including
three freshwater species of Conlarium, and one Riomyces
species collected from freshwater.
Clade 6 represents the order Xenospadicoidales which
was introduced by Réblová et al. (2018). Five freshwater
fungal strains obtained in this study cluster together and
represent an independent lineage with strong support
(100% ML) within Xenospadicoidaceae. A new genus
Neospadicoides is introduce herein based on phylogeny
and morphology.
Clade 7 represents the order Sporidesmiales with a
single family Sporidesmiaceae. Most of the species in
Sporidesmiaceae are reported from freshwater habitats. For
the first time, Zhang et al. (2017a, b) provided the sexual
morph for the genus Sporidesmium. In this study, we
introduce two new species, Sporidesmium lageniforme and
Sporidesmium lignicola, based on both morphological
characters and phylogenetic evidence, while S. lignicola is
reported with both of asexual and sexual morphs.
Clade 8 represents the family Rhamphoriaceae established by Réblová and Štěpánek (2018) which includes four
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b Fig. 1 Phylogram generated from maximum likelihood analysis
based on combined LSU, SSU, RPB2 and TEF1a sequence data for
the species of Sordariomycetes. RAxML bootstrap support values
equal to or greater than 75% are given above the nodes. Newly
generated sequences are in red. Strains of known species collected
from freshwater habitats are marked as ‘‘green circle’’ followed by
strain number. Ex-type strains are in bold
genera. We introduce a new Rhodoveronaea species which
is the first species described from freshwater in this family.
Clade 9 represents the family Acrodictyaceae established by Xia et al. (2017) with a single genus Acrodictys.
A previously described species, Barbatosphaeria aquatica
MFLUCC 18–0356 (Hyde et al. 2018), is synonymized
under Acrodictys aquatica. We introduce a new species,
Acrodictys fluminicola.
Clade 10 represents the monotypic genus Pseudostanjehughesia introduced by Yang et al. (2018a) with P.
aquitropica as the type species and it was collected from
freshwater habitats. We introduce the second species, P.
lignicola sp. nov.
Clade 11 represents the family Papulosaceae established
by Winka and Eriksson (2000). Papulosaceae is typified by
the monotypic marine genus Papulosa (Winka and Eriksson 2000).
Clade 12 represents the species Sporidesmium tropicale
(HKUCC 10838, MFLUCC 16–0185). The strain
MFLUCC 16–0185 was obtained from the specimen collected from a freshwater habitat (Yang et al. 2018a).
Clade 13 represents the typical freshwater genus Bullimyces introduced by Ferrer et al. (2012).
Clade 14 represents the family Barbatosphaeriaceae
which was introduced by Zhang et al. (2017a, b). We
introduce a new species, Barbatosphaeria lignicola based
on morphology and phylogeny.
Clade 15 represents the new order Distoseptisporales
introduced herein. Su et al. (2016) introduced the family
Distoseptisporaceae to accommodate Sporidesmium-like
taxa based on morphology and phylogeny. Phylogenetic
analysis based on combined LSU, SSU, RPB2 and TEF1a
sequence data show that the species of Distoseptisporales
cluster together with strong support and form a distinct,
strongly supported clade within Diaporthomycetidae.
Clade 16 represents two strains of the genus Cyanoannulus. Zhang et al. (2017a, b) established the family
Woswasiaceae to accommodate Woswasia, Xylochrysis
and Cyanoannulus in Diaporthomycetidae families incertae sedis with weak support. In our phylogenetic analysis,
species of Cyanoannulus formed a distinct clade basal to
the order Distoseptisporales and therefore we suggest to
place Cyanoannulus in Diaporthomycetidae genera incertae sedis.
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Clade 17 represents the order Annulatascales introduced
by Maharachchikumbura et al. (2015). The family Annulatascaceae is accepted in this order. Some species of
Annulatascaceae are lacking sequence data and therefore it
is not possible to resolve their phylogenetic affinities.
Clade 18 represents the order Myrmecridiales established by Crous et al. (2015) with a single family,
Myrmecridiaceae. We introduce a new species Myrmecridium aquaticum. Presently, three species (M. aquaticum,
M. fluviae, M. montsegurinum) are collected from freshwater habitats.
Clade 19 represents the family Ophioceraceae, with
eight species which were reported from freshwater habitats.
We introduce a new species, Ophioceras submersum.
Clade 20 represents the new family Ceratosphaeriaceae
introduced herein. The order Magnaporthales was introduced by Thongkantha et al. (2009) to accommodate
Magnaporthaceae based on morphological characters of
both asexual and sexual morphs together with phylogenetic
analyses of combined LSU and SSU sequence data. There
are four families accepted in Magnaporthales, viz. Magnaporthaceae, Ophioceraceae, Pseudohalonectriaceae,
Pyriculariaceae. We introduce the new family Ceratosphaeriaceae to accommodate Ceratosphaeria species.
Clade 21 represents the family Pseudohalonectriaceae
established by Hongsanan et al. (2017) based on the genus
Pseudohalonectria. Five strains collected from freshwater
habitats are included in our phylogentic tree.
Clade 22 represents the family Magnaporthaceae introduced by Cannon (1994). We introduce a new monotypic
genus Aquafiliformis.
Clade 23 represents the family Tirisporellaceae with a
freshwater species, Thailandiomyces bisetulosus (BCC
00018, BCC 00200) which was introduced by Pinruan et al.
(2008).
Clade 24 represents the family Jobellisiaceae with a
freshwater species, Jobellisia guangdongensis (GD14–4)
collected from southern China (Liu et al. 2011a, b).
Clade 25 represents the freshwater genus Hyalorostratum established by Raja et al. (2010) with Hyalorostratum
brunneisporum as the type species.
Clade 26 represents the family Chaetosphaeriaceae
established by Réblová et al. (1999) based on Chaetosphaeria (Tulasne and Tulasne 1863). We introduce 11
new species in this family, viz. Chaetosphaeria aquatica,
C. catenulata, C. guttulata, C. submersa, Codinaea yunnanensis, Dictyochaeta cangshanensis, D. ellipsoidea, D.
lignicola, D. submersa, Tainosphaeria lunata and T.
obclavata. Chloridium aseptatum (MFLUCC 11–0216)
(Wei et al. 2018) is synonymized under Chloridium
gonytrichii.
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Fig. 1 continued
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Fig. 1 continued
Clade 27 represents the family Phyllachoraceae with
five strains. Phyllachoraceae sp. WF33A was collected
from freshwater habitats.
Clade 28 represents the single isolate of Clohiesia. The
species Clohiesia corticola (HKUCC 3712) which was
collected from freshwater, clusters with species of Phyllachoraceae with weak support.
Clade 29 represents the order Sordariales established by
Hawksworth and Eriksson (1986) and comprises three
families, viz. Chaetomiaceae, Sordariaceae and
Lasiosphaeriaceae sensu lato. Six freshwater species are
represented in this clade and most of them are in
Lasiosphaeriaceae.
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Clade 30 represents the family Cordanaceae. We introduce two new species herein, Cordana aquatica and C.
lignicola, based on morphology and phylogeny. We also
provide descriptions, illustrations and molecular sequence
data for Cordana abramovii and C. terrestris.
Clade 31 represents the family Coniochaetaceae with
five Coniochaeta species and Coniochaeta gigantospora
(ILLS 60816) which was originally collected from a
freshwater habitat (Raja et al. 2012).
Clade 32 represents the family Sporocadaceae and we
introduce a new species, Seiridium aquaticum.
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Fig. 1 continued
Clade 33 represents the family Amphisphaeriaceae
established by Winter (1885). We introduce a new species,
Lepteutypa aquatica.
Clade 34 represents the family Apiosporaceae established by Hyde et al. (1998a). A new species, Arthrinium
aquaticum is introduced.
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Fig. 1 continued
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Fig. 1 continued
Clade 35 represents a new species Peroneutypa lignicola. The genus Peroneutypa has been accomodated in
Diatrypaceae (Shang et al. 2018). In our multi-gene phylogenetic analysis, the new species forms a distinct
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subclade basal to Lopadostomataceae but close to Diatrypaceae, although this relationship is not supported.
Clade 36 represents the species Sporidesmium gyrinomorphum (MFLUCC 16–0186) introduced by Yang
et al. (2018a, b).
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Fig. 1 continued
Clade 37 represents the family Hypoxylaceae with four
Hypoxylon strains. We introduce a new species Hypoxylon
lignicola based on morphology and phylogeny. Wendt
et al. (2018) emended Hypoxylaceae and accepted 14
genera in this family.
Clade 38 represents the family Nectriaceae including 66
strains. Lombard et al. (2015) accepted 47 genera in this
family based on molecular sequence data. We introduce a
new species Cosmospora aquatica, based on morphology
and phylogeny. DNA sequence data for Aquanectria
jacinthicolor, A. penicillioides, Chaetopsina beijingensis,
Mariannaea samuelsii, M. superimposita and Paracremonium binnewijzendii are provided.
Clade 39 represents the family Stachybotriaceae established by Crous et al. (2014a) in the order Hypocreales to
accommodate the genera Myrothecium, Peethambara and
Stachybotrys. In this study, morphological characterization
and DNA sequence data for Stachybotrys chartarum and S.
chlorohalonata are provided.
Clade 40 represents the family Halosphaeriaceae. In this
study, we introduce a new Ascosacculus species, A. fusiformis, based on morphology and phylogeny.
Clade 41 represents the new family Triadelphiaceae that
we introduce herein based on its distinct morphology and
molecular phylogeny.
Clade 42 represents the family Reticulascaceae. We
introduced a new species herein, Cylindrotrichum submersum sp. nov. and synonymise Blastophorum aquaticum
(Hyde et al. 2016) under Cylindrotrichum aquaticum.
Clade 43 represents one of most common and typical
freshwater family Pleurotheciellaceae established by
Réblová et al. (2016a). We introduce a new species,
Phaeoisaria filiformis.
Clade 44 represents another typical freshwater fungal
family Savoryellaceae. We introduce a new genus Dematiosporium in this family.
Clade 45 represents the order Conioscyphales established by Réblová et al. (2016a) with a single family
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Conioscyphaceae. We introduce two new species, Conioscypha aquatica and C. submersa.
Clade 46 represents the monotypic order Fuscosporellales established by Yang et al. (2016a) based on the family
Fuscosporellaceae. Most of the species in this order are
reported from freshwater habitats.
Clade 47 represents the genus Flammispora established
by Pinruan et al. (2004a).
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Taxonomy
Based on the outline and multigene phylogeny of freshwater Sordariomycetes provided in this study, freshwater
Sordariomycetes are scattered in six sub-classes, viz. Diaporthomycetidae, Hypocreomycetidae, Lulworthiomycetidae, Savoryellomycetidae, Sordariomycetidae and
Xylariomycetidae. In this section, we provide the notes for
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Fig. 1 continued
each freshwater genera and species including information
on known distribution, herbarium and sequence data. The
outline of freshwater Sordariomycetes based on literature
up to November 2018 was provided in Supplementary
material 2. Descriptions, illustrations and sequence data for
the fungi collected from freshwater habitats in China and
Thailand from 2013 to 2018 are also provided.
Detailed information of freshwater Sordariomycetes are as
follows:
Diaporthomycetidae Senan. et al.
Annulatascales D’souza et al.
Annulatascaceae Wong et al.
Annulatascus K.D. Hyde, Aust. Syst. Bot. 5(1): 118 (1992)
Asexual morph Undetermined. Sexual morph Updated
generic
description
and
illustrations
see
Maharachchikumbura et al. (2016).
Type species: Annulatascus velatisporus K.D. Hyde,
Aust. Syst. Bot. 5(1): 118 (1992)
Notes: Hyde (1992a) introduced the genus Annulatascus
to accommodate two ascomycete species, A. velatisporus
and A. bipolaris, with the former as the type species and
both of them were collected from submerged decaying
wood in Australia. There are 19 epithets of Annulatascus
listed in Index Fungorum (December, 2018), however
Hyde et al. (1999a, b) transferred Annulatascus bipolaris to
the genus Cataractispora as C. bipolaris based on the
morphological characters. Subsequently, Campbell and
Shearer (2004) established a new genus Annulusmagnus for
Annulatascus triseptatus based on LSU sequence data, and
Luo et al. (2015) combined Annulatascus biatriisporus as
Pseudoannulatascus biatriisporus. Réblová et al. (2018)
synonymised Annulatascus biatriisporus and Pseudoannulatascus biatriisporus under Torrentispora biatriispora,
based on phylogenetic analysis. Currently, 16 species are
accepted in Annulatascus and most of them are reported
from freshwater habitats in tropical areas (Barbosa et al.
2008; Shearer et al. 2010; Boonyuen et al. 2012; Hu et al.
2012a) and only two species (A. citrisporus and A.
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Fig. 1 continued
licualae) are known from terrestrial habitats on palm
rachides (Fröhlich and Hyde 2000).
Annulatascus apiculatus F.R. Barbosa & Gusmão
Distribution: Brazil, on submerged decaying wood in a
stream (Barbosa et al. 2008)
Asexual morph: Undetermined
Notes: Holotype HUEFS 134723. Sequence data is not
available.
Annulatascus aquaticus Ho et al.
Distribution: China, Hong Kong, Tai Po Kau Forest
stream, on submerged decaying wood (Ho et al. 1999a).
Asexual morph: Undetermined
123
470
Fungal Diversity (2019) 99:451–660
Fig. 1 continued
Notes: Holotype HKU (M) 4526 (now in IFRD).
Sequence data is not available.
Annulatascus aquatorba Boonyuen & Sri-indrasutdhi
Distribution: Thailand, Narathiwat Province, Sirindhorn Peat Swamp Forest, on submerged wood test block of
Erythrophleum teysmannii (Boonyuen et al. 2012).
Asexual morph: Undetermined
Notes: Holotype BBH 29936. LSU and SSU sequence
data are available.
Annulatascus fusiformis K.D. Hyde & S.W. Wong
Distribution: Australia, Queensland, on submerged
wood (Vijaykrishna and Hyde 2006); China, Yunnan
Province, Yiliang county, Xiao Bai Long Mountain, on
submerged culms of Phyllostachys bambusoides in a
stream (Cai et al. 2006a); Philippines, Mindawao, Bukidnon, Impalutao, Natigbasan creek, on submerged wood
(Hyde and Wong 2000).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 3102 (now in IFRD).
Sequence data is not available.
Annulatascus hongkongensis Ho et al.
Distribution: China, Hong Kong, Plover Cove Reservoir, on submerged wood (Ho et al. 1999b).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 4702 (now in IFRD). LSU
sequence data is available.
Annulatascus joannae Tsui et al.
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Tsui et al. 2002).
Asexual morph: Undetermined
Notes: Holotype IFRD 8648. Sequence data is not
available.
123
Annulatascus lacteus Tsui et al.
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Tsui et al. 2002).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 4623 (now in IFRD).
Sequence data is not available.
Annulatascus liputii L. Cai & K.D. Hyde
Distribution: Philippines, on submerged bamboo in
Liput River (Cai et al. 2003b).
Asexual morph: Undetermined
Notes: Holotype PDD 75038. Sequence data is not
available.
Annulatascus menglensis Hu et al.
Distribution: China, Yunnan Province, Mengla,
Wudaoban Stream, on submerged wood (Hu et al. 2012a).
Asexual morph: Undetermined
Notes: Holotype IFRDSC 002–017. Sequence data is not
available.
Annulatascus nilensis Abdel-Wahab & Abdel-Aziz
Distribution: Egypt, Sohag, Nile River, on decayed
submerged stems of Phragmites australis (Abdel-Wahab
et al. 2011).
Asexual morph: Undetermined
Notes: Holotype IMI 397966. LSU sequence data is
available.
Annulatascus palmietensis Goh et al.
Distribution: Brunei, Tutong River, on submerged wood
(Fryar et al. 2004; Hyde and Sarma 2006); South Africa,
Durban, Palmiet River, on submerged wood (Hyde et al.
1998b).
Asexual morph: Undetermined
Fungal Diversity (2019) 99:451–660
Notes: Holotype IFRD 8649; Sequence data is not
available.
Annulatascus saprophyticus Z.L. Luo & K.D. Hyde
Distribution: Thailand, Chiang Mai Province, on submerged wood in a stream (Luo et al. 2015).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–0070, isotype HKAS 86445.
ITS, LSU, SSU, RPB2 and TEF1a sequence data are
available and obtained from ex-type culture.
Annulatascus tropicalis Ranghoo & K.D. Hyde
Distribution: China, Hong Kong, Tai Po, Plover Cove
Reservoir, on submerged wood (Tsui et al. 2002).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 5253 (now in IFRD).
Sequence data is not available.
Annulatascus velatisporus K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Hyde 1992a; Dayarathne et al. 2016); Brunei,
Temburong, Kuala Belalong Field Studies Centre, Sungai
Anak, on submerged wood, China, Hong Kong, New
Territories, Tai Po Kau Country Park, on twigs submerged
in a stream; Philippines, Mindanao, Bukidnon, Impalutao,
Natigbasan Creek, on submerged wood (Wong et al.
1999a); Japan, Koito River, on submerged wood (Tsui
et al. 2001a); India, Western Ghats, Kali River, submerged
woody litter (Sudheep and Sridhar 2011); Malaysia, Lipur
Lentang Nature Reserve, on submerged wood in a stream
(Ho et al. 2001); South Africa, Durban, Palmiet River, on
submerged wood (Hyde et al. 1998b); Thailand, Chiang
Mai Province, on submerged wood (Hu et al. 2012b).
Asexual morph: Undetermined
Notes: Holotype BRIP 17373, epitype MFLU 16–2204.
LSU and SSU sequence data are available.
Annulusmagnus J. Campb. & Shearer, Mycologia 96(4):
826 (2004)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell and Shearer (2004).
Type species: Annulusmagnus triseptatus (Wong et al.)
J. Campb. & Shearer, Mycologia 96(4): 826 (2004)
Notes: Campbell and Shearer (2004) introduced the
genus Annulusmagnus for Annulatascus triseptatus based
on LSU sequence data. Annulusmagnus triseptatus was first
collected from submerged wood in Brunei (Wong et al.
1999a) and subsequently reported from Australia, Canada
and the USA, on submerged corticated or decorticated
wood in both lotic and lentic habitats (Campbell and
Shearer 2004). There is only one species reported in this
genus (Campbell and Shearer 2004).
Annulusmagnus triseptatus (Wong et al.) J. Campb. &
Shearer
: Annulatascus triseptatus Wong et al., Mycol. Res.
103(5): 563 (1999)
471
Distribution: Australia, Koah, Clohiesy River, on submerged wood (Campbell and Shearer 2004); Brunei,
Temburong, Kuala Belalong Field Studies Centre, Sungai
Anak, on submerged wood (Wong et al. 1999a); Canada,
Ontario and Manitoba, on submerged wood; USA, on
submerged wood; Venezuela, on submerged wood
(Campbell and Shearer 2004).
Asexual morph: Undetermined
Notes: Holotype IFRD 8650. LSU, SSU and RPB2
sequence data are available.
Aqualignicola Ranghoo et al., Mycol. Res. 105(5): 628
(2001)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Ranghoo et al. (2001) and Hu et al.
(2012a).
Type species: Aqualignicola hyalina Ranghoo et al.,
Mycol. Res. 105(5): 628 (2001)
Notes: Aqualignicola was introduced by Ranghoo et al.
(2001) with Aqualignicola hyalina as type species. Currently, two species are accepted in this genus (Ranghoo
et al. 2001; Hu et al. 2012a), and both species were collected from freshwater habitats in Asia (Ranghoo et al.
2001; Sudheep and Sridhar 2011; Hu et al. 2012a).
Aqualignicola hyalina Ranghoo et al.
Distribution: China, Hong Kong, New Territories, Tai
Po, the Lam Tsuen River and Plover Cove Reservoir, on
submerged wood (Ranghoo et al. 2001); India, Western
Ghats, Kali River, submerged woody litter (Sudheep and
Sridhar 2011).
Asexual morph: Undetermined
Notes: Holotype IFRD 8680. Sequence data is not
available.
Aqualignicola vaginata Hu et al.
Distribution: China, Yunnan Province, Mengla,
Wudaoban Stream, on submerged wood (Hu et al. 2012a).
Asexual morph: Undetermined
Notes: Holotype IFRDC 021–043. Sequence data is not
available. Aqualignicola vaginata was introduced by Hu
et al. (2012a) and it is so far only known from the type
locality.
Ascitendus J. Campb. & Shearer, Mycologia 96(4): 829
(2004)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell and Shearer (2004).
Type species: Ascitendus austriacus (Réblová et al.) J.
Campb. & Shearer, Mycologia 96(4): 829 (2004)
Notes: Réblová and Winka (2001) introduced a new
species, Ascolacicola austriaca from a stream in Australia.
Campbell and Shearer (2004) examined 27 collections of
Ascolacicola austriaca and found its morphology to be
different from the type species of the genus. Based on the
morphological differences between Ascolacicola austriaca
123
472
and A. aquatica and phylogenetic analysis, a new genus
Ascitendus was proposed for Ascolacicola austriaca
(Campbell and Shearer 2004). Hyde et al. (2018) introduced the second species for this genus. Currently, two
species of Ascitendus are accepted and both were collected
from freshwater habitats (Réblová and Winka 2001;
Campbell and Shearer 2004; Hyde et al. 2018).
Ascitendus aquaticus Dayarathne et al.
Distribution: Australia, North Queensland, Mulgrave
River, on decaying wood submerged in a River (Hyde et al.
2018).
Asexual morph: Undetermined
Notes: Holotype MFLU 18–0143. LSU and SSU
sequence data are available.
Ascitendus austriacus (Réblová et al.) J. Campb. &
Shearer
: Ascolacicola austriaca Réblová et al., Mycologia 93:
486 (2001)
Distribution: Australia, Wien 19, Herman-nskogel, on
wood of Fagus sylvatica submerged in a stream (Réblová
and Winka 2001), Canada, Ontario and Manitoba, on
submerged wood; USA, on submerged wood; Venezuela,
on submerged wood (Campbell and Shearer 2004).
Asexual morph: Undetermined
Notes: Holotype PRM 842991. LSU sequence data is
available.
Ayria Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 248
(2004)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Fryar and Hyde (2004), Raja et al.
(2009a).
Type species: Ayria appendiculata Fryar & K.D. Hyde,
Cryptog. Mycol. 25(3): 248 (2004)
Notes: Fryar and Hyde (2004) established the genus
Ayria with A. appendiculata as type species, the specimen
was collected from rotting wood submerged in brackish
and sea water, in Brunei. Raja et al. (2009a) introduced the
second species A. nubispora in this genus.
Ayria appendiculata Fryar & K.D. Hyde
Distribution: Distribution: USA, Florida, on submerged
decaying wood (Raja et al. 2009b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8706. The type specimen was
collected from brackish, sea water. Raja et al. (2009b)
reported this species from freshwater habitats but without
description, illustration and information for specimens.
Sequence data is not available.
Ayria nubispora Raja, Ferrer & Shearer
Distribution: Costa Rica, Heredia, La Selva stream, on
submerged decorticated woody debris and Limon, Las
Palmas Stream, on submerged decorticated wood; USA,
Florida, Marion County, Ocala National Forest, Fore
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Fungal Diversity (2019) 99:451–660
Lake,on submerged decorticated woody debris (Raja et al.
2009a).
Asexual morph: Undetermined
Notes: Holotype ILL 40594. Sequence data is not
available.
Cataractispora Hyde et al., Mycol. Res. 103(8): 1019
(1999)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Hyde (1992a) and Hyde et al. (1999).
Type species: Cataractispora aquatica Hyde et al.,
Mycol. Res. 103(8): 1020 (1999)
Notes: The genus Cataractispora was introduced by
Hyde et al. (1999) with three new species and one new
combination and all the species were collected from
freshwater habitats. Ho et al. (2004) introduced one new
species C. receptaculorum which was collected from
freshwater habitats in Hong Kong, China. Five species are
accepted in this genus and all species are reported from
freshwater habitats around the world (Hyde 1992a; Hyde
et al. 1999b; Ho et al. 2004; Raja et al. 2009b).
Cataractispora appendiculata Hyde et al.
Distribution: Australia, north Queensland, Cow Bay,
freshwater stream, on submerged wood; Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Esu,
on submerged wood; Seychelles, Mahe, River St Marie
Louise, on submerged wood (Hyde et al. 1999b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8724. Sequence data is not
available.
Cataractispora aquatica Hyde et al.
Distribution: Australia, north Queensland, Cow Bay,
freshwater stream, on submerged wood; Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Esu,
on submerged wood; Seychelles, Mahe, River St Marie
Louise, on submerged wood (Hyde et al. 1999b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8725. Sequence data is not
available.
Cataractispora bipolaris (K.D. Hyde) Hyde et al.
: Annulatascus bipolaris K.D. Hyde, Aust. Syst. Bot.
5(1): 120 (1992)
Distribution: Australia, north Queensland, Clohesy
River, on submerged wood (Hyde 1992a); USA, Florida,
on submerged decaying wood (Raja et al. 2009b).
Asexual morph: Undetermined
Notes: Holotype BRIP 17374. Sequence data is not
available.
Cataractispora receptaculorum Ho et al.
Distribution: China, Hong Kong, Plover Cove Reservoir, on bamboo submerged in freshwater (Ho et al. 2004)
Asexual morph: Undetermined
Notes: Holotype HKU (M) 5239 (now in IFRD).
Sequence data is not available.
Fungal Diversity (2019) 99:451–660
Cataractispora viscosa Hyde et al.
Distribution: Australia, north Queensland, Cow Bay,
freshwater stream, on submerged wood; China, Hong
Kong, New Territories, Tai Po Kau Country Park, on twigs
submerged in a stream; Mauritius, Black River National
Park, Black River (Hyde et al. 1999b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8726. Sequence data is not
available.
Chaetorostrum Zelski et al., Mycosphere 2(5): 594 (2011)
Asexual morph Taeniolella-like. Sexual morph Description and illustrations see Zelski et al. (2011a)
Type species: Chaetorostrum quincemilense Zelski
et al., Mycosphere 2(5): 595 (2011)
Notes: The genus Chaetorostrum was erected by Zelski
et al. (2011) to accommodate a holomorph species collected from freshwater stream in Peru. There is only one
species in this genus and known only from the type
locality.
Chaetorostrum quincemilense Zelski et al.
Distribution: Peru, Camanti, Stream at Quincemil Trail
1, on submerged woody debris (Zelski et al. 2011).
Asexual morph: Taeniolella-like, see Zelski et al. (2011)
Notes: Holotype ILL 40822. Sequence data is not
available.
Longicollum Zelski et al., Mycosphere 2(5): 540 (2011)
Asexual morph: Undetermined. Sexual morph: Description and illustrations see Zelski et al. (2011).
Type species: Longicollum biappendiculatum Zelski
et al., Mycosphere 2(5): 540 (2011)
Notes: Zelski et al. (2011) introduced the genus Longicollum with single species L. biappendiculatum, which was
collected from both lentic and lotic freshwater habitats in
America.
Longicollum biappendiculatum Zelski et al.
Distribution: Brazil, Bahia, stream at Serra da Jibóia, on
submerged bark debris; Costa Rica, Heredia, La Selva
Biological Station, La Selva stream, on submerged bark
debris; Peru, Camanti, stream at Quincemil Trail 1, on
submerged woody debris; USA, Florida, Wildcat Lake,
Ocala National Forest, on submerged woody debris (Zelski
et al. 2011).
Asexual morph: Undetermined
Notes: Holotype ILL 40794. Sequence data is not
available.
Submersisphaeria K.D. Hyde, Nova Hedwigia 62(1–2):
172 (1996)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Hyde (1996) and Campbell et al.
(2003a, b)
473
Type species: Submersisphaeria aquatica K.D. Hyde,
Nova Hedwigia 62(1–2): 172 (1996)
Notes: The genus Submersisphaeria was introduced by
Hyde (1996) with Submersisphaeria aquatica as type
species. Presently, five species were accepted in this genus
and only one species has been reported from freshwater
habitats. Submersisphaeria aquatica was originally
described from Queensland, Australia (Hyde 1996), and
Campbell et al. (2003a, b) reported it from six states in the
USA. Fournier et al. (2016) found S. aquatica on submerged wood of Pseudotsuga menziesii in France.
Submersisphaeria aquatica K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Hyde 1996); France, Nièvre: Arleuf, Préperny, on
submerged wood of Pseudotsuga menziesii (Fournier et al.
2016); USA, Georgia, Louisiana, Mississippi, New
Hampshire, North Carolina, Tennessee, Wisconsin, on
submerged wood (Campbell et al. 2003a).
Asexual morph: Undetermined
Notes: Holotype BRIP 22743, other specimens collected
from freshwater habitats: ILL 40186, ILL 40260. LSU
sequence data is available.
Vertexicola Hyde et al., Mycologia 92(5): 1019 (2000)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Ranghoo et al. (2000) and Barbosa
et al. (2013).
Type species: Vertexicola caudatus Hyde et al.,
Mycologia 92(5): 1020 (2000)
Notes: The genus Vertexicola is characterized by asci
with a refractive apical ring and a tail-like pedicel and
distoseptate ascospores with relatively thick walls and
lacking appendages or a sheath (Ranghoo et al. 2000).
There are two species accepted in this genus and both were
collected from freshwater habitats.
Vertexicola ascoliberatus Shearer & F.R. Barbosa
Distribution: Costa Rica, Heredia, La Selva Biological
Station, Salto 30, on submergd wood (Barbosa et al. 2013).
Asexual morph: Undetermined
Notes: Holotype ILL 40815. Sequence data is not
available.
Vertexicola caudatus Hyde et al.
Distribution: China, Hong Kong, Tai Po, Plover Cove
Reservoir, submerged wood (Ranghoo et al. 2000).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 3108 (now in IFRD).
Sequence data is not available.
Atractosporales Zhang et al.
Atractosporaceae Zhang et al.
Atractospora Réblová & J. Fourn., Mycol Prog 15: 8
(2016)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Réblová et al. (2016a).
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Fungal Diversity (2019) 99:451–660
Fig. 2 Atractospora aquatica (MFLU 18–2322, holotype) a Ascomata on decaying wood. b Section of the ascoma. c Structure of peridium.
d Paraphyses. e–h Asci. i–l Ascospores. m Germinating ascospore. Scale bars: b 100 lm, c, d 25 lm, e–f 50 lm, i–m 10 lm
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Fungal Diversity (2019) 99:451–660
Type species: Atractospora reticulata Réblová & J.
Fourn., Mycol Prog 15(21): 10 (2016)
Notes: The genus Atractospora was described for
perithecial ascomycetes occurring on decaying wood submerged in freshwater habitats (Réblová et al. 2016b). Five
species are accepted in this genus, of which Atractospora
ellipsoidea and A. thailandensis are reported from Asia and
USA (Ho et al. 1999c; Campbell and Shearer 2004; Fryar
et al. 2004), while other three species were collected from
Europe (Réblová et al. 2016b).
Atractospora aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF555639, Facesoffungi
number: FoF 05411, Fig. 2
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–2322
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 157–251 lm high, 154–234 lm diam., dark
brown to black, solitary, semi-immersed to superficial,
globose to subglobose, unilocular. Ostiole periphysate.
Peridium 20–34 lm thick, consisting two-layers, Outer
layer consisting of thick-walled, mid brown cells of textura
prismatica, towards the interior grading into several layers
of thin-walled subhyaline, flattened cells. Paraphyses
hyaline, unbranched, cylindrical. Asci 110–168 9 9–
13 lm (
x = 139 9 11 lm, n = 20), 8-spored, unitunicate,
cylindrical, with a slender, tapering stipe, apex broadly
rounded, with a conspicuous, non-amyloid apical ring.
Ascospores 15–19 9 7–9 lm (
x = 17 9 8 lm, n = 20),
uniseriate or obliquely uniseriate, fusiform, guttulate,
aseptate, hyaline, smooth-walled.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a
freshwater stream, July 2017, H.W. Shen, S-1398 (MFLU
18–2322, holotype).
Notes: Atractospora aquatica resembles A. thailandensis in having globose to subglobose, unilocular, dark brown
to black ascomata, hyaline, unbranched paraphyses,
cylindrical asci with a slender, tapering stipe and conspicuous, non-amyloid, apical ring and fusiform, hyaline,
aseptate, guttulate ascospores (Zhang et al. 2017a, b).
However, Atractospora aquatica differs from A. thailandensis in having shorter asci (110–168 vs. 175–215 lm)
and smaller ascospores (15–19 vs. 20–26 lm) with smaller
guttules. Atractospora aquatica also resembles A. decumbens and A. reticulata in having solitary, dark brown
ascomata, unitunicate, 8-spored, cylindrical asci and fusiform, hyaline ascospores uniseriate or obliquely uniseriate
in the ascus (Réblová et al. 2016b). However, Atractospora
aquatica differs from A. decumbens and A. reticulata in
having shorter asci and aseptate ascospores, while A.
475
decumbens and A. reticulata have 3-septate ascospores.
Phylogenetic results show that Atractospora aquatica is
distinct from other species of Atractospora (Fig. 1, clade
3).
Atractospora decumbens Réblová & J. Fourn
Distribution: France, Midi-Pyrénées: Ariège, Rimont,
L’ Estanque brook, on submerged wood of Sambucus nigra
in a stream (Réblová et al. 2016b).
Asexual morph: Undetermined
Notes: Holotype PRM 934676. ITS, LSU, SSU and
RPB2 sequence data are available.
Atractospora ellipsoidea (Ho et al.) Réblová & J. Fourn
: Aquaticola ellipsoidea Ho et al., Fungal Divers 3: 90
(1999)
Distribution: Brunei, Tutong River, on submerged wood
(Fryar et al. 2004); China, Hong Kong, Tai Po, Tai Po Kau
Forest Stream, on submerged wood of Machilus velutina
(Ho et al. 1999c); USA, North Carolina (Campbell and
Shearer 2004).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 6033 (now in IFRD). LSU
sequence data is available.
Atractospora reticulata Réblová & J. Fourn
Distribution: France, Midi-Pyrénées: Ariège, Rimont,
Peyrau brook, on submerged wood of Sambucus nigra in a
stream (Réblová et al. 2016b).
Asexual morph: Undetermined
Notes: Holotype PRM 934677. ITS, LSU and RPB2
sequence data are available.
Atractospora thailandensis Dong et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged wood in a small River (Zhang et al. 2017).
Asexual morph: Undetermined
Notes: Holotype HKAS 96226. ITS, LSU, SSU, RPB2
and TEF1a sequence data are available.
Atractospora verruculosa Réblová & J. Fourn
Distribution: France, Pyrénées-Atlantiques, Lescun,
Borde Bayé, Lescun stream, on submerged wood of Alnus
glutinosa (Réblová et al. 2016b).
Asexual morph: Undetermined
Notes: Holotype PRM 934679. ITS, LSU, SSU and
RPB2 sequence data are available.
Rubellisphaeria Réblová & J. Fourn., Mycol Prog 15: 13
(2016)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Réblová et al. (2016a).
Type species: Rubellisphaeria abscondita Réblová &
J.Fourn., Mycol Prog 15: 13 (2016)
Notes: Réblová et al. (2016a) introduced the genus
Rubellisphaeria with single species Rubellisphaeria
abscondita, collected from freshwater in France. This is a
monotypic genus and known only from the type locality.
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476
Rubellisphaeria abscondita Réblová & J. Fourn
Distribution: France, Loire: Saint Jean la Vêtre, Les
Yverneaux, on submerged twigs of Abies alba in a peat bog
(Réblová et al. 2016b).
Asexual morph: Undetermined
Notes: Holotype PRM 934681. ITS, LSU, SSU and
RPB2 sequence data are available.
Jobellisiales M.J. D’souza & K.D. Hyde
Jobellisiaceae Réblová
Jobellisia M.E. Barr, Mycotaxon 46: 60 (1993)
Asexual morph Undetermined. Sexual morph Ascomata
superficial or basally immersed, nonstromatic, globose to
subglobose to obpyriform to lageniform, brown or externally with yellowish pigments, glabrous or slightly rugose,
with short to long papilla or with long upright neck.
Peridium comprising two or three layers. Paraphyses
numerous, septate, hyaline. Asci 8-spored, unitunicate,
cylindrical-clavate, with nonamyloid apex and distinct
refractive apical annulus, with basally swollen stipe, stipe
remnants attached to the ascogenous hyphae after dehiscence. Ascospores ellipsoidal to reniform to navicular,
aseptate or transversely 1-septate with one or two polar
germ pores, brown.
Type species: Jobellisia luteola (Ellis & Everh.) M.E.
Barr, Mycotaxon 46: 61 (1993)
Notes: The genus Jobellisia currently comprises nine
species and they are found in tropical and temperate zones
2008;
of
the
northern
hemisphere
(Réblová
Maharachchikumbura et al. 2016). Three species have been
found in freshwater habitats.
Jobellisia guangdongensis F. Liu & L. Cai
Distribution: China, Guangdong Province, on submerged decaying wood (Liu et al. 2011).
Asexual morph: Undetermined
Notes: Holotype HMAS 251240. ITS and LSU sequence
data are available.
Jobellisia luteola (Ellis & Everh.) M.E. Barr
: Letendraea luteola Ellis & Everh., Proc. Acad. nat.
Sci. Philad. 47: 415 (1895)
Distribution: USA, on submerged decaying wood (Raja
et al. 2009b).
Asexual morph: Undetermined
Notes: Holotype Morgan 1109. LSU sequence data is
available. Jobellisia luteola was originally collected from
decayed wood on terrestrial habitats. Raja et al. (2009b)
reported this species from freshwater habitats during their
studies on latitudinal, habitat and substrate distribution
patterns of freshwater ascomycetes in Florida, USA.
Jobellisia viridifusca K.M. Tsui & K.D. Hyde
Distribution: China, Hong Kong, Lantau Island, Tung
Chung River, on submerged decaying wood (Ranghoo
et al. 2001)
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Asexual morph: Undetermined
Notes: Holotype HKU (M) 80455 (now in IFRD).
Sequence data is unavailable.
Diaporthales Nannf.
Diaporthaceae Hohn. ex Wehm.
Diaporthe Nitschke, Pyrenomycetes Germanici 2: 240
(1870)
Asexual morph Coelomycetous, pycnidial, ostiolate,
brown to black, scattered or aggregated, globose to subglobose. Conidiophores cylindrical, sometimes filiform,
aseptate or septate, cylindrical, sometimes branched.
Conidiogenous cells phialidic, cylindrical, terminal. Conidia dimorphic, hyaline, smooth, with usually fusiform and
biguttulate alpha conidia and usually filiform, hamate, nonguttulate beta conidia. Sexual morph Ascomata globose to
subglobose, coriaceous, immersed to semi-immersed, single to clustered, brown to black. Neck cylindrical, black.
Peridium comprising compressed cells of textura angularis. Paraphyses cylindrical, longer than asci, septate. Asci
8-spored, unitunicate, thin-walled, apedicellate, broad
cylindrical to obclavate, with a minute apical ring. Ascospores overlapping biseriate, ellipsoidal to fusiform,
septate, hyaline, thin-walled, smooth-walled.
Type species: Diaporthe eres Nitschke, Pyrenomyc.
Germ. 2: 245 (1870)
Notes: Species of Diaporthe have broad host ranges and
are widely distributed (Udayanga et al. 2012). Currently,
MycoBank (accessed December 2018) lists more than 900
names in the genus Diaporthe, but this was reduced to 171
species (Dissanayake et al. 2017). Hu et al. (2012c) introduced a new Diaporthe species D. aquatica, collected from
submerged wood in stream in Yunnan Province, China.
Diaporthe aquatica Hu et al.
Distribution: China, Guizhou Province, Guiyang city,
on submerged wood in a small ditch (Hu et al. 2012c)
Asexual morph: undetermined
Notes: Holotype IFRD 021–018; ITS sequence data are
available, ex-type strain IFRDCC 3051. D. aquatica is the
only Diaporthe species restricted to freshwater habitats (Hu
et al. 2012c).
Gnomoniaceae G. Winter
Ambarignomonia Sogonov, Stud. Mycol. 62: 35 (2008)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Senanayake et al. (2018).
Type species: Ambarignomonia petiolorum (Schwein.)
Sogonov, Stud Mycol 62: 36 (2008)
Notes: The genus Ambarignomonia was introduced by
Sogonov et al. (2008) to accommodate Sphaeria petiolorum. Presently, there is only one species accepted in this
genus. Sogonov et al. (2008) mentioned the monotypic
Ambarignomonia with A. petiolorum and is restricted to
one plant host, Liquidambar styraciflua, whereas other
Fungal Diversity (2019) 99:451–660
genera of the Gnomoniaceae do not show such consistency
in host associations.
Ambarignomonia petiolorum (Schwein.: Fr.) Sogonov
: Sphaeria petiolorum Schwein.: Fr., Schr. Naturf.
Ges. Leipzig 1: 41. 1822: Syst. Mycol. 2: 517 (1823)
: Gnomonia petiolorum (Schwein.: Fr.) Cooke, Grevillea 7: 54 (1878)
: Gnomoniella amoena var. petiolorum (Schwein.: Fr.)
Sacc., Syll. Fung. 1: 414 (1882)
Distribution: USA, Wisconsin, Trout lake/Big Muskellunge lake, on submerged partially decomposed Acer
rubrum leaves (Fallah and Shearer 2001).
Asexual morph: Undetermined
Notes: Lectotype BPI 800519, Epitype BPI 844274,
specimens collected from freshwater habitats: ILLS 54015,
ILLS 54016. ITS, LSU, RPB2, TEF1a and b-tubulin
sequence data are available. Fallah and Shearer (2001)
collected two specimens (ILLS 54015 and ILLS 54016)
from freshwater habitats and provided descriptions and
illustrations and identified the collections as Gnomonia
petiolorum. Sogonov et al. (2008) combined this species as
Ambarignomonia petiolorum.
Gnomonia Ces. & De Not., Comm. Soc. crittog. Ital. 1(4):
231 (1863)
Asexual morph see Sivanesan and Shaw (1977). Sexual
morph Description see Maharachchikumbura et al. (2016).
Type species: Gnomonia vulgaris Ces. & De Not.,
Comm. Soc. crittog. Ital. 1(fasc. 4): 232 (1863)
Notes: The genus Gnomonia was introduced by Cesati
and De Notaris (1863) and typified by Gnomonia gnomon.
Gnomonia comprises 273 species and only one species has
been reported from freshwater habitats (Sivanesan and
Shaw 1977; Fallah and Shearer 2001; Senanayake et al.
2018).
Gnomonia papuana Sivan. & D.E. Shaw
Distribution: Papua New Guinea, on submerged leaves
(Sivanesan and Shaw 1977).
Asexual morph: Sesquicillium-like, see Sivanesan and
Shaw (1977)
Notes: Holotype IMI 197503. Sequence data is not
available.
Gnomoniella Sacc., Michelia 2 (7): 312 (1881)
Asexual morph Undetermined. Sexual morph Ascomata
globose to subglobose, immersed. Paraphyses deliquescence. Asci cylindrical, subsessiles. Ascospores fusiform,
ellipse, hyaline, septate.
Type species: Gnomoniella tubaeformis (Tode) Sacc.,
Syll. fung. (Abellini) 1: 413 (1882)
Notes: Kirk et al. (2008) accepted 13 species in the
genus Gnomoniella. Two species have been found in
freshwater habitats and both were collected from Asia (Ho
et al. 2001).
477
Gnomoniella microspora M. Monod
Distribution: Malaysia, on submerged wood (Ho et al.
2001).
Asexual morph: Undetermined
Notes: ITS sequence data are available. Gnomoniella
microspora was originally collected from terrestrial habitats (Monod 1983). Ho et al. (2001) reported Gnomoniella
microspora from freshwater habitats and we consider this
species as freshwater fungus.
Gnomoniella rubicola Pass.
Distribution: Brunei, on submerged wood (Ho et al.
2001).
Asexual morph: Undetermined
Notes: Sequence data is not available. Same as Gnomoniella microspora, the original collection of G. rubicola
was not collected from freshwater habitats. However, we
consider this species as freshwater fungus as Ho et al.
(2001) found it on submerged wood in Brunei.
Diaporthales genera incertae sedis
Phruensis Pinruan, Mycologia 96 (5): 1165 (2004)
Asexual morph Hyphomycetous. Colonies on PDA effuse,
brown mycelium party immersed, nonstromatic. Conidiophores semimacronematous, mononematous, branched,
straight or slightly flexuous, pale brown to brown, smooth.
Conidiogenous cells monophialidic, determinate, with
small collarettes. Conidia straight or curved, oblong, hyaline, smooth. Sexual morph Ascomata immersed, subglobose, black, coriaceous, ostiolate, with long central
cylindrical neck. Peridium composed of 2 layers, with
outer layer parenchymatous, intensely brown and merging
with the host cells, with inner layer, cells elongate and
hyaline. Paraphyses hyaline, broad, septate. Asci unitunicate, cylindrical to fusiform, apedicellate, apically rounded,
J-, subapical ring. Ascospores cylindrical, straight or
curved, versicolorous, transseptate, brown with hyaline or
pale brown end cells.
Type species: Phruensis brunneispora Pinruan,
Mycologia 96(5): 1165 (2004)
Notes: Pinruan et al. (2004b) introduced the genus
Phruensis with a single species P. brunneispora, which
was collected from a submerged palm in Sirindhorn Peat
Swamp Forest in Thailand. No more species reported for
this genus since it was introduced.
Phruensis brunneispora Pinruan
Distribution: Thailand, Narathiwat Province, Sirindhorn peat swamp forest, on submerged palm in freshwater
stream (Pinruan et al. 2004b).
Asexual morph: Phialophora-like, see Pinruan et al.
(2004b)
Notes: Holotype BBH, Pinruan Wah 113.1. SSU
sequence data is available.
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Fig. 3 Distoseptispora appendiculata (DLU B95, holotype) a Colonies on wood. b–e Conidiophores with conidia. f Conidiogenous cells with
conidia. g Conidiogenous cells. h–k Conidia. l Germinating conidium. Scale bars: b–f 30 lm, g–l 20 lm
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479
Fig. 4 Distoseptispora guttulata (B-43) a Colonies on wood. b–d Conidiophores with conidia. e–g Conidia. h Germinating conidium. Culture on
PDA from above (i) and reverse (j). Scale bars: b–h 30 lm
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480
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Fig. 5 Distoseptispora lignicola (MFLU 18–1458, holotype). a, b Colonies on wood. c–d Conidiophores with conidia. e–f Conidiogenous cells
and conidia. g–j Conidia. k Germinating conidium. Culture on PDA from surface (l) and reverse (m). Scale bars: d 30 lm, c, e–k 20 lm
123
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481
Fig. 6 Distoseptispora multiseptata (B-37) a, b Colonies on wood. c–f Conidiophores with conidia. g–k Conidia. l Conidiophores with
conidiogenous cells. m Germinating conidium. Culture on PDA from above (n) and reverse (o). Scale bars: c–k, m 30 lm, l 10 lm
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482
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Fig. 7 Distoseptispora neorostrata (DLU B103, holotype) a, b Colonies on wood. c, d Conidiophores with conidia. e, f Conidiogenous cells with
conidia. g–j Conidia. k Germinating conidium. Culture on MEA from above (l) and reverse (m). Scale bars: c–k 30 lm
Distoseptisporales Z.L. Luo, H.Y. Su & K.D. Hyde, ord.
nov
Index Fungorum number: IF 555640, Facesoffungi
number: FoF 05412
Asexual morph Mycelium mostly immersed, composed of
branched, septate, smooth, pale brown hyphae. Conidiophores macronematous, mononematous, septate, unbranched, erect, straight or flexuous, smooth. Conidiogenous
cells monoblastic, integrated, determinate, terminal,
cylindrical. Conidia acrogenous, solitary, euseptate or
123
distoseptate, obclavate or cylindrical with rounded apex.
Conidial
secession
schizolytic.
Sexual
morph
Undetermined.
Type family: Distoseptisporaceae K.D. Hyde &
McKenzie, Fungal Divers 80:402 (2016)
Notes: Distoseptisporaceae was established by Su et al.
(2016) with a single genus Distoseptispora based on morphology and phylogeny. Phylogenetic analysis based on
combined LSU, SSU, RPB2 and TEF1a sequence data here
shows
that
species
of
Distoseptispora
and
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483
Fig. 8 Distoseptispora obclavata (DLU B71, holotype) a, b Colonies on wood. c–g Conidiophores with conidia. h, i Conidiogenous cells with
conidia. j–m Conidia. n Germinating conidium. Culture on PDA from above (o) and reverse (p). Scale bars: c–h 30 lm, i–n 20 lm
123
484
Aquapteridospora cluster together with strong support and
form a distinct clade within subclass Diaporthomycetidae.
We therefore raise Distoseptisporaceae to order
Distoseptisporales.
Distoseptisporaceae K.D. Hyde & McKenzie
Distoseptispora Hyde et al., Fungal Divers 80: 402 (2016)
Asexual morph Description and illustration see Su et al.
(2016) and Yang et al. (2018). Sexual morph
Undetermined.
Type species: Distoseptispora aquatica Luo et al.,
Fungal Divers 80: 402 (2016)
Notes: Su et al. (2016) introduced the genus Distoseptispora to accommodate two Sporidesmium-like species.
Yang et al. (2017) emended the description of the genus
Distoseptispora. Currently, there are 13 species in Distoseptispora with ten species collected from freshwater
habitats (Su et al. 2016; Hyde et al. 2016b; Luo et al.
2018a; Yang et al. 2018a).
Distoseptispora appendiculata D.F. Bao, Z.L. Luo & H.Y.
Su, sp. nov.
Index Fungorum number: IF 556690, Facesoffungi
number: FoF 06302, Fig. 3
Etymology: Referring to its gelatinous conidia
appendage
Holotype: DLU B95
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, olivaceous or
mid-brown, hairy, velvety. Mycelium mostly immersed,
consisting of branched, septate, smooth, subhyaline to pale
brown hyphae. Conidiophores 62–86 lm long, 4.5–5.5 lm
wide (
x = 74 9 5 lm, n = 10), macronematous, mononematous, solitary, erect, straight or flexuous, olivaceous or
brown, 5–6-septate, smooth. Conidiogenous cells
monoblastic, holoblastic, terminal, dark brown. Conidia
67–89 lm long, 10–16 lm wide (
x = 78 9 13 lm,
n = 20), acrogenous, solitary, obpyriform or obclavate,
13–17-distoseptate, thick-walled, olivaceous or dark brown
below, hyaline towards apex, truncate at base, slender and
rounded at apex, smooth, with a conspicuous, gelatinous,
hyaline sheath around tip. Sexual morph Undetermined.
Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon,
saprobic on decaying wood submerged in a freshwater
stream, 1st October 2017, Z.L. Luo, B-95 (DLU B95,
holotype), ex-type living culture MFLUCC 18–0259.
Notes: Distoseptispora appendiculata resembles D.
obpyriformis in having effuse, olivaceous or mid-brown,
hairy, velvety colonies, macronematous, straight or flexuous, olivaceous or brown, septate conidiophores and
obpyriform, distoseptate, olivaceous or dark brown conidia
(Luo et al. 2018a). However, Distoseptispora appendiculata is easily distinguished from D. obpyriformis and other
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Fig. 9 Aquapteridospora fusiformis (MFLU 18–1601, holotype). c
a Colonies on substrate. b–d Conidiophore with conidia. e, f
Conidiogenous cells with conidia g–j Conidia, k Germinating
conidium. Culture on PDA from surface (l) and reverse (m). Scale
bars: b–d 50 lm, e, f 20 lm, g–k 10 lm
species in Distoseptispora by its conspicuous, gelatinous,
hyaline appendage sheath. Phylogenetic results show that
Distoseptispora appendiculata is distinct from other species of Distoseptispora (Fig. 10).
Distoseptispora aquatica Luo et al.
Distribution: China, Yunnan Province, Dali, Cangshan
Mountain, on submerged wood in stream (Su et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKAS 83991. ITS and LSU sequence
data are available.
Distoseptispora cangshanensis Luo et al.
Distribution: China, Yunnan Province, Dali, Cangshan
Mountain, on submerged wood (Luo et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0474. ITS, LSU and TEF1a
sequence data are available.
Distoseptispora fluminicola McKenzie et al.
Distribution: China, Yunnan Province, Dali, on submerged wood (Su et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKAS 84006. ITS, LSU and TEF1a
sequence data are available.
Distoseptispora guttulata J. Yang & K.D. Hyde
Facesoffungi number: FoF 03357, Fig. 4
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, brown to black,
hairy or velvety. Mycelium partly superficial, partly
immersed, consisting of branched, septate, smooth, subhyaline to pale brown hyphae. Conidiophores 28–84 lm
long, 4–5 lm wide (
x = 56 9 4.5 lm, n = 20), macronematous, mononematous, mid or dark brown, 3–4(–7)-septate, solitary or caespitose, smooth, straight or slightly
flexuous, cylindrical, rounded at the apex. Conidiogenous
cells monoblastic, integrated, terminal, determinate, mid to
dark brown, cylindrical, sometimes proliferating percurrently. Conidia 70–190(–520) lm long, 8.5–10.5 lm wide
(
x = 130 9 9.5 lm, n = 20), acrogenous, solitary,
holoblastic, obclavate or lanceolate, rostrate, curved,
9–14(–27)-euseptate, mid to dark brown, or olivaceous,
smooth, truncate at the base, tapering to the apex. Sexual
morph Undetermined.
Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon,
saprobic on decaying wood submerged in a freshwater
stream, 1st October 2017, Z.L. Luo, B-43, living culture
DLUCC B43.
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123
486
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b Fig. 10 Phylogram generated from maximum likelihood analysis
based on ITS, LSU, RPB2 and TEF1a sequence data for species of
Distoseptisporales (with Sordaria fimicola as outgroup). The best
scoring RAxML tree with a final likelihood value of - 30852.86243
is presented. RAxML bootstrap support values equal to or greater
than 75% are given before the forward slash (black). Bayesian
posterior probability equal to or higher than 0.95 are given after the
forward slash (red). Hyphen (‘-’) indicates a value lower than 75% for
RAxML and Bayesian posterior probability lower than 0.95. Newly
generated sequences are in red. Ex-type or ex-epitype strains are in
bold
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Yang
et al. 2018a)
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0852, isotype GZAAS 17–
0005. ITS, LSU, SSU and TEF1a sequence data are
available. Distoseptispora guttulata was introduced by
Yang et al. (2018a) based on a collection obtained from a
freshwater stream in Thailand. Morphologically, our isolate fits well with the characters of D. guttulata (Yang et al.
2018a). Phylogenetic analysis also shows that our isolate
clusters with ex-type of D. guttulata with good support
(Fig. 10).
Distoseptispora lignicola D.F. Bao, Z.L. Luo, H.Y. Su &
K.D. Hyde, sp. nov.
Index Fungorum number: IF 555641, Facesoffungi
number: FoF 05413, Fig. 5
Etymology: Referring to this taxon dwelling on wood
Holotype: MFLU 18–1458
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, scattered, hairy,
pale brown to brown. Mycelium mostly immersed, composed of branched, septate, brown, smooth hyphae. Conidiophores
84–124 lm
long,
4–5 lm
wide
(
x = 104 9 4.5 lm, n = 20), macronematous, mononematous, solitary or in groups, erect, straight or slightly flexuous, 6–10-septate, unbranched, cylindrical, brown,
smooth. Conidiogenous cells monoblastic, integrated, terminal, determinate, brown, cylindrical. Conidia 60–
108 lm long, 7–9 lm wide (
x = 84 9 8 lm, n = 20),
acrogenous, solitary or catenate, obclavate, truncate at
base, tapering towards the apex, straight or slightly curved,
5–9-euseptate, slightly constricted at septa, guttulate,
brown, smooth. Sexual morph Undetermined.
Material examined: THAILAND, SaiKhu Waterfall, on
submerged decaying wood, 28 August 2017, C.G. Lin, B-2
(MFLU 18–1458, holotype), ex-type living culture
MFLUCC 18–0198.
Notes: Distoseptispora lignicola resembles D. guttulata
in having macronematous, mononematous, unbranched,
cylindrical, septate conidiophores, solitary or in groups on
natural substrata, monoblastic, integrated, terminal, determinate conidiogenous cells and acrogenous, obclavate,
487
rostrate, euseptate, guttulate conidia (Yang et al. 2018a).
However D. lignicola differs from D. guttulata in having
longer conidiophores (84–124 vs. 55–90 (–145) lm) and
5–9-euseptate conidia, while D. guttulata has 11–14(–20)euseptate conidia. Phylogenetically, Distoseptispora guttulata clusters in Distoseptispora and sister to D. leonensis
(HKUCC 10822), but is distinct from other Distoseptispora
species (Fig. 1, clade 15).
Distoseptispora multiseptata J. Yang & K.D. Hyde
Facesoffungi number: FoF 02244, Fig. 6
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, dark olivegreen, hairy or velvety. Mycelium mostly immersed,
comprised of branched, septate, smooth, hyaline to pale
brown hyphae. Conidiophores 29–47 lm long, 4–6 lm
wide (
x = 38 9 5 lm, n = 20), macronematous, mononematous, solitary, brown, 2–3-septate, straight or slightly
flexuous, erect, slightly tapering distally, truncate at the
apex, olive-green to dark brown. Conidiogenous cells
holoblastic, monoblastic, integrated, terminal, brown,
determinate, cylindrical. Conidia 147–185 lm long, 12–
14 lm wide (
x = 16 9 13 lm, n = 20), acrogenous, solitary, obclavate, rostrate, multi-distoseptate, tapering
towards the apex, dark olivaceous green. Conidial secession schizolytic. Sexual morph Undetermined.
Material examined: THAILAND, Sai khu waterfall,
Prachuap khiri Khan., saprobic on decaying wood submerged in a freshwater stream, 1st September 2017, Vinit
Kumar, B-37, living culture MFLUCC 18–0215.
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, on submerged wood in a stream (Hyde et al.
2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1144, isotype HKAS 95045.
ITS, LSU, SSU, RPB2 and TEF1a sequence data are
available. Distoseptispora multiseptata was introduced by
Yang et al. (2018a) based on a collection obtained from a
freshwater stream in Thailand. Morphologically, our isolate fits well with the characters of D. multiseptata (Yang
et al. 2018a). Phylogenetic analysis also shows that our
isolate clusters with ex-type of D. multiseptata (Fig. 10).
Distoseptispora neorostrata D.F. Bao, Z.L. Luo & H.Y.
Su, sp. nov.
Index Fungorum number: IF 556691, Facesoffungi
number: FoF 06135, Fig. 7
Etymology: Referring to its morphological similarity to
D. rostrata
Holotype: DLU B103
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, dark olivaceous,
hairy. Mycelium partly superficial, partly immersed in the
substrate, comprised of branched, septate, hyaline to pale
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b Fig. 11 Ceratosphaeria aquatica (MFLU 18–2323, holotype) a Ap-
pearance of necks on substrate. b Section through ascomata. c,
d Section through neck. e Structure of peridium. f Paraphyses. g–
j Asci. k Apical ring. l–o Ascospores. Scale bars: b, c 100 lm,
d 50 lm, e–j, l–o 20 lm, k 10 lm
brown hyphae. Conidiophores 93–117 lm long, 5.5–
6.5 lm wide (
x = 105 9 6 lm, n = 10), macronematous,
mononematous, solitary, brown, 5–7-septate, straight or
flexuous, tapering distally, truncate at the apex. Conidiogenous cells monoblastic, integrated, terminal, brown,
determinate. Conidia 109–147 lm long, 13–15 lm wide
(
x = 128 9 14 lm, n = 20), acrogenous, solitary, elongate,
obclavate, rostrate, multi-distoseptate, tapering towards the
rounded apex, dark olivaceous to mid or dark brown, pale
brown towards apex, truncate at the base, guttulate,
smooth-walled. Sexual morph Undetermined.
Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon,
saprobic on decaying wood submerged in a freshwater
stream, 1st October 2017, Z.L. Luo, B-103 (DLU B103,
holotype), ex-type living culture MFLUCC 18–0376.
Notes: Distoseptispora neorostrata shares similar morphological characters with D. rostrata in the shape, colour
and size of its conidiophores and conidia (Luo et al.
2018a). However, the multi-gene phylogenetic analyses
showed that they are different species (Fig. 10).
Distoseptispora obclavata D.F. Bao, Z.L. Luo & H.Y. Su,
sp. nov.
Index Fungorum number: IF 556689, Facesoffungi
number: FoF 06296, Fig. 8
Etymology: Referring to its obclavate conidia
Holotype: DLU B71
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, olivaceous or
dark brown, hairy, velvety. Mycelium mostly immersed,
consisting of branched, septate, smooth, subhyaline to pale
brown hyphae. Conidiophores 117.5–162.5 lm long, 5–
7 lm wide (
x = 140 9 6 lm, n = 20), macronematous,
mononematous, pale to dark brown, solitary, 5–10-septate,
erect, straight or slightly flexuous, unbranched, smooth,
cylindrical, rounded at the apex. Conidiogenous cells
monoblastic, integrated, terminal, determinate, pale to dark
brown, cylindrical. Conidia 46–66 lm long, 9–11 lm wide
(
x = 56 9 10 lm, n = 25), acrogenous, solitary, obclavate,
9–11-distoseptate, thick-walled, olivaceous to pale or dark
brown, apering towards the rounded apex, slightly curved,
truncate at the base, guttulate, smooth-walled. Sexual
morph Undetermined.
Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon,
saprobic on decaying wood submerged in a freshwater
489
stream, 1st October 2017, Z.L. Luo, B-71 (DLU B71,
holotype), ex-type living culture MFLUCC 18–0329.
Notes: Distoseptispora obclavata resembles D. appendiculata in having effuse, olivaceous or mid-brown, hairy,
velvety colonies, macronematous, straight or flexuous,
septate conidiophores and obclavate, distoseptate, olivaceous or dark brown conidia. However, Distoseptispora
obclavata is easily distinguished from D. appendiculata by
its shorter conidia (46–66 vs. 67–89 lm) without appendage sheath. Phylogenetic results show that Distoseptispora appendiculata is distinct from other species of
Distoseptispora (Fig. 10).
Distoseptispora obpyriformis Z.L. Luo & H.Y. Su
Distribution: China, Yunnan Province, Nujiang River,
on submerged wood (Luo et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0476, paratype MFLU 18–
0477. ITS, LSU, RPB2 and TEF1a sequence data are
available.
Distoseptispora phangngaensis Yang et al.
Distribution: Thailand, Phang Nga Province, Bann Tom
Thong Khang, on decaying wood submerged in a freshwater stream (Yang et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0855, isotype GZAAS 17–
0008. ITS, LSU and TEF1a sequence data are available.
Distoseptispora rostrata Luo et al.
Distribution: China, Yunnan Province, Nujiang River,
on submerged wood (Luo et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0479, paratype MFLU
18–0475. ITS, LSU, RPB2 and TEF1a sequence data are
available.
Distoseptispora submersa Luo et al.
Distribution: China, Yunnan Province, Nujiang River,
on submerged wood (Luo et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0478, isotype HKAS 92806.
ITS, LSU, RPB2 and TEF1a sequence data are available.
Distoseptispora suoluoensis Yang et al.
Distribution: China, Guizhou Province, Anshun city,
Gaodang village, on decaying wood submerged in Suoluo
River (Yang et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0853, isotype GZAAS 17–
0006, paratype MFLU 17–0854. ITS, LSU and TEF1a
sequence data are available.
Distoseptisporales genera incertae sedis
Aquapteridospora Yang et al., Cryptog. Mycol. 36(4): 474
(2015)
Asexual morph Descriptions and illustration see Yang
et al. (2015). Sexual morph Undetermined.
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490
Type species: Aquapteridospora lignicola J. Yang, K.D.
Hyde & Maharachch, Cryptog Mycol 36: 474 (2015)
Notes: Yang et al. (2015) introduced the genus
Aquapteridospora with single asexual species, A lignicola,
which was collected from freshwater stream in northern
Thailand. In this study, we introduce the second species
collected from freshwater. Aquapteridospora was placed as
Diaporthomycetidae genera incertae sedis by Yang et al.
(2015). In our phylogenetic analysis, Aquapteridospora
species form a distinct clade within Distoseptisporales and
basal to Distoseptisporaceae, and we therefore treat this
genus as Distoseptisporales genera incertae sedis, and its
familial placement needs further studies.
Aquapteridospora lignicola Yang et al.
Distribution: Thailand, Chiang Rai Province, stream
flowing in Tham Luang Nang Non Cave, on submerged
wood (Yang et al. 2015).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1172. LSU sequence data is
available.
Aquapteridospora fusiformis Z.L. Luo, D.F. Bao, H.Y. Su
& K.D. Hyde, sp. nov.
Index Fungorum number: IF 555642, Facesoffungi
number: FoF 05414, Fig. 9
Etymology: Referring to the fusiform conidia of this
fungus.
Holotype: MFLU 18–1601
Saprobic on decaying wood submerged in freshwater.
Asexual morph Colonies on the natural substrate effuse,
hairy, pale brown to brown. Mycelium superficial or partly
immersed, composed of branched, septate, pale brown to
brown, smooth, thin-walled hyphae. Conidiophores (88–
)134–188 lm long, 5–7 lm wide (
x = 161 9 6 lm,
n = 20), macronematous, mononematous, solitary, erect,
straight or slightly flexuous, unbranched, cylindrical, septate, smooth, thick-walled, brown at the base, paler towards
apex. Conidiogenous cells polyblastic, terminal, later
becoming intercalary, pale brown, integrated, with several
sympodial proliferations, bearing tiny, protuberant, circular
scars. Conidia 14–18 lm long, 5–7 lm wide
(
x = 16 9 6 lm, n = 20), solitary, fusiform, obtuse at both
ends, mostly 3-septate, sometimes 4-septate, slightly constricted at septa, brown to dark brown in central cells and
subhyaline at end cells, smooth. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Jizu
Mountain, saprobic on decaying wood submerged in a
freshwater stream, July 2016, S.M. Tang, S-889 (MFLU
18–1601, holotype), ex-type living culture MFLUCC
18–1606.
Notes: Aquapteridospora fusiformis resembles A. lignicola in having macronematous, solitary, unbranched, septate conidiophores which are brown at the base and paler
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towards apex, polyblastic, terminal conidiogenous cells
and fusiform, septate conidia. Both of these species also
share similar size of conidiophores and conidia (Yang et al.
2015). However, A. fusiformis differs from A. lignicola in
having pale brown to brown colonies, smooth conidia
without a sheath, while the conidia of A. lignicola has large
guttules in the middle cells and a conspicuous sheath.
Phylogenetic analysis also shows that A. fusiformis and A.
lignicola are distinct from other species, but they cluster
together with strong support (Fig. 1, clade 15). To further
support A. fusiformis as a new species, we compared
nucleotide differences with A. lignicola (MFLU 15–1172)
following the guidelines of Jeewon and Hyde (2016).
Comparison of the 789 nucleotides across the LSU region
reveals 9 bp differences. Based on the differences of
morphology and DNA nucleotide, we introduce our isolate
as new species in Aquapteridospora.
Magnaporthales Thongk et al.
Ceratosphaeriaceae Z.L. Luo, H.Y. Su & K.D. Hyde,
fam. nov.
Index Fungorum: IF 555643; Facesoffungi number: FoF
05415
Asexual morph Harpophora-like. Phialides or short
conidiophores arising on aerial hyphae, with conidial heads
slimy, inconspicuous, and transparent. Conidiogenous cells
phialidic, ampulliform to lageniform, terminal or intercalary, cylindrical. Conidia cylindrical, hyaline, aseptate,
smooth. Sexual morph Stromata absent. Ascomata globose
to pyriform, deeply immersed to almost superficial, dark
brown to black, carbonaceous, with a long cylindrical,
black or yellow crystals neck. Periphyses well-developed.
Peridium composed of a large number of layers of very
thick-walled rather small cells in the neck region. Interascal tissue of paraphyses thin-walled, probably evanescent
at maturity. Asci 8-spored, unitunicate, cylindrical, fairly
thin-walled, the apex truncate, with a conspicuous J-apical
ring. Ascospores arranged biseriately, narrowly cylindricfusiform, or filiform, the ends acute, thin-walled, hyaline,
septate, guttulate, smooth-walled.
Type genus: Ceratosphaeria Niessl, Verh. nat. Ver.
Brünn 14: 203 (1876)
Notes: Réblová (2006) accommodated Ceratosphaeria
in Magnaporthaceae based on their phylogenetic analyses
of combined LSU and SSU sequence data. Phylogenetically, our multi-gene analyses show that Ceratosphaeria
species formed a distinct clade in Magnaporthales and
shares close relationship with Pseudohalonectriaceae
(Fig. 15). Morphologically, Pseudohalonectriaceae is
characterized by erumpent to immersed ascomata with a
protruding, greenish yellow, bright yellow to brown neck,
cylindrical to clavate asci with a nonamyloid, thimbleshaped, refractive, apical apparatus and cylindrical or
Fungal Diversity (2019) 99:451–660
Fig. 12 Ceratosphaeria lignicola (MFLU 18–1457, holotype). a,
b Appearance of neck on substrate. c, d Section of ascoma. e Structure
of peridium. f Paraphyses. g–k Asci. l–n Ascospores. o Germinating
491
ascospore. culture on PDA from surface (p) and reverse (q). Scale
bars: c 100 lm, d, g 50 lm, h–o 30 lm, e, f 20 lm
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b Fig. 13 Aquafiliformis lignicola (MFLU 18–2325, holotype) a Ap-
pearance of necks on substrate. b Section through ascoma. c Structure
of peridium. d Paraphyses. e, f Asci. g–j Ascospores. Culture on PDA
from surface (k) and reverse (l). Scale bars: b 150 lm, c 50 lm, d–
f 30 lm, g–j 20 lm
ellipsoidal, straight to curved, multi-septate ascospores
(Hongsanan et al. 2017). Ceratosphaeriaceae is distinct
from Pseudohalonectriaceae in having narrowly cylindricfusiform to filiform, longer ascospores. We therefore
introduce a new family Ceratosphaeriaceae to accommodate Ceratosphaeria.
Ceratosphaeria Niessl, Verh. nat. Ver. Brünn 14: 203
(1876)
Asexual morph Harpophora-like. Phialides or short
conidiophores arising on aerial hyphae, conidial heads
slimy, inconspicuous, and transparent. Conidiogenous cells
phialidic, ampulliform to lageniform, terminal or intercalary, hyaline to subhyaline, cylindrical. Conidia cylindrical with curvature, hyaline, narrowly rounded at both
ends, aseptate, smooth. Sexual morph Stromata absent.
Ascomata globose to pyriform, often irregular in shape due
to compression, deeply immersed to almost superficial,
dark brown to black, carbonaceous, with a long cylindrical
black or yellow crystals, thick-walled neck which is easily
detached, scattered to densely aggregated. Peridium composed of a large number of layers of very thick-walled
rather small cells in the neck region. Interascal tissue of
paraphyses thin-walled, probably evanescent at maturity,
periphyses well-developed. Asci 8-spored, unitunicate,
cylindrical, fairly thin-walled, the truncate apex, with a
conspicuous, J-, apical ring. Ascospores arranged biseriately, narrowly cylindric-fusiform, filiform, the ends acute,
often slightly curved, thin-walled, hyaline, guttulate,
smooth-walled.
Type species: Ceratosphaeria lampadophora (Berk. &
Broome) Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)
Notes: The genus Ceratosphaeria was introduced by
Niessl (1876) with C. lampadophora as the type species
and it is morphologically characterized in having globose
to pyriform, immersed to almost superficial, dark coloured
stromatic ascomata, leathery to fragile perithecial walls,
cylindric-clavate, short-stipitate asci, truncate to broadly
rounded at the apex, with an apical annulus and allantoid to
suballantoid, pale brown, aseptate ascopores (Niessl 1876;
Réblová 2006). In this study, we introduce two new species
in Ceratosphaeria. Five species in this genus have been
recorded from freshwater habitats.
Ceratosphaeria aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555644, Facesoffungi
number: FoF 05416, Fig. 11
493
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–2323
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 269–361 lm high, 196–284 lm diam., deeply
immersed with neck erumpent through host surface, globose to subglobose, brown to yellow brown, occurring
solitary. Neck long, surface smooth, at times with yellow
crystals. Peridium 29–43 lm thick, composed of an inner
layer of flattened hyaline cells, a middle layer of small,
polygonal to irregular, pale brown cells, an outer layer of
irregular, yellow brown, pseudoparenchymatic cells. Paraphyses 4–7 lm wide, longer than asci, long tapering
above. Asci 86–124 9 13–21 lm (
x = 105 9 17 lm,
n = 30), 8-spored, unitunicate, cylindrical to broadly fusiform, with a narrow apical ring. Ascospores 89–95 9 4–
7 lm (
x = 92 9 5.5 lm, n = 30), filiform, mostly 3-septate, guttulate, hyaline, smooth-walled.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in a freshwater river, April
2015, Z.L. Luo, S-639 (MFLU 18–2323, holotype; HKAS
92859, isotype), ex-type living culture MFLUCC 18–1337.
Notes: Ceratosphaeria aquatica resembles C. phialidica
in having deeply immersed ascomata, and filiform, hyaline
ascospores (Huhndorf et al. 2008). However, Ceratosphaeria aquatica differs from C. phialidica in having
broader cylindrical asci (13–21 vs. 5.5–6.5 lm) and guttulate, septate, larger ascospores (89–95 9 4–7 vs.
65–85 9 1.3–1.7 lm). Ceratosphaeria aquatica also
shares similar morphological characters with C. lignicola
with filiform, guttulate, hyaline ascospores. However,
Ceratosphaeria aquatica differs from C. lignicola in having 3-septate, wider ascospores (4–7 vs. 3.5–4.5 lm).
Phylogenetic analysis also shows that C. aquatica is distinct from C. lignicola (Fig. 15).
Ceratosphaeria lampadophora (Berk. & Broome) Niessl
: Sphaeria lampadophora Berk. & Broome, Ann. Mag.
nat. Hist., Ser. 3 3: 372 (1859)
Distribution: Australia, on submerged wood (Hyde
et al. 1997)
Asexual morph: Harpophora-like. Detailed descriptions
and illustrations refer to Réblová (2006).
Notes: LSU sequence data is available.
Ceratosphaeria phialidica (Shearer) Huhndorf et al.
: Pseudohalonectria phialidica Shearer Can. J. Bot.
67(7): 1950 (1989)
Distribution: USA, submerged wood (Shearer 1989).
Asexual morph: Harpophora-like.
Notes: Holotype NY-01050492 and NY-01050493,
other specimen collected from freshwater habitats: NY03380687. Sequence data is not available.
Ceratosphaeria pusilla (Fuckel) Sacc
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Fig. 14 Ophioceras submersum (MFLU 18–1459, holotype). a Appearance of neck on substrate. b–c Section of ascoma. d Structure of
peridium. e Paraphyses. f–h Asci. i–l Ascospores. Scale bars: b 150 lm, c 50 lm, d–l 20 lm
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495
123
496
b Fig. 15 Phylogram generated from maximum likelihood analysis
based on LSU and TEF1a sequence data for species of Magnaporthales (with Amplistroma erinaceum and A. caroliniana as
outgroup). The best scoring RAxML tree with a final likelihood
value of - 10365.588175 is presented. RAxML bootstrap support
values equal to or greater than 60% are given before the forward slash
(black). Bayesian posterior probability equal to or higher than 0.95
are given after the forward slash (red). Hyphen (‘-’) indicates a value
lower than 60% for RAxML and Bayesian posterior probability lower
than 0.95. Newly generated sequences are in red. Ex-type or exepitype strains are in bold
: Ceratostoma pusillum Fuckel Jb. nassau. Ver. Naturk. 23-24: 128 (1870) [1869-70]
Distribution: USA, submerged wood in southern Illinois
swamps (Shearer and Crane 1986).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Ceratosphaeria lignicola D.F. Bao, Z.L. Luo, H.Y. Su &
K.D. Hyde, sp. nov.
Index Fungorum number: IF 555645, Facesoffungi
number: FoF 05417, Fig. 12
Etymology: Referring to this fungus dwelling on wood.
Holotype: MFLU 18–1457
Saprobic on decaying wood, submerged wood in
freshwater. Asexual morph Undetermined. Sexual morph
Ascomata 390–470 lm diam., 500–600 lm high, solitary
or aggregated, deeply immersed, subglobose, dark brown
to black, with a yellow, cylindrical, periphysate neck.
Peridium 13.5–17.5 lm wide, composed of an inner layer
of flattened hyaline cells, a middle layer of small, polygonal to irregular, brown cells, an outer layer of irregular,
yellow brown, pseudoparenchymatic cells. Paraphyses
4.5–6.5 lm wide, numerous, septate, hyaline, tapering
distally,
smooth.
Asci
106–116 9 11–13 lm
(
x = 111 9 12 lm, n = 20), 8-spored, unitunicate, broadly
cylindrical, thin-walled, with a refractive apical apparatus.
Ascospores
94–102 9 3.5–4.5 lm
(
x = 98 9 4 lm,
n = 30), hyaline, filiform, tapering at both ends, guttulate,
smooth-walled.
Material examined: THAILAND, Khwaeng Hua Mak,
Khet Bang Kapi Krung Thep Maha Nakhon, saprobic on
decaying wood submerged in a freshwater stream, October
2017, Z.L. Luo, B-96, (MFLU 18–1457, holotype), ex-type
living culture, MFLUCC 18–0342.
Notes: Ceratosphaeria lignicola resembles C. phialidica
in having deeply immersed ascomata in the substrate
(Huhndorf et al. 2008). Ceratosphaeria lignicola differs
from C. phialidica in having broader cylindrical asci
(11–13 vs. 5.5–6.5 lm) and larger, guttulate, ascospores
(94–102 9 3.5–4.5 vs. 65–85 9 1.3–1.7 lm). Ceratosphaeria lignicola also shares similar morphological
characters with C. lampadophora (Huhndorf et al. 2008).
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However, Ceratosphaeria lignicola differs from C. lampadophora in having longer, filiform, aseptate ascospores
(94–102 vs. 52–72 lm), while C. lampadophora have
fusiform, 5–7-septate ascospores. Phylogenetic analysis
also shows that Ceratosphaeria lignicola is distinct from C.
lampadophora (Fig. 15).
Magnaporthaceae P.F. Cannon
Aquafiliformis Z.L. Luo, K.D. Hyde & H.Y. Su, gen. nov.
Index Fungorum number: IF 555646, Facesoffungi
number: FoF 05418
Etymology: ‘‘Aqua’’ referring to the aquatic habitats,
‘‘filiformis’’ referring to the filiform ascospores.
Saprobic on decaying wood. Asexual morph Undetermined. Sexual morph Ascomata immersed with neck
erumpent through host surface, globose to subglobose, dark
brown to black, solitary. Peridium composed of an inner
layer of flattened hyaline cells and an outer layer of
irregular, heavily pigmented, thick-walled, pale brown to
dark brown cells of textura angularis. Paraphyses hyaline,
septate, unbranched. Asci 8-spored, unitunicate, cylindrical
to clavate, hyaline. Ascospores filiform, aseptate, guttulate,
hyaline, smooth-walled.
Type species: Aquafiliformis lignicola Z.L. Luo, K.D.
Hyde & H.Y. Su
Notes: Aquafiliformis morphologically resembles Ceratosphaeria in having globose to subglobose, dark brown
to black ascomata, cylindrical, 8-spored, unitunicate asci
and filiform, hyaline, guttulate ascospores (Huhndorf et al.
2008). However, the phylogenetic analysis of combined
LSU, SSU, RPB2 and TEF1a sequence data show that our
newly obtained strains (MFLUCC 16–1341, MFLUCC
18–1338) clusters in Magnaporthaceae, while Ceratosphaeria belongs to the newly introduced Ceratosphaeriaceae (Fig. 1, clade 20, 22). Wijayawardene
et al. (2018) accepted 23 genera in the family Magnaporthaceae. Twenty genera with available molecular
sequence data are included in our phylogenetic analysis
and 17 of them clusters together, while the newly obtained
strains MFLUCC 16–1341 and MFLUCC 18–1338 form a
separate clade in Magnaporthaceae and close to Muraeriata (SMH 2008a, SMH 2008b) (Fig. 15). However, our
strain differs from Muraeriata species in having globose to
subglobose ascomata, a peridium composed of an inner
layer of flattened hyaline cells and an outer layer of
irregular, heavily pigmented, thick-walled, pale brown to
dark brown cells of textura angularis and filiform, aseptate
ascospores, while Muraeriata species have lageniform to
globose ascomata, a peridium with a middle layer of large,
polygonal to irregular, hyaline cells that collapse or rupture
creating large empty pockets, with an external brown crust
and narrowly fusiform, septate ascospores (Huhndorf et al.
2008). The genera Clavatisporella and Herbampulla were
Fungal Diversity (2019) 99:451–660
also placed in Magnaporthaceae, but the sequence data for
these two genera are unavailable, and morphology of our
fungus are different from those of Clavatisporella and
Herbampulla by its filiform ascospores (Scheuer and
Nograsek 1993; Hyde 1995a). Therefore, we introduce a
new genus Aquafiliformis to accommodate our collections.
Aquafiliformis lignicola Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555647, Facesoffungi
number: FoF 05419, Fig. 13
Etymology: Referring to this fungus dwelling on wood.
Holotype: MFLU 18–2325
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 275–335 lm high, 300–356 lm diam.,
immersed with erumpent neck through host surface, globose to subglobose, dark brown to black, solitary. Peridium
24–34 lm thick, composed of an inner layer of flattened
hyaline cells and an outer layer of irregular, heavily pigmented, thick-walled, pale brown to dark brown cells of
textura angularis. Paraphyses 4.5–7.5 lm wide, hyaline,
septate,
unbranched.
Asci
76–86 9 8–12 lm
(
x = 81 9 10 lm, n = 25), 8-spored, unitunicate, cylindrical to clavate, hyaline. Ascospores 57–69 9 2.5–3.5 lm
(
x = 63 9 3 lm, n = 20), filiform, guttulate, aseptate,
hyaline, smooth-walled.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in a freshwater River, April
2015, X.C. Tao, S-478 (MFLU 18–2325, holotype; HKAS
92814, isotype), ex-type living culture MFLUCC 16–1341;
Ibid., saprobic on decaying wood submerged in a freshwater stream in Cangshan Mountain, August 2016, H.W.
Shen, S-717 (MFLU 18–2324, paratype), living culture
MFLUCC 18–1338.
Notes: Aquafiliformis lignicola resembles Neogaeumannomyces bambusicola in having immersed, globose to
subglobose, solitary ascomata, unitunicate, cylindrical,
hyaline asci and filiform, hyaline, guttulate ascospores (Liu
et al. 2015a, b). However, Aquafiliformis lignicola differs
from Neogaeumannomyces bambusicola in having different sizes of ascomata, asci and ascospores. Aquafiliformis
lignicola has aseptate ascospores, while Neogaeumannomyces bambusicola has 2–3-septate ascospores. Phylogenetic analysis also support that they belong to different
genera (Fig. 15).
Ophioceraceae Klaubauf et al.
Ophioceras Sacc., Syll. fung. (Abellini) 2: 358 (1883)
Asexual morph Undetermined. Sexual morph Ascomata
globose to elongate-globose, superficial to immersed,
black, with a long neck. Peridium thick, blackened. Paraphyses hypha-like, hyaline, septate, numerous. Asci
8-spored, cylindrical, with small, refractive, apical rings.
497
Ascospores filiform, narrowly fusoid to cylindrical, septate,
hyaline.
Type species: Ophioceras dolichostomum (Berk. &
M.A. Curtis) Sacc., Syll. fung. (Abellini) 2: 358 (1883)
Notes: Saccardo (1883) introduced Ophioceras based on
O. dolichostomum (Berk. & M.A. Curtis) Sacc. Ophioceras
species are commonly encountered on decaying woody
substrates in freshwater habitats worldwide (Hyde 1992b;
Hyde and Goh 1998a; Shearer et al. 1999; Tsui et al.
2001a,b; Thongkantha et al. 2009).
Ophioceras aquaticus Hu et al.
Distribution: China, Yunnan Province, on submerged
wood (Hu et al. 2012c).
Asexual morph: Undetermined
Notes: Holotype IFRD 021–055. ITS, LSU and SSU
sequence data obtained from ex-type culture are available.
Ophioceras arcuatisporum Shearer et al.
Distribution: Canada, Manitoba, Lake Rosabella, on
submerged grasses; USA, Minnesota, Lake Itasca State
Park, Elk lake, on Typha sp., Shagawa lake, on submerged
herbaceous debri (Shearer et al. 1999).
Asexual morph: Undetermined
Notes: Holotype ILLS, Crane A-167–1. Sequence data
is not available.
Ophioceras commune Shearer et al.
Distribution: China, Hong Kong, Lam Tsuen River, on
submerged wood (Tsui et al. 2001b); Egypt, River Nile, on
submerged wood (Abdel-Aziz 2016); Japan, Koito River,
on submerged wood (Tsui et al. 2001a); Panama, Barro
Colorado Islanda, on twig submerged in Allee Creek
(Shearer et al. 1999); USA, Illinois, Louisiana, Minnesota,
on submerged wood (Shearer et al. 1999), Florida, on
submerged decaying wood (Raja et al. 2009b).
Asexual morph: Undetermined
Notes: Holotype ILLS, Shearer 408–2, other specimens
collected from freshwater habitats: HKAS 92587, HKAS
92640, HKAS 92569. ITS, LSU, SSU, RPB2 and TEF1a
sequence data are available.
Ophioceras dolichostomum (Berk. & M.A. Curtis) Sacc
: Sphaeria dolichostoma Berk. & M.A. Curtis, Soc.,
Bot. 10(no. 46): 388 (1868) [1869]
Distribution: Australia, north Queensland, submerged
wood (Hyde 1992b); Japan, Koito River, on submerged
wood (Tsui et al. 2001a); USA, Florida, on submerged
wood (Conway and Barr 1977); Seychelles, Riviere St
Marie-Louis, on submerged wood (Hyde and Goh 1998a).
Asexual morph: Undetermined
Notes: Holotype anon. 51 (anon. 580), other specimen
collected from freshwater habitats: BRIP 19330. ITS, LSU,
SSU, TEF1a and MCM7 sequence data are available.
Ophioceras fusiforme Shearer et al.
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b Fig. 16 Myrmecridium aquaticum (MFLU 18–1595, holotype) a,
b Colony on natural substrate. c, d Conidiophore with conidia. e,
f Conidiogenous cells with conidia. g–i Conidia. j Germinating
conidium. Culture on PDA from above (k) and reverse (l). Scale bars:
c, d 100 lm, e, f 30 lm, g–j 10 lm
Distribution: USA, Indiana, Shades State Park, small
stream, on submerged decorticated woody debris (Shearer
et al. 1999).
Asexual morph: Undetermined
Notes: Holotype ILLS, Shearer 837–1. SSU sequence
data obtained from ex-type culture is available.
Ophioceras guttulatum Tsui et al.
Distribution: China, Hong Kong, Tai Po. Lain Tsuen
River, on submerged wood (Tsui et al. 2001c); China,
Yunnan Province, on submerged bamboo (Cai et al.
2006a).
Asexual morph: Undetermined
Notes: Holotype IFRD 8819. Sequence data is not
available.
Ophioceras hongkongense Tsui et al.
Distribution: China, Hong Kong, Tai Po. Lain Tsuen
River, on submerged wood (Tsui et al. 2001c)
Asexual morph: Undetermined
Notes: Holotype IFRD 8820. LSU and SSU sequence
data are available.
Ophioceras leptosporum (S.H. Iqbal) J. Walker
Synonym: Gaeumannomyces leptosporus S.H. Iqbal,
Trans. Br. mycol. Soc. 58(2): 346 (1972)
Distribution: UK, submerged plant stalks (Iqbal 1972).
Asexual morph: Undetermined
Notes: Holotype K(M) 35072. ITS, LSU, SSU, RPB1,
TEF1a and MCM7 sequence data are available.
Ophioceras submersum D.F. Bao, Z.L. Luo, H.Y. Su &
K.D. Hyde, sp. nov.
Index Fungorum number: IF 555648, Facesoffungi
number: FoF 05420, Fig. 14
Etymology: Referring to the submerged habitats of the
fungus
Holotype: MFLU 18–1459
Saprobic on decaying wood, submerged wood in
freshwater. Asexual morph Undetermined. Sexual morph
Ascomata 500–600 lm diam., 300–400 lm high, scattered,
solitary, deeply immersed, subglobose or ellipsoidal, coriaceous, black, with a long black neck. Ostiole central, with
straight upright neck at one end, black, periphysate.
Peridium 25–31 lm, thick-walled, composed two layers,
inner layer of hyaline, small pseudoparenchyma cells, outer
layer of pseudoparenchyma cells occluded with brown
amorphous material, dark brown cells of textura angularis.
Paraphyses 7–10 lm wide, hyaline, septate, constricted at
septa, broader at base, tapering to the end, longer than asci,
499
smooth. Asci 115–137 9 10–11 lm (
x = 126 9 10.5 lm,
n = 15), 8-spored, unitunicate, cylindrical, subhyaline,
apically
rounded.
Ascospores
87–109 9 3–4 lm
(
x = 98 9 3.5 lm, n = 20), overlapping in ascus, filiform,
slightly curved, thin-walled, multi-septate, rounded at both
ends, hyaline, smooth-walled.
Material examined: THAILAND, Sai Khu Waterfall, on
submerged decaying wood, 1 September 2017, C.G. Lin,
B-25 (MFLU 18–1459, holotype), ex-type living culture,
MFLUCC 18–0211.
Notes: Ophioceras submersum clusters in Ophioceras
based on multi-gene phylogenetic analyses and is related to
O. dolichostomum, O. hongkongense and O. venezuelense
(Fig. 15). Ophioceras submersum resembles O. hongkongense in having subglobose, black ascomata with a long
black neck, hyaline, septate paraphyses, unitunicate,
cylindrical, subhyaline asci and filiform, hyaline, septate,
smooth ascospores (Tsui et al. 2001c). However, Ophioceras submersum differs from O. hongkongense by its
smaller ascomata and longer asci (Tsui et al. 2001c).
Phylogenetic analysis also shows that they are distinct
species (Fig. 1, clade 19).
Ophioceras tenuisporum Shearer et al.
Distribution: Panama, Barro Colorado Islanda, on twig
submerged in Allee Creek (Shearer et al. 1999);
Asexual morph: Undetermined
Notes: Holotype ILLS, Shearer 652–1. LSU, SSU and btubulin sequence data are available.
Ophioceras venezuelense Shearer et al.
Distribution: Venezuela, on submerged decorticated
wood (Shearer et al. 1999).
Asexual morph: Undetermined
Notes: Holotype ILLS, Crane A-109–1. SSU sequence
data is available.
Pseudohalonectriaceae Hongsanan & K.D. Hyde
Pseudohalonectria Minoura & T. Muroi, Trans. Mycol.
Soc. Japan 19(2): 132 (1978)
Asexual morph Hyphomycetous, phialidic. Phialides
hyaline, micronematous, flask-shaped. Conidia allantoid,
hyaline, aseptate. Sexual morph Ascomata immersed or
partially immersed, with a long neck, globose to subglobose. Peridium membranous. Paraphyses numerous, septate, hyaline. Asci unitunicate, cylindrical, straight or
curved, with J-, thimble-shaped apical ring. Ascospores
overlapping uniseriate to biseriate, multi-seriate, filiformes,
septate.
Type species: Pseudohalonectria lignicola Minoura &
T. Muroi, Trans. Mycol. Soc. Japan 19(2): 132 (1978)
Notes: The genus Pseudohalonectria was introduced to
accommodate P. lignicola (Minoura and Muroi 1978).
Hongsanan et al. (2017) introduced Pseudohalonectriaceae
as a new family within Magnaporthales to accommodate
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Pseudohalonectria. Sixteen species are accepted in this
genus, of which six species have been reported from
freshwater habitats worldwide (Minoura and Muroi 1978;
Shearer 1989a, b; Hyde et al. 1998b; Cai et al. 2002a).
Pseudohalonectria adversaria Shearer
Distribution: South Africa, Durban, Palmiet River, on
submerged wood (Hyde et al. 1998b); USA, Illinois, on
submerged woody debris from Deer Pond (Shearer
1989a, b).
Asexual morph: Undetermined
Notes: Holotype ILLS, CS-603–1. Sequence data is not
available.
Pseudohalonectria falcata Shearer
Fig. 17 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data for species of
Myrmecridium (with Acrodictys elaeidicola as outgroup). The best
scoring RAxML tree with a final likelihood value of - 4631.789675
is presented. RAxML bootstrap support values equal to or greater than
75% are given before the forward slash (black). Maximum parsimony
bootstrap support values equal to or greater than 75% are given after
the forward slash (red). Hyphen (‘-’) indicates a value lower than 75%
for RAxML and Maximum parsimony. Newly generated sequences
are in red. Ex-type strains are in bold
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Distribution: USA, Illinois, on submerged twig collected
from Quiver Creek (Shearer 1989).
Asexual morph: Undetermined
Notes: Holotype NY-01050484, paratype NY01050483, other specimens collected from freshwater
habitats NY-03380667, NY-03380668. LSU, SSU and
TEF1a sequence data are available.
Pseudohalonectria fuxianii Cai et al.
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Cai et al. 2002a).
Asexual morph: Undetermined
Notes: Holotype IFRD 8838. Sequence data is not
available.
Pseudohalonectria lignicola Minoura & T. Muroi
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Cai et al. 2002a); Japan, Koito River, on
submerged wood (Tsui et al. 2001a), Lake Biwa, on submerged wood (Minoura and Muroi 1978); USA, Illinois,
Indiana, on submerged wood in streams or Rivers (Shearer
1989a, b).
Asexual morph: Undetermined
Notes: Holotype HUT 40005. ITS, LSU, SSU, RPB1,
TEF1a, MCM7 and b-tubulin sequence data are available.
Pseudohalonectria longirostrum Shearer
Distribution: Panama, a twig submerged in Shannon
Creek (Shearer 1989).
Asexual morph: Undetermined
Notes: Holotype CS-656-1, NY. Sequence data is not
available.
Pseudohalonectria lutea Shearer
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Cai et al. 2002a); Chile, submerged
wood collected from a small stream (Shearer 1989a).
Asexual morph: Undetermined
Notes: Holotype NY-01050490, NY-01050491. LSU
sequence data is available.
Myrmecridiales Crous
Myrmecridiaceae Crous
Myrmecridium Arzanlou et al., Stud. Mycol. 58: 84 (2007)
Asexual morph Colonies flat, with immersed mycelium.
Conidiophores arising vertically and clearly distinct from
creeping hyphae, unbranched, straight or flexuose, septate,
thick-walled. Conidiogenous cells polyblastic, integrated,
terminally, cylindrical. Conidia solitary, subhyaline,
smooth or finely verrucose, obovoidal or fusiform, conidial
secession schizolytic. Sexual morph Ascomata solitary or
aggregated in small groups, immersed, hyaline to pale
brown. Papilla or short necks centrally located, opening
flush with the wood surface or slightly projecting. Ostiole
periphysate. Clypeus positioned slightly beneath the wood
surface. Ascomatal wall two layered. Paraphyses hyaline,
septate, slightly constricted at the septa, cylindrical. Asci
501
cylindrical, with a slender, long stipe. Ascospores ellipsoidal, delicately verruculose, septate, hyaline.
Type species: Myrmecridium schulzeri (Sacc.) Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 84 (2007)
Notes: The genus Myrmecridium was introduced by
Arzanlou et al. (2007) to accommodate Ramichloridiumlike taxa with hyaline mycelium, and pale to unpigmented
conidiophores with pimple-like conidiogenous denticles at
their apices. Twelve species are accepted in this genus
(Peintner et al. 2016; Réblová et al. 2016b; Tibpromma
et al. 2017) and two of them were collected from freshwater habitats (Réblová et al. 2016b; Tibpromma et al.
2017).
Myrmecridium aquaticum Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555649, Facesoffungi
number: FoF 05421, Fig. 16
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1595
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, hairy, brown. Mycelium immersed,
composed of septate, branched, smooth, hyaline hyphae.
Conidiophores 211–308 lm long, 5–7 lm wide
(
x = 258 9 6 lm, n = 20), macronematous, mononematous, erect, unbranched, multi-septate, straight or flexuous,
cylindrical, percurrently proliferating, brown, paler
towards the apex, smooth, nodose at the tip. Conidiogenous
cells holoblastic, polyblastic, integrated, terminal, later
becoming intercalary, subhyaline to pale brown. Conidia
14–16 lm long, 4–6 lm wide (
x = 15 9 5 lm, n = 20),
acropleurogenous, dry, obovoid, rounded at the apex,
pointed at the base, 3-septate, subhyaline, smooth-walled.
Sexual morph Undetermined
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April
2015, X.C. Tao, S-448 (MFLU 18–1595, holotype; HKAS
92833, isotype), ex-type living culture MFLUCC 15–0366,
KUMCC 15–0340; Jizu Mountain, saprobic on decaying
wood submerged in a freshwater stream, April 2017, N
Zhao, S-1158, living culture MFLUCC 18–1489; Cangshan
Mountain, saprobic on decaying wood submerged in a
freshwater stream, March 2014, X.Y. Liu, S-001.
Notes: Myrmecridium aquaticum resembles M. sorbicola in having solitary, erect, unbranched, multi-septate,
brown conidiophores, integrated, terminal and intercalary
conidiogenous cells and obovoid, smooth conidia rounded
at the apex (Crous et al. 2018). However, Myrmecridium
aquaticum differs from M. sorbicola in having larger
conidiophores (211–308 vs. 50–200 lm), 3-septate and
longer conidia (14–16 vs. 8–10 lm), while M. sorbicola
has almost 1-septate conidia with mucoid sheath
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Fig. 18 Sporidesmium lageniforme (MFLU 18–1594, holotype) a, b Colonies on substrate. c, d Conidiophore with conidia. e Conidiophores.
f Conidiogenous cells g–i Conidia j Germinating conidium. Culture on PDA from above (k) and reverse (l). Scale bars: c–e 50 lm, f–l 20 lm
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503
Fig. 19 Sexual morph of Sporidesmium lignicola (MFLU 18–2326, holotype) a, b Appearance of necks on substrate. c Section through
ascoma. d, e Structure of peridium. f Paraphyses. g–i Asci. j–o Ascospores. Scale bars: c 100 lm, d 50 lm, f–i 30 lm, j–o 5 lm
123
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Fig. 20 Asexual morph of Sporidesmium lignicola (DLU 1376) a Colonies on natural substrate. b Conidiophore with conidia. c–m Conidia.
Scale bars: b–m 20 lm
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surrounding conidium in median region. Phylogenetic
analysis shows that Myrmecridium aquaticum is distinct
from other Myrmecridium species (Fig. 17).
Myrmecridium fluviae Hyang B. Lee & T.T.T. Nguyen
Distribution: Korea, Jeonnam Province, Yeongsan
River located in Gwangju, from a freshwater sample
(Tibpromma et al. 2017).
Sexual morph: Undetermined
Notes: Holotype CNUFC YR61–1; ITS and LSU
sequence data are available.
Myrmecridium montsegurinum Réblová & J. Fourn
Distribution: France, Midi-Pyrénées: Ariège, Montségur, Le Lasset stream along D9 road, on submerged
wood of Fraxinus excelsior, Fagus sylvatica, Hedera helix
and Alnus glutinosa (Réblová et al. 2016b)
Asexual morph: Undetermined
Notes: Holotype PRM 934684, other specimens collected from freshwater habitats: PRM 934685, PRM
934686. ITS, LSU, SSU and RPB2 sequence data are
available.
Ophiostomatales Benny & Kimbr.
Ophiostomataceae Nannf.
Subbaromyces Hesselt., Bull. Torrey bot. Club 80: 511
(1953)
Asexual morph Conidiophores branched, septate. Conidia
hyaline, smooth-walled, asepate, exogenously formed,
ellipsoid. Sexual morph Ascomata partially submerged,
later superficial, membranous, syringe-shaped, beak divided into two portions by a large pronounced collar, with
upper portion tapering to a small ostiole, surrounded by a
fringe of hyphae. Paraphyses absent. Asci 8-spored, unitunicate, clavate, with autodigestible wall. Ascospores
2-celled, with cells of equal size, hyaline, surrounded by a
gelatinous envelope, released as a mucus-like droplet at tip
of perithecium.
Type species: Subbaromyces splendens Hesselt., Bull.
Torrey bot. Club 80: 511 (1953)
Notes: The genus was established by Hesseltine (1953)
for a taxon collected from trickling filter rocks in New
York, USA. Two species were accepted within this genus
(Hesseltine 1953; Chary and Ramarao 1974). The other
accepted species, S. aquaticus, was introduced by Chary
and Ramarao (1974) for a species isolated from water
samples collected in India. A phylogenetic analysis based
on SSU sequence data of S. splendens was provided by
Jones et al. (1999), and it showed that S. splendens is
closely related to Curvularia brachyspora. In updated
classifications, Subbaromyces was placed in the family
Ophiostomataceae (Ophiostomatales, Sordariomycetes)
(Maharachchikumbura et al. 2015, 2016; Wijayawardene
et al. 2017, 2018).
Subbaromyces aquaticus Manohar. & P. R. Rao
505
Distribution: India, on submerged wood in freshwater
(Chary and Ramarao 1974).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Phomatosporales Senan. et al.
Phomatosporaceae Senan. & K.D. Hyde
Phomatospora Sacc., Nuovo G. bot. ital. 7: 306 (1875)
Asexual morph Sporothrix-like, reported from culture
(Rappaz 1992). Sexual morph Ascomata solitary to rarely
gregarious, immersed or becoming erumpent with age,
globose or subglobose, light brown, dark brown to black,
coriaceous, sometimes developing under a small blackened
clypeus, ostiolate, papillate. Papilla short or rarely somewhat long, central or eccentric, cylindrical, sometimes
covered with black, amorphous material around the upper
region, periphyses hyaline, short, filiform. Peridium comprising small, brown pseudoparenchymatous cells forming
a textura angularis to textura prismatica or inner, hyaline,
thick-walled cells of textura angularis and outer, brown,
cells of textura angularis. Paraphyses comprising hyphalike, filamentous, septate or aseptate, slightly constricted at
the septa, distally tapering, hyaline. Asci 8-spored, unitunicate, cylindrical or oblong-fusiform, thin-walled, short
stalked or sessile, apex oblong with J-, apical apparatus.
Ascospores uniseriate, rarely biseriate, overlapping uniseriate to biseriate, ellipsoidal to fusiform, 0–3-septate, not
constricted at the septum, sometimes bi-guttulate, guttules
located at the ends of the cell, or longitudinally striate,
sometimes with filamentous appendages at both ends,
hyaline.
Type species: Phomatospora berkeleyi Sacc., Nuovo G.
bot. ital. 7(4): 306 (1875)
Notes: Phomatospora was placed in Ascomycota genera
incertae sedis based on phylogenetic analysis (Vijaykrishna et al. 2006). Senanayake et al. (2016) established the
family Phomatosporaceae (Phomatosporales) to accommodate the genera Phomatospora, Lanspora and
Tenuimurus. Members of the genus Phomatospora are
widely distributed in freshwater, marine and terrestrial
habitats. Seven species of Phomatospora are known from
freshwater habitats (Shearer and Raja http://fungi.life.uiuc.
edu/world_records; Nordén et al. 2015).
Phomatospora aquatica Minoura & T. Muroi
Distribution: Australia, Queensland, submerged wood
in stream (Vijaykrishna and Hyde 2006); Japan, on basal
wood submerged in Pond Nanatsuike (Minoura and Muroi
1978).
Asexual morph: Undetermined
Notes: Holotype HUT 40004. Sequence data is not
available.
Phomatospora berkeleyi Sacc
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506
Distribution: Australia, Queensland, submerged wood
in stream (Vijaykrishna and Hyde 2006); USA, Illinois,
Tinley Park Forest Preserve, Typha pond, on submerged
stems of Typha latifolia; Wisconsin, Trout lake, on submerged stems of Carex comosa, Big Muskellunge lake, on
submerged stems of Scirpus brevicaudatus, Allequash lake,
on submerged stems of Typha latifolia (Fallah and Shearer
1998).
Asexual morph: Undetermined
Notes: Holotype K(M) 49573, other specimens were
collected from freshwater: ILLS 53088, ILLS 53089, ILLS
53091. Sequence data is not available. Phomatospora
berkeleyi was originally collected from dead stalks of
Solanum on terrestrial habitats (Saccardo 1875). Fallah and
Shearer (1998) collected this species from freshwater
habitats and provided descriptions and illustrations.
Phomatospora helvetica H. Wegelin
Distribution: Norway, Aust-Agder, Arendal, Langevoll,
Nedenes, on submerged wood in small stream in temperate
deciduous forest (Nordén et al. 2015).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Phomatospora luteotingens J. Fourn. & Lechat
Distribution: France, Ariège: Caussou, Caussou Brook,
on submerged branch of Fraxinus excelsior, Hautes Pyrénées: Asque, La Gourgue, Arros stream, on submerged
twig of Alnus glutinosa; Spain, Asturias: road to La Endriga, on submerged wood of Alnus glutinosa (Fournier and
Lechat 2010)
Asexual morph: Undetermined
Notes: Holotype LIP, Fournier 09145. Sequence data is
not available.
Phomatospora muskellungensis Fallah & Shearer
Distribution: USA, Wisconsin, Big Muskellunge lake,
submerged wood of Typha latifolia, Allequash lake, on
submerged stems of Typha latifolia (Fallah and Shearer
1998).
Asexual morph: Undetermined
Notes: Holotype ILLS 53011. Sequence data is not
available.
Phomatospora striatigera Scheuer
Distribution: Australia, on submerged Carex gracili in
freshwater (Scheuer 1988)
Asexual morph: Undetermined
Notes: Holotype GZU, Scheuer, 18 Jun. 1981, Epitype
CBS 133932. ITS and LSU sequence data are available.
Phomatospora triseptata Raja & Shearer
Distribution: USA, Florida, Big Cypress National Preserve, on submerged soft, decorticated woody debris (Raja
and Shearer 2008).
Asexual morph: Undetermined
Notes: Holotype ILL 40114. Sequence data is not
available.
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Sporidesmiales Crous
Sporidesmiaceae Fr.
Sporidesmium Link, Mag. Gesell. naturf. Freunde, Berlin
3(1–2): 41 (1809)
Asexual morph Descriptions and illustrations refer to Su
et al. (2016). Sexual morph Descriptions and illustrations
refer to Zhang et al. (2017a, b).
Type species: Sporidesmium atrum Link, Mag. Gesell.
naturf. Freunde, Berlin 3(1–2): 41 (1809)
Notes: The genus Sporidesmium was established by
Link (1809) with S. atrum Link as the type. Sporidesmium
is a large and heterogeneous genus with 483 epithets
referred to the genus in Index Fungorum (December 2018).
However, many previously described species were revised
and transferred to over 30 genera (Iturriaga et al. 2008).
Studies based on phylogenetic analyses have been carried
out to further re-examine the classification of Sporidesmium-like taxa, given that the generic delimitations
based on morphological characters appear to be questionable, and it has been found that Sporidesmium and its
morphology similar genera are clearly not monophyletic,
and even they are distributed among different families and
orders within Dothideomycetes and Sordariomycetes
(Shenoy et al. 2006; Su et al. 2016; Yang et al. 2018a).
Sporidesmium was only found as an asexual morph genus
until Zhang et al. (2017a, b) introduced the first sexual
morph Sporidesmium thailandense which was collected
from freshwater habitats in Thailand.
Sporidesmium aquaticivaginatum J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, stream flowing outside Kaeng Krachan National
Park, on submerged wood (Hyde et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1159, isotype HKAS 95046.
ITS, LSU, SSU, RPB2 and TEF1a sequence data are
available.
Sporidesmium cangshanense Z.L. Luo & K.D. Hyde,
nom. nov.
Facesoffungi number: FoF 05422
: Sporidesmium aquaticum H.Y. Su, Z.L. Luo & K.D.
Hyde, Fungal Divers 80:398 (2016)
Distribution: China, Yunnan Province, Dali city, on
submerged wood in a freshwater stream in Cangshan
Mountain (Su et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKAS 84029. LSU sequence data is
available. Sporidesmium aquaticum was introduced by
Arambarri et al. (1989). Su et al. (2016) introduced a new
Sporidesmium species also called Sporidesmium aquaticum
which is a homonym and thus invalid. We replace the latter
invalid name, Sporidesmium aquaticum, as Sporidesmium
cangshanense.
Sporidesmium dulongense Luo et al.
Fungal Diversity (2019) 99:451–660
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in Dulong River (Hyde et al.
2019).
Sexual morph: Undetermined
Notes: Holotype HKAS 92659. ITS, LSU, RPB2 and
TEF1a sequence data are available.
Sporidesmium fluminicola H.Y. Su & K.D. Hyde
Distribution: China, Yunnan Province, Dali, Cangshan
Mountain, saprobic on decaying wood submerged in
stream (Su et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKAS 84003. ITS, LSU and SSU
sequence data are available.
Sporidesmium gyrinomorphum Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Yang
et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0851, isotype GZAAS
17–0004. ITS, LSU, SSU, RPB2 and TEF1a sequence data
are available.
Sporidesmium lageniforme Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555650, Facesoffungi
number: FoF 05423, Fig. 18
Etymology: Referring to the lageniform conidia of the
fungus
Holotype: MFLU 18–1594
Saprobic on decaying wood submerged in freshwater.
Asexual morph Colonies on the substratum superficial,
effuse, scattered, hairy, black. Mycelium mostly immersed,
comprising of branched, septate, smooth-walled, brown
hyphae. Conidiophores 105–141 lm long, 4–6 lm wide
(
x = 123 9 5 lm, n = 20), macronematous, mononematous, erect, straight or flexuous, unbranched, 4–6-septate,
greyish brown to dark brown, smooth. Conidiogenous cells
monoblastic, integrated, terminal, determinate, cylindrical,
dark brown. Conidia 38–48 lm long, 13–17 lm wide
(
x = 43 9 15 lm, n = 20), acrogenous, solitary, lageniform, truncate at base, tapering at apex, 5–7-septate, subhyaline to greyish at immature, dark olivaceous-brown at
maturity, hyaline towards the apex, smooth-walled. Sexual
morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on decaying submerged wood, October
2016, Z.L. Luo, S-880 (MFLU 18–1594, holotype), ex-type
living culture DLUCC 0880.
Notes: Sporidesmium lageniforme resembles S. dulongense in having unbranched, dark brown, septate conidiophores, monoblastic, terminal, determinate conidiogenous
cells and acrogenous, solitary, septate conidia truncate at
base, tapering at apex. Sporidesmium lageniforme differs
from S. dulongense in having shorter conidia (38–48 vs.
507
50–58 lm) and conidia of S. dulongense have sphaerical
guttules in each cell while conidia of S. lageniforme mostly
have no guttules. Phylogenetic analysis also shows that
Sporidesmium lageniforme and S. dulongense are distinct
species (Fig. 1, clade 7).
Sporidesmium lignicola Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555651, Facesoffungi
number: FoF 05424, Figs. 19, 20
Etymology: Referring to the fungus dwelling on wood.
Holotype: MFLU 18–2326
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse on natural substrate, scattered, pale brown to brown. Mycelium
immersed, composed of septate, branched, brown, smooth
hyphae. Conidiophores 50–70 lm long, 3–4 lm wide
(
x = 60 9 3.5 lm, n = 20), macronematous, mononematous, unbranched, erect, straight or flexuous, brown to dark
brown, septate, smooth. Conidiogenous cells holoblastic,
monoblastic, terminal, determinate, dark brown, cylindrical. Conidia 21–27 lm long, 4.5–6.5 lm wide
(
x = 24 9 5.5 lm, n = 20), acrogenous, solitary, dry, clavate or cylindrical, straight or slight curved, truncate at
base, brown, mostly 3-septate, formed in chains, smooth.
Sexual morph Ascomata 190–330 lm high, 160–340 lm
diam., immersed with neck erumpent through host surface,
subglobose to ellipsoid, scattered, 2–4 locules, pale brown
to brown. Ostiole 185–365 lm long, 78–122 lm wide,
cylindrical, central or lateral, brown at the base, becoming
hyaline towards the apex. Peridium 30–44 lm thick, twolayered, outer layer comprising pale brown to brown,
oblong and rounded cells, inner layer comprising several
layers of hyaline, large cells of textura angularis or
irregular cells. Paraphyses 2.5–4.5 lm wide, hyaline,
unbranched, septate, slightly constricted at the septum. Asci
87–115 9 8–10 lm (
x = 101 9 9 lm, n = 35), 8-spored,
unitunicate, cylindrical, with a tapering pedicel, apically
rounded, with a distinct, relatively small, refractive,
wedge-shaped apical ring. Ascospores 13–15 9 6–8 lm
(
x = 14 9 7 lm, n = 35), uni-seriate, fusiform, aseptate or
uniseptate, guttulate, hyaline, smooth-walled.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in a freshwater river, April
2015, X.C. Tao, S-429 (MFLU 18–2326, holotype), extype living culture KUMCC 15–0266; Cangshan Mountain,
saprobic on decaying wood submerged in a freshwater
stream, July 2017, H.W. Shen, S-1376 (DLU 1376, paratype), living culture DLUCC 1376.
Notes: The sexual morph of Sporidesmium lignicola
resembles Sporidesmium thailandense in having subglobose to ellipsoid, scattered, immersed ascomata with necks
erumpent through the host surface, hyaline, unbranched,
123
508
septate paraphyses, cylindrical asci apically rounded with
an apical ring and fusiform, hyaline ascospores (Zhang
et al. 2017). However, Sporidesmium lignicola differs from
S. thailandense in having ascomata with 2–4 locules, a
two-layered peridium, smaller asci (87–115 9 8–10 vs.
160–220 9 11–14 lm), and aseptate or uniseptate, smaller
ascospores (13–15 9 6–8 vs. 23–28 9 8–10 lm) with
small guttules, while S. thailandense has 3-septate
Fungal Diversity (2019) 99:451–660
ascospores with 2–5 prominent guttules. The asexual
morph of Sporidesmium lignicola can be easily distinguished from other Sporidesmium asexual morph species in
having clavate or cylindrical conidia formed in chains.
Phylogenetically, our two newly obtained strains cluster
together with high support (100% ML) and is distinct from
other Sporidesmium species (Fig. 1, clade 7). We therefore
Fig. 21 Neospadicoides aquatica (MFLU 18–1605, holotype) a–c Conidiophores with conidia. d, e Conidiogenous cells with percurrent
proliferations. f–k Conidium. Scale bars: a–c 40 lm, d, e 25 lm, f–k 10 lm
123
Fungal Diversity (2019) 99:451–660
introduce a new species Sporidesmium lignicola with both
asexual and sexual morphs.
Sporidesmium olivaceoconidium J. Yang & K.D. Hyde
Distribution: Thailand, Chiang Rai Province, stream
flowing in Tham Luang Nang Non Cave, on submerged
wood (Hyde et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1175. ITS, LSU, SSU,
RPB2 and TEF1a sequence data are available.
Sporidesmium pyriformatum J. Yang & K.D. Hyde
Distribution: Thailand, Khiri Khan Province, Hua Hin,
stream flowing outside Kaeng Krachan National Park, on
submerged wood (Hyde et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1155. ITS, LSU, SSU,
RPB2 and TEF1a sequence data are available.
Sporidesmium submersum H.Y. Su & K.D. Hyde
Distribution: China, Yunnan Province, Dali, Cangshan
Mountain, saprobic on decaying wood submerged in
stream (Su et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKAS 84004. ITS, LSU and SSU
sequence data are available.
Sporidesmium thailandense Dong et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged wood in a small River (Zhang et al.
2017a, b; Yang et al. 2018a).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2709, other specimen collected from freshwater habitats MFLU 15–1152. ITS, LSU,
SSU, RPB2 and TEF1a sequence data are available.
Sporidesmium tropicale M.B. Ellis
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Yang
et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype IMI 37498, other specimen collected
from freshwater MFLU 17–0850. ITS, LSU, SSU, RPB2
and TEF1a sequence data are available. Sporidesmium
tropicale was found on dead branches of woody plants and
is widely distributed in tropical areas (Ellis 1958; Wu and
Zhuang 2005). Yang et al. (2018a, b) collected this species
from freshwater habitats in Thailand and provided the
illustrations and descriptions.
Tirisporellales Suetrong et al.
Tirisporellaceae Suetrong et al.
Thailandiomyces Pinruan, Sakayaroj, Hyde & Jones,
Fungal Diversity 29: 91 (2008)
Asexual morph Colonies on natural substrate effuse,
black. Mycelium superficial. Conidiophores macronematous, mononematous, erect, brown, paler towards the apex,
straight or flexuous, branched or unbranched.
509
Conidiogenous cells enteroblastic, monophialidic, integrated, terminal, with a large and distinct collarette; collarette funnel-shaped. Conidia cylindrical, ellipsoid or
obovoid, thick-walled, brown, aseptate. Sexual morph
Ascomata scattered to gregarious, semi-immersed to
superficial, globose, black, coriaceous, ostiolate, with long
cylindrical necks, periphysate with short hyaline cells.
Peridium composed of compressed cells of textura angularis, black outside, becoming brown inwardly. Paraphyses
present but deliquescent, irregular in width, rarely septate,
tapering towards the apices, embedded in a mucilaginous
matrix. Asci 8-spored, unitunicate, cylindrical to clavate,
apedicellate, free-floating, apically truncate, with a Jsubapical ring. Ascospores uniseptate, obliquely overlapping 2-seriate, fusoid, straight or broadly curved, with
bipolar appendages, hyaline, smooth-walled.
Type species: Thailandiomyces bisetulosus Pinruan
et al., Fungal Divers 29: 91 (2008)
Notes: The genus Thailandiomyces was introduced by
Pinruan et al. (2008) to accommodate one ascomycete
species collected from submerged trunk of Licuala longicalycata in Thailand. This species was apparently linked
with a Craspedodidymum asexual morph in culture. This
genus remains monotypic. Based on phylogenetic analyses,
Suetrong et al. (2015) introduced a new family
Tirisporellaceae, typified by a new genus Tirisporella
E.B.G. Jones, K.D. Hyde & Alias. The genus Thailandiomyces phylogenetically resides in this family.
Thailandiomyces bisetulosus Pinruan et al.
Distribution: Thailand, on submerged trunk of Licuala
longicalycata (Pinruan et al. 2008).
Asexual morph: Craspedodidymum licualae Pinruan
Notes: Holotype BBH, Pinruan, Wah 110. LSU and SSU
sequence data are available.
Togniniales Senan. et al.
Togniniaceae Réblová et al.
Phaeoacremonium Gams et al., Mycologia 88: 789 (1996)
Asexual morph Mycelium consisting of branched, septate
hyphae, single or bundled. Conidiophores branched in the
basal region or unbranched, arising from aerial or submerged hyphae, erect, nearly cylindrical when unbranched,
slightly tapering, straight or flexuous, variable in length, up
to 7-septate, mostly pale brown, paler towards the tip.
Conidiogenous cells mostly monophialidic, discrete or
integrated, terminal or intercalary, sometimes polyphialidic, sparsely warted, pale brown to hyaline, verruculose
or smooth. Conidia aggregated into round, slimy heads at
the apices of phialides, aseptate, hyaline, smooth-walled;
oblong-ellipsoidal to obovate, cylindrical, allantoid or
reniform, uncommonly fusiform-ellipsoidal or globose,
becoming guttulate with age. Sexual morph Ascomata
aggregated or solitary, superficial to immersed,
123
510
Fungal Diversity (2019) 99:451–660
Fig. 22 Neospadicoides lignicola (MFLU 18–1606, holotype) a, b Conidiophores with conidia. c Conidophore apex with discrete conidiogenous
cells. d–f Conidia. g Germinating conidium. Culture on PDA from above (h) and reverse (i). Scale bars: a, b 50 lm, c 30 lm, d–g 20 lm
123
Fungal Diversity (2019) 99:451–660
511
Fig. 23 Neospadicoides yunnanensis (MFLU 18–2329, holotype) a Colonies on substrate. b Conidiophore with conidiogenous cells. c,
d Conidiophore with conidia. e–h Conidia. Scale bars: c, d 50 lm, b, e 20 lm, f–h 5 lm
123
512
nonstromatic, globose to subglobose, dark, opaque, longnecked. Paraphyses abundant, broadly cellular, slightly
constricted at the septa, branching, hyaline, slightly tapering apically or thread-like towards the apex. Asci 8-spored,
unitunicate, arising in acropetal succession, appearing
spicate when mature, ascal apex thickened without a discharge mechanism, basally bluntly obtuse, sessile. Ascospores mostly biseriate or in a single row, allantoid,
reniform, cylindrical or oblong-ellipsoidal, aseptate,
hyaline.
Type species: Phaeoacremonium parasiticum (Ajello,
Georg & C.J.K. Wang) Gams et al., Mycologia 88(5): 794
(1996)
Notes: Phaeoacremonium has recently been monographed, and comprises 46 species (Gramaje et al. 2015;
Maharachchikumbura et al. 2016). Phaeoacremonium
species are saprobic on plants, or pathogenic on human and
animals or soil fungi (Mostert et al. 2006;
Maharachchikumbura et al. 2016). Hu et al. (2012c)
introduced a new species, Togninia aquatica, collected
from freshwater habitats. Gramaje et al. (2015) combined
this species as Phaeoacremonium aquaticum.
Phaeoacremonium aquaticum (Hu et al.) Gramaje et al.
: Togninia aquatica Hu et al. Mycologia 104(6): 1482
(2012)
Distribution: China, Yunnan Province, Jinghong City,
Mengla County, on submerged wood in a small stream (Hu
et al. 2012c).
Asexual morph: Undetermined
Notes: Holotype IFRD 023–047. ITS sequence data is
available.
Phaeoacremonium ovale Huang et al.
Distribution: China, Yunnan Province, Baoshan City,
on submerged decayed wood in a stream along the roadside
(Huang et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype HKAS 99550, isotype MFLU 18–1076.
ITS, actin and b-tubulin sequence data are available.
Trichosphaeriales M.E. Barr
Trichosphaeriaceae G. Winter
Brachysporium Sacc., Syll. fung. (Abellini) 4: 423 (1886)
Asexual morph Colonies effuse, brown, velvety. Mycelium mostly immersed, composed of branched, septate,
subhyaline to brown hyphae. Conidiophores mononematous, macronematous, erect, straight or slightly flexuous,
smooth, thick-walled, septate, unbranched, cylindrical,
brown in the bottom, paler and tapering toward the apex.
Conidiogenous cells holoblastic, terminal, integrated,
hyaline, denticulate, proliferating sympodially. Conidia
acropleurogenous, septate, smooth, thick-walled, fusoid to
limoniform, polar cells subhyaline, narrowing at the apex,
median cells brown. Sexual morph Ascomata partly
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Fungal Diversity (2019) 99:451–660
immersed to superficial, solitary, globose to subglobose,
conical around the ostiole, papillate, dark brown to black,
glabrous. Peridium leathery to fragile, consisting of two
regions; outer region of carbonaceous, dark brown, angular
to rectangular cells; inner region of hyaline, thin-walled,
elongated, compressed cells. Ostiolar canal periphysate.
Paraphyses persistent, branched, hyaline, septate, irregular
in width. Asci 8-spored, unitunicate, cylindrical-clavate,
with long, slender stipe, broadly rounded to truncate at the
apex, with distinct refractive apical annulus. Ascospores
biseriate, ellipsoidal to fusiform to oblong-lemon-shaped,
at first aseptate, followed by formation of a median septum,
hyaline, smooth-walled.
Type species: Brachysporium obovatum (Berk.) Sacc.,
Syll. fung. (Abellini) 4: 427 (1886)
Notes: The asexual morph genus Brachysporium was
established by Saccardo in 1880. Réblová and Seifert
(2004a) introduced a new genus Cryptadelphia, with type
species C. groenendalensis (Sacc., E. Bomm. & M. Rouss.)
Réblová & Seifert to accommodate six sexual morphs of
Brachysporium. Many Brachysporium species were
reported saprobic on decaying wood of different substrates
(Réblová and Seifert 2004a; Markovskaja and Treigien
2007). Some species were also described from marine
habitats, e.g. B. belgolandicum Schaumann and B. helgolandicum Schaumann. Among the accepted Brachysporium species, only two are known from freshwater habitats
(Lamore and Goos 1978; Raja et al. 2009b).
Brachysporium obovatum (Berk.) Sacc.
: Helminthosporium obovatum Berk., Annals and
Magazine of Natural History 6: 434 (1841)
Distribution: USA, Florida, on submerged substrate in
lentic habitat in Ocala National Forest (Raja et al. 2009b).
Sexual morph: see Réblová and Seifert (2004a)
Notes: Sequence data is not available.
Brachysporium nigrum (Link) S. Hughes
Distribution: USA, Rhode Island, on wood in the Saugatucket River (Lamore and Goos 1978).
Sexual morph: see Réblová and Seifert (2004a).
Notes: LSU, SSU and RPB2 sequence data are
available.
Unisetosphaeria Pinnoi et al., Mycoscience 44: 377 (2003)
Asexual morph Undetermined. Sexual morph Ascomata
immersed to superficial, scattered, pyriform, hyaline to
light brown, dark brown near the apex, coriaceous, ostiolate, papillate. Papilla periphysate, surrounded by short
dark hairs. Seta single, composed of several rows of brown
cells, arising from the ostiolar region. Peridium composed
of angular brown-walled cells. Paraphyses sparse, obscure,
comprising short rows of ovoid to oblong cells. Asci
8-spored, unitunicate, clavate, short pedicellate, apically
Fungal Diversity (2019) 99:451–660
truncate, with a refractive, J-, apical ring. Ascospores
2-seriate, septate, hyaline.
Type species: Unisetosphaeria penguinoides Pinnoi
et al., Mycoscience 44(5): 378 (2003)
Notes: This monotypic genus was introduced by Pinnoi
et al. (2003). Its taxonomic placement was between
Chaetosphaeriaceae and Trichosphaeriaceae. However,
Unisetosphaeri penguinoides has several incompatible
characters of ascomata, paraphyses and asci which are
against the Chaetosphaeriaceae. The characters of ascomata, peraphyses, asci and ascospores are more congruent
for Trichosphaeriaceae (Pinnoi et al. 2003). Thus, it was
suggested to assign this genus in Trichosphaeriaceae based
on morphology. This suggestion was followed by
Maharachchikumbura et al. (2015, 2016).
Unisetosphaeria penguinoides Pinnoi et al.
Distribution: Thailand, Narathiwat Province, on submerged petiole of Eleiodoxa conferta (Pinnoi et al. 2003).
Asexual morph: Undetermined
Notes: Holotype BBH Aom 103. Sequence data is not
available.
Xenospadicoidales Hern.-Restr. et al.
Xenospadicoidaceae Hern.-Restr. et al.
Neospadicoides Z.L. Luo, K.D. Hyde & H.Y. Su, gen. nov.
Index Fungorum number: IF 555652, Facesoffungi
number: FoF 05425
Etymology: Referring to the genus similar to
Spadicoides.
Saprobic on decaying wood. Asexual morph Colonies
effuse, hairy, brown to dark brown. Mycelium partly
superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores
macronematous, mononematous, solitary or in groups,
erect, unbranched, septate, straight or flexuous, cylindrical,
brown, paler towards the apex, smooth. Conidiogenous
cells holoblastic, enteroblastic, percurrent, polytretic,
integrated, terminal. Conidia acrogenous or acropleurogenous, fusiform, obovoid, septate, smooth-walled. Sexual
morph Undetermined.
Type species: Neospadicoides lignicola Z.L. Luo, K.D.
Hyde & H.Y. Su
Notes: Neospadicoides is morphologically similar to
Spadicoides in having effuse, hairy colonies on natural
substrate, mycelium composed of septate, branched,
smooth hyphae, conidiophores macronematous, mononematous, erect, straight or flexuous, cylindrical, smooth,
conidiogenous cells integrated, terminal and smooth conidia (Hughes 1958; Seifert et al. 2011). However,
Neospadicoides differs from Spadicoides in having
unbranched conidiophores and acrogenous or acropleurogenous, septate conidia while Spadicoides have branched
or unbranched conidiophores, tretic conidiogenous cells
513
and acropleurogenous, aseptate or septate conidia (Hughes
1958; Goh and Hyde 1996a; Seifert et al. 2011). The
phylogenetic analysis show that our Neospadicoides species cluster together with good support value and form a
distinct subclade from Spadicoides within Xenospadicoidaceae (Xenospadicoidales) (Fig. 1, clade 6).
Neospadicoides aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555654, Facesoffungi
number: FoF 05426, Fig. 21
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–1605
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, brown to dark
brown. Mycelium partly superficial, partly immersed,
composed of septate, branched, smooth, pale brown
hyphae. Conidiophores (80–)126–194(–218) lm long, 5–
7 lm wide (
x = 160 9 6 lm, n = 20), macronematous,
mononematous, solitary or in groups, erect, unbranched,
septate, straight or flexuous, cylindrical, brown, paler
towards the apex, smooth, pecurrently proliferating.
Conidiogenous cells holoblastic, integrated, terminal, subhyaline, percurrently proliferating. Conidia 18–22 lm
long, 7–9 lm wide (
x = 20 9 8 lm, n = 20), acrogenous
or acropleurogenous, subhyaline to pale brown when
young, brown to dark brown when matured, fusiform to
cylindrical, rounded at the apex, truncate at the base,
2-septate, with thick and dark band at septa, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Gaoligong
Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2015, X.J. Su, S-701 (MFLU 18–1605,
holotype), ex-type living culture MFLUCC 17–2217.
Notes: Neospadicoides aquatica resembles Spadicoides
americana in having macronematous, mononematous,
erect, unbranched, septate, solitary or in groups conidiophores paler towards the apex, integrated, terminal conidiogenous cells and 2-septate, smooth conidia (Wongsawas
et al. 2008). However, Neospadicoides aquatica differs
from Spadicoides americana in having larger conidia
(18–22 9 7–9 vs. 10–14.5 9 4–6.5 lm) which are rounded at the apex, truncate at the base.
Neospadicoides lignicola Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555653, Facesoffungi
number: FoF 05427, Fig. 22
Etymology: Referring to this fungus dwelling on wood.
Holotype: MFLU 18–1606
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, brown to dark
brown. Mycelium partly superficial, partly immersed,
composed of septate, branched, smooth, pale brown
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Fungal Diversity (2019) 99:451–660
b Fig. 24 Acrodictys fluminicola (DLU 0274, holotype) a, b Colonies
on natural substrate. c, d Conidiophores with conidia. e, f Conidiophores with conidiogenous cells. g–o Conidia. p Germinating conidium. Culture on MEA, q from above, r from reverse. Scale bars: c–
f 50 lm, g–p 15 lm
hyphae. Conidiophores (70–)134–276(–303) lm long, 5–
7 lm wide (
x = 205 9 6 lm, n = 20), macronematous,
mononematous, solitary or in groups, erect, unbranched,
septate, straight or flexuous, cylindrical, brown, paler
towards the apex, smooth. Conidiogenous cells holoblastic,
discrete, terminal and intercalary, subhyaline to pale
brown. Conidia 20–24 lm long, 9–13 lm wide
(
x = 22 9 11 lm, n = 20), acrogenous or acropleurogenous, pale brown to brown, obovoid, rounded at the apex,
truncate at the base, almost 2–3-septate, with dark band at
septa, guttulate, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on decaying wood submerged in
a freshwater stream, July 2015, X.J. Su, S-735 (MFLU
18–1606, holotype), ex-type living culture MFLUCC
17–2444.
Notes: Neospadicoides lignicola resembles N. yunnanensis in having macronematous, mononematous, solitary
or in groups, erect, unbranched, septate, straight or flexuous, cylindrical, brown conidiophores paler towards the
apex, integrated, terminal conidiogenous cells and obovoid,
septate, guttulate conidia (this study). However,
Neospadicoides lignicola differs from N. yunnanensis in
having
larger
conidia
(20–24 9 9–13
vs.
7.5–10.5 9 4–6 lm), and conidiogenous cells of N. yunnanensis are with pale-colored pores at the conidiogenous
loci. Phylogenetic analysis also shows that N. lignicola and
N. yunnanensis are different species (Fig. 1, clade 6). To
further support Neospadicoides lignicola as a new species,
we compared nucleotide differences with N. yunnanensis
following the guidelines of Jeewon and Hyde (2016).
Comparison of the 521 nucleotides across the ITS region
reveals 40 bp differences (7.67%) including 3 gaps.
Neospadicoides yunnanensis Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555655, Facesoffungi
number: FoF 05428, Fig. 23
Etymology: Referring to the location where this fungus
was collected, Yunnan Province, China.
Holotype: MFLU 18–2329
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on decaying wood
effuse, pale brown, hairy. Mycelium partly immersed in the
substrata, composed of pale brown, septate, smooth,
branched hyphae. Conidiophores 113–153 lm long, 4–
515
6 lm wide (
x = 133 9 5 lm, n = 20), macronematous,
mononematous, erect, unbranched, dark brown, paler
towards the apex, straight or slightly flexuous, cylindrical,
septate, smooth, thick-walled, occasionally swollen at the
apex. Conidiogenous cells polytretic, integrated, terminal
and intercalary, with pale colored pores remaining at the
conidiogenous loci. Conidia 7.5–10.5 lm long, 4–6 lm
wide (
x = 9 9 5 lm, n = 20), acropleurogenous, solitary,
pale brown, obovoid, guttulate, mostly 2-septate, comprising a proximal euseptum and a distal distoseptum, with
dark band at basal septa, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on decaying wood submerged,
December 2017, X.J. Su, S-1499 (MFLU 18–2329, holotype), ex-type living culture DLUCC 1499.
Notes: Neospadicoides yunnanensis resembles Spadicoides hodgkissa in having macronematous, mononematous, erect, solitary, unbranched, dark brown, paler towards
the apex, straight or slightly flexuous, cylindrical, septate,
smooth conidiophores occasionally swollen at the apex,
polytretic, integrated, terminal and intercalary conidiogenous cells with pale colored pores remaining at the
conidiogenous loci and solitary, obovoid, guttulate conidia
comprising a proximal euseptum and a distal distoseptum
(Ho et al. 2002b). However, Neospadicoides yunnanensis
differs from Spadicoides hodgkissa in having pale brown
conidia with dark band at basal euseptum while Spadicoides hodgkissa has versicolored conidia comprising a
proximal euseptum and a distal distoseptum and the distal
distoseptum possesses a conspicuous central pore which is
surrounded by a pigmented ring, appearing as dolipores in
side view.
Spadicoides S. Hughes, Can. J. Bot. 36: 805 (1958)
Asexual morph Descriptions and illustrations refer to
Réblová et al. (2018). Sexual morph Undetermined.
Type species: Spadicoides bina (Corda) S. Hughes, Can.
J. Bot. 36: 806 (1958)
Notes: The genus Spadicoides was introduced by
Hughes (1958) with S. bina as the type species. Goh and
Hyde (1996a) briefly discussed the generic concept of
Spadicoides. Thirty-one Spadicoides species were revised,
of which 21 species were accepted. Presently, 55 epithets
are listed in Index Fungorum (December 2018), and most
of the species are reported from terrestrial habitats
(Subramanian and Vittal 1974; Kuthubutheen and Nawawi
1991a; Wong et al. 2002; Li 2010; Xia et al. 2014; Ma et al.
2016). Seven species are known from freshwater habitats
(Goh and Hyde 1996a; Ho et al. 2002b; Cai et al.
2004a, 2006a, b, c; Zhuang 2001).
Spadicoides americana C.J.K. Wang
123
516
Fig. 25 Barbatosphaeria lignicola (HKAS 84005, holotype) a Specimen. b, c Appearance of necks on substrate. d Horizontal section
through ascomata. e, f Section through ascomata. g, h Structure of
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Fungal Diversity (2019) 99:451–660
peridium. i Paraphyses. j–n Ascus. o–p Ascospore. Scale bars: e,
f 150 lm, g–i 30 lm, j–p 5 lm
Fungal Diversity (2019) 99:451–660
Distribution: China, Zhejiang Province, Hangzhou city,
on submerged wood in a stream at Nine Creeks (Wongsawas et al. 2008).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater habitats
HMZFW 00002. Sequence data is not available.
Spadicoides atra (Corda) S. Hughes
: Chloridium atrum Corda, Icones fungorum hucusque
cognitorum 1: 17, t. 4:235 (1837)
: Psilonia atra (Corda) Corda, Icones fungorum
hucusque cognitorum 4: 27, t. 6:84 (1840)
: Catenularia atra (Corda) Sacc., Sylloge Fungorum 4:
304 (1886)
Distribution: China, Hong Kong, on submerged wood
(Zhuang 2001).
Sexual morph: Undetermined
Notes: LSU, SSU and RPB2 sequence data are
available.
Spadicoides bambusicola Zhou et al.
Distribution: China, Yunnan Province, on submerged
wood (Cai et al. 2006a).
Sexual morph: Undetermined
Notes: Holotype HKU (M) 8333 (now in IFRD).
Sequence data is unavailable.
Spadicoides cordanoides Goh & K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Goh and Hyde 1996a).
Sexual morph: Undetermined
Notes: Holotype BRIP 23201. Sequence data is not
available. This species is known only from Australia on
submerged wood in freshwater habitats (Goh and Hyde
1996a).
Spadicoides hodgkissa Ho et al.
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream (Ho et al. 2002b)
Sexual morph: Undetermined
Notes: Holotype HKU(M) 6155 (now in IFRD).
Sequence data is unavailable.
Spadicoides minuta Cai et al.
Distribution: China, Yunnan Province, on Phyllostachys
bambusoides submerged in a small stream (Cai et al.
2004a).
Sexual morph: Undetermined
Notes: Holotype IFRD 8850. Sequence data is
unavailable.
Spadicoides obovata (Cooke & Ellis) S. Hughes
: Acrothecium obovatum Cooke & Ellis, Grevillea 5
(34): 50 (1876)
: Spondylocladium obovatum (Cooke & Ellis) S.
Hughes, Can. J. Bot. 31 (5): 634 (1953)
Distribution: China, Hong Kong, on submerged wood
(Zhuang 2001).
Sexual morph: Undetermined
517
Notes: Sequence data is unavailable.
Torrentispora Hyde et al., Mycol. Res. 104(11): 1399
(2000)
Asexual morph Undetermined. Sexual morph Updated
description refer to Réblová et al. (2018).
Type species: Torrentispora fibrosa Hyde et al., Mycol.
Res. 104(11): 1399 (2000)
Notes: Hyde et al. (2000) introduced the genus Torrentispora with single species, T. fibrosa. Réblová et al.
(2018) revisited Torrentispora with nine species in this
genus and assigned it to the new family Xenospadicoidaceae within the new order Xenospadicoidales based
on morphology and DNA sequence data. Torrentispora
comprises seven species from freshwater habitats.
Torrentispora aquatica (Vijaykr. et al.) Réblová & A.N.
Mill
: Fusoidispora aquatica Vijaykr. et al., Sydowia
57:272 (2005)
Distribution: China, Hong Kong, New Territories, Tai
Po Kau Country Park, Tai Po Kau Forest Stream, on submerged wood (Vijaykrishna et al. 2005).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 17484 (now in IFRD), isotype PDD 78746. LSU sequence data is available.
Torrentispora biatriispora (K.D. Hyde) Réblová & A.N.
Mill.
: Annulatascus biatriisporus K.D. Hyde, Nova Hedwigia 61: 120 (1995)
: Pseudoannulatascus biatriisporus (K.D. Hyde) Z.L.
Luo et al., Phytotaxa 239:179 (2015)
Distribution: Australia, on submerged wood (Hyde
1995b); China, Hong Kong, Tsuen Wan, Shing Mun
Reservoir, on submerged wood (Tsui et al. 2002); Costa
Rica, on submerged wood (Barbosa et al. 2013); Seychelles, on submerged wood (Hyde and Goh 1998a).
Asexual morph: Undetermined
Notes: Holotype BRIP 21481, other specimen collected
from freshwater habitats: ILL 40816. ITS, LSU, SSU and
RPB2 sequence data are available.
Torrentispora calembola Réblová & A.N. Mill.
Distribution: France, on submerged wood of Fraxinus
excelsior (Réblová et al. 2018).
Asexual morph: Undetermined
Notes: Holotype PRA-12744 (M.R. 3726), isotype PRA12745. ITS, LSU and SSU sequence data are available.
Torrentispora crassiparietis Fryar & K.D. Hyde
Distribution: Brazil, on submerged bark (Barbosa et al.
2013); Brunei, on submerged wood (Fryar and Hyde 2004;
Fryar et al. 2004); Costa Rica, on submerged wood (Barbosa et al. 2013).
Asexual morph: Undetermined
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Fungal Diversity (2019) 99:451–660
b Fig. 26 Phylogram generated from maximum likelihood analysis
based on combined ITS, LSU and RPB2 sequence data for species of
Barbatosphaeria (with Natantiella ligneola as outgroup). The best
scoring RAxML tree with a final likelihood value of - 9327.825160
is presented. RAxML bootstrap support values equal to or greater than
75% are given before the forward slash (black). Maximum parsimony
bootstrap support values equal to or greater than 75% are given after
the forward slash (red). Hyphen (‘-’) indicates a value lower than 75%
for RAxML and maximum parsimony. Newly generated sequences
are in red. Ex-type strains are in bold
Notes: Holotype IFRD 8863, other specimens collected
from freshwater habitats: HUEFS 158094, ILL 40817, ILL
40818, ILL 40819, ILL 40820. Sequence data is
unavailable.
Torrentispora fibrosa Hyde et al.
Distribution: China, Hong Kong, New Territories, Tai
Po, Lam Tsuen River and Tai Po Kau Forest Reserve, Tai
Po Kau Forest Stream, on submerged decaying wood
(Hyde et al. 2000); New Zealand, Lake Christabel track,
on decaying wood of Nothofagus sp. (Réblová et al. 2018).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 4519 (now in IFRD), other
specimen collected from freshwater habitats PDD 110879.
LSU, SSU and RPB2 sequence data are available.
Torrentispora fusiformis Fryar & K.D. Hyde
Distribution: Brunei, on submerged wood (Fryar and
Hyde 2004).
Asexual morph: Undetermined
Notes: Holotype IFRD 8864. Sequence data is not
available. This species is known only from Brunei on
submerged wood in brackish and freshwater habitats (Fryar
and Hyde 2004).
Torrentispora pilosa Shearer & F.R. Barbosa
Distribution: Costa Rica, on submerged wood (Barbosa
et al. 2013).
Asexual morph: Undetermined
Notes: Holotype ILL 40814. Sequence data is not
available. This species is known only from Costa Rica, on
submerged wood (Barbosa et al. 2013).
Diaporthomycetidae family incertae sedis
Acrodictyaceae J.W. Xia & X.G. Zhang
Acrodictys M.B. Ellis, Mycol. Pap. 79: 6 (1961)
Asexual morph Descriptions and illustrations refer to Xia
et al. (2017). Sexual morph Undetermined.
Type species. Acrodictys bambusicola M.B. Ellis,
Mycol. Pap. 79: 6 (1961)
Notes: Acrodictys was introduced by Ellis (1961), with
A. bambusicola M.B. Ellis as the type species. There are 45
epithets listed in Index Fungorum (December 2018). Until
2016, identification of Acrodictys-like species was based
on morphology. Xia et al. (2017) provided ITS, LSU, SSU
519
and b-tubulin sequence data, detailed descriptions and
illustrations for eight Acrodictys species, including a new
species. In this study, we introduce a new species and a
new combination for this genus. Three Acrodictys species
have been reported from freshwater habitats.
Acrodictys aquatica (N.G. Liu & K.D. Hyde) Z.L. Luo &
K.D. Hyde, comb. nov.
: Barbatosphaeria aquatica N.G. Liu & K.D. Hyde, in
Hyde et al., Mycosphere 9(2): 384 (2018)
Index Fungorum: IF 655656; Facesoffungi number: FoF
05438
Descriptions and illustrations refer to Hyde et al. (2018).
Distribution: Thailand, Chiang Rai Province, Muang,
Ban Nang Lae Nai, on decaying wood submerged in a
freshwater stream (Hyde et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0040. ITS and LSU
sequence data are available. Hyde et al. (2018) introduced
the species Barbatosphaeria aquatica based on a collection
obtained from freshwater habitat in Thailand. Our phylogenetic analysis based on multi-genes shows that the strain
from ex-type culture of Barbatosphaeria aquatica clusters
in the genus Acrodictys (Fig. 1, clade 9). Morphology of
Barbatosphaeria aquatica also fits well with Acrodictys
species in having mononematous, erect, unbranched conidiophores; monoblastic, terminal, integrated conidiogenous
cells; solitary, dry, clavate, green to brown conidia, with
transverse septa, deeply constricted at the septa, distinctly
verruculose (Hyde et al. 2018). Based on morphology and
phylogenetic analyses, we synonymize Barbatosphaeria
aquatica under Acrodictys aquatica.
Acrodictys fluminicola Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555657, Facesoffungi
number: FoF 05429, Fig. 24
Etymology: Referring to this fungus dwelling in a
stream.
Holotype: DLU 0274
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, dark brown to
black. Conidiophores 98–142 lm long, 4–6 lm wide
(
x = 120 9 5 lm, n = 20), macronematous, mononematous, erect, unbranched, straight or flexuous, septate, dark
brown at the base, narrower and paler toward the apex,
smooth, thick-walled. Conidiogenous cells monoblastic,
integrated, terminal, cylindrical, lageniform to doliiform,
subhyaline to pale brown, smooth. Conidia 24–30 lm
long, 13–17 lm wide (
x = 27 9 15 lm, n = 20), acrogenous, solitary, muriform, broadly clavate, obovoid to
pyriform, usually with 2–3 transverse septa and a few
longitudinal septa, with conspicuous pores in each cells,
slightly constricted at the septa, pale brown at the basal
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Fig. 27 Sporidesmiella aquatica (MFLU 18–1602, holotype) a Colony on natural substrate. b, c Conidiophore with conidia. d, e Conidiogenous
cells with conidia. f–h Conidia. i Germinating conidium. Culture on PDA from above (j) and reverse (k). Scale bars: b, c 100 lm, d–i 50 lm
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521
Fig. 28 Sporidesmiella novae-zelandiae (MFLU 18–1604) a Colony on natural substrate. b, c Conidiophore with conidia. d, e Conidiogenous
cells with conidia. f–i Conidia. j Germinating conidium. Culture on PDA from surface (k) and reverse (l). Scale bars: b, c 30 lm, d–i 15 lm
123
522
cells and brown at the other part, basal cell obconical,
truncate at base. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a
freshwater stream, October 2014, L.W. Wang, S-274 (DLU
0274, holotype), ex-type living culture KUMCC 15–0240.
Notes: Acrodictys fluminicola resembles the generic
type A. bambusicola in having macronematous, unbranched and septate conidiophores dark brown at the base,
narrower and paler toward the apex, solitary, muriform,
obovoid to pyriform conidia slightly constricted at the
septa, with obconical basal cell and the size of conidiophores and conidia of these two species are almost similar
(Xia et al. 2017). However, A. fluminicola differs from A.
bambusicola in has 2–3 transverse septa conidia with
conspicuous guttulae in each cell, while A. bambusicola
have 3–4 transverse septa. Phylogenetic analysis also
shows that Acrodictys fluminicola and A. bambusicola are
distinct species (Fig. 1, clade 9). Phylogenetically, A.
aquatica clusters with A. hainnensis (Fig. 1, clade 9),
however, Acrodictys fluminicola morphologically differs
from A. hainnensis in having longer conidiophores (98–142
vs. up to 35 lm) and larger conidia (24–30 9 13–17 vs.
15–22 9 7–13 lm).
Acrodictys liputii Cai et al.
Distribution: Philippines, Negros Occidental, Bario
Alegria, Liput River, on submerged bamboo culm (Cai
et al. 2002b).
Sexual morph: Undetermined
Notes: Holotype PDD 74789, isotype IFRD 8640. ITS,
LSU and SSU sequence data are available.
Barbatosphaeriaceae Zhang et al.
Barbatosphaeria Réblová, Mycologia 99(5): 727 (2008)
[2007]
Asexual morph Undetermined. Sexual morph Description
and illustrations refer to Réblová et al. (2015a).
Type species: Barbatosphaeria barbirostris (Dufour)
Réblová, Mycologia 99(5): 727 (2008)
Notes: Barbatosphaeria is a non-stromatic perithecial
ascomycete introduced for a single species Barbatosphaeria barbirostris and placed in the Sordariomycetes genera incertae sedis based on nuc28S rDNA
sequences (Réblová 2007). Réblová et al. (2015a) revisited
Barbatosphaeria and introduced seven species for this
genus. In this study, we introduce one new species and this
is the only known Barbatosphaeria species collected from
freshwater habitats.
Barbatosphaeria lignicola Z.L. Luo, H.Y. Su & K.D.
Hyde, sp. nov.
Index Fungorum number: IF 555658, Facesoffungi
number: FoF 05430, Fig. 25
Etymology: Referring to this fungus dwelling on wood
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Holotype: HKAS 84005
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata immersed, globose to subglobose, dark brown to
black, 380–400 lm high, 270–320 lm diam., aggregated
in circular groups, occasionally solitary. Neck black,
cylindrical, straight to slightly flexuous, converging radially. Peridium 25–40 lm thick, leathery to fragile, 2-layered. Outer layer consisting of thick-walled, brown,
polyhedral to elongate cells of textura prismatica to textura
angularis, towards the interior grading into several layers of
thin-walled pale brown to subhyaline flattened cells. Paraphyses septate, slightly constricted at the septa, wider
near the base, tapering toward the tip, longer than asci. Asci
19–31 9 4–6 lm (
x = 25 9 5 lm, n = 30), 8-spored,
unitunicate, clavate in sporiferous part, tapering toward the
stipe, ascal apex broadly rounded to obtuse. Ascospores 5–
7 9 1–2 lm (
x = 6 9 1.5 lm, n = 20), arranged obliquely
1–2-seriately in the sporiferous part of the ascus, allantoid,
U- to horseshoe shaped, narrow to pinted at both ends,
aseptate, non-constricted, hyaline.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in
Heilong stream, 15 March 2014, Z.L. Luo, S-105, (HKAS
84005, holotype).
Notes: Barbatosphaeria lignicola mostly resembles B.
hippocrepida in having globose to subglobose, dark brown
to black ascomata, black, cylindrical neck, septate paraphyses wider near the base, tapering toward the tip, longer
than asci and slightly constricted at the septa, unitunicate,
8-spored asci and allantoid, U- to horseshoe shaped, hyaline, aseptate, non-constricted, similar size of ascospores
(Réblová and Štěpánek 2009). However, B. lignicola differs from B. hippocrepida in having smaller ascomata
(380–400 9 270–320
vs.
350–500 9 400–650 lm),
ascospores tending to group in the apex of the sporiferous
part while ascospores of B. hippocrepida tend to group in
the middle of the sporiferous part, leaving the apex empty.
Phylogenetic analyses show that Barbatosphaeria lignicola
is distinct from other Barbatosphaeria species (Fig. 26).
Conlariaceae Zhang et al.
Conlarium F. Liu & L. Cai, Mycologia 104(5): 1180
(2012)
Asexual morph Hyphomycetous, for description and
illustration refer to Zhang et al. (2017a, b). Sexual morph
Description and illustration refer to Liu et al. (2012).
Type species: Conlarium dupliciascospora F. Liu & L.
Cai, Mycologia 104(5): 1180 (2012)
Notes: Currently, two species are accepted in the genus
Conlarium, and both C. aquaticum and C. dupliciascospora
are reported from freshwater habitats in China and Thailand respectively (Liu et al. 2012; Zhang et al. 2017a, b).
Fungal Diversity (2019) 99:451–660
523
Fig. 29 Sporidesmiella hyalosperma (MFLU 18–1603) a Colony on natural substrate. b Conidiophores with conidia. c Conidiogenous cells. d,
e Conidia. f, g Germinating conidia. Culture on PDA from above (h) and reverse (i). Scale bars: b 50 lm, c–g 20 lm
Conlarium aquaticum Dong et al.
Distribution: Thailand, Prachuap Khiri Khan, on submerged wood in a small River (Zhang et al. 2017).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–2703. ITS, LSU, SSU and
TEF1a sequence data obtained from ex-type culture are
available.
Conlarium dupliciascosporum F. Liu & L. Cai
Distribution: China, Guangdong Province, Zhaoqing,
Dinghu Mountain, on submerged wood in a stream (Liu
et al. 2012).
Asexual morph: See Liu et al. (2012)
Notes: Holotype HMAS 243129. ITS, LSU and SSU
sequence
data
are
available,
ex-type
strain
CGMCC3.14938.
Riomyces Ferrer et al., Mycologia 104(4): 876 (2012)
Asexual morph Undetermined. Sexual morph Descriptions and illustration refer to Ferrer et al. (2012).
Type species: Riomyces rotundus A. Ferrer, A.N. Mill.,
Sarmiento & Shearer, Mycologia 104(4): 876 (2012)
Notes: There is only one species was accepted in the
genus Riomyces and was only known from the type
locality.
Riomyces rotundus Ferrer et al.
Distribution: Costa Rica, Alajuela and Heredia, on
submerged wood (Ferrer et al. 2012).
Asexual morph: Undetermined
Notes: Holotype ILL AF303–1, other specimen collected from freshwater habitats: ILL AF303–3. LSU and
SSU sequence data are available.
Junewangiaceae J.W. Xia & X.G. Zhang
Dictyosporella Abdel-Aziz, Fungal Divers 75: 144 (2015)
Asexual morph Description and illustrations see Ariyawansa et al. (2015). Sexual morph Description and illustrations see Zhang et al. (2017a, b).
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Fungal Diversity (2019) 99:451–660
b Fig. 30 Fluminicola thailandensis (MFLU 15–0085) a Specimen. b,
c Appearance of necks on substrate. d Vertical section of ascomata.
e Structure of peridium. f Paraphyses. g–i Ascus. j–m Ascospores.
n Germinating ascospore. o Colony on PDA. Scale bars: d, g–
i 40 lm, e, f, n 20 lm, j–m 10 lm
Type species: Dictyosporella aquatica Abdel-Aziz,
Fungal Divers 75: 145 (2015)
Notes: Ariyawansa et al. (2015) introduced the genus
Dictyosporella to accommodate the asexual morph species
D. aquatica Abdel-Aziz, collected on submerged decaying
stems of Phragmites australis in Nile River, Egypt, and
accommodated it in family Annulatascaceae. Zhang et al.
(2017a, b) introduced a sexual morph species Dictyosporella thailandensis W. Dong, H. Zhang & K.D. Hyde
for this genus which was collected from freshwater, and
they moved this genus from Annulatascaceae to Diaporthomycetidae genera incertae sedis. Song et al. (2018a)
introduced the third species Dictyosporella hydei H.Y.
Song & D.M. Hu based on morphology and phylogenetic
analyses, but the placement of this genus was not mentioned in their study. Our phylogenetic analysis based on
multi-gene shows that the genus Dictyosporella clusters in
Junewangiaceae with strong support (Fig. 1, clade 1).
According to our phylogenetic result, Dictyosporella is
transferred from Diaporthomycetidae genera incertae sedis
to Junewangiaceae. Three species were accepted in this
genus and all were collected from freshwater habitats.
Dictyosporella aquatica Abdel-Aziz
Distribution: Egypt, Sohag, River Nile, on submerged
decayed stem of Phragmites australis (Ariyawansa et al.
2015).
Sexual morph: Undetermined
Notes: Holotype CBS H-22127, isotype MFLU
15–1510. LSU and SSU sequence data are available.
Dictyosporella hydei H.Y. Song & D.M. Hu
Distribution: China, Yunnan Province, Mengla, on
submerged wood in a small stream (Song et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype IFRD 021–044. LSU sequence data is
available. This species is only known from the type
locality.
Dictyosporella thailandensis Dong et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged bamboo in a small River (Zhang et al.
2017a, b).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2706. ITS, LSU, SSU and
RPB2 sequence data are available.
Junewangia W.A. Baker & Morgan-Jones, Mycotaxon 81:
307 (2002)
525
Asexual morph Description and illustrations see Xia et al.
(2018). Sexual morph Undetermined.
Type species: Junewangia sphaerospora W.A. Baker &
Morgan-Jones, Mycotaxon 81: 312 (2002)
Notes: Junewangia was established by Baker et al.
(2002) to accommodate four species of Acrodictys M.B.
Ellis. The species of Junewangia are characterized by
slightly flared conidiophores with annellidic, percurrent
proliferation, cylindrical or narrowly cuneate conidiogenous cells, subglobose to globose conidia with a
prominent, protruding basal cell, uniformly pigmented, and
a series of largely angular or oblique septa, with schizolytic
secession. Seven species are accepted in the genus
Junewangia and only one of them was collected from
freshwater habitats in China (Song et al. 2018b).
Junewangia aquatica H.Y. Song & D.M. Hu
Distribution: China, Yunnan Province, Mengla, stream
in rubber trees field, on submerged wood (Song et al.
2018b).
Sexual morph: Undetermined
Notes: Holotype HFJAU 0700; ITS, LSU, SSU
sequence data are available. This species is only known
from the type locality.
Sporidesmiella P.M. Kirk
Asexual morph Descriptions and illustrations see Wu and
Zhuang (2005); Sexual morph Undetermined.
Type species: Sporidesmiella claviformis P.M. Kirk,
Trans. Br. Mycol. Soc. 79: 479 (1982)
Notes: Kirk (1982) introduced the genus Sporidesmiella
P.M. Kirk to accommodate the species previously described in the genera Sporidesmium and Endophragmia.
Sporidesmiella resembles Repetophragma but differs in its
distoseptate conidia (Kirk 1982; Subramanian 1992). In our
phylogenetic analysis, Sporidesmiella clusters in
Junewangiaceae while Repetophragma belongs in Pseudosporidesmiaceae (Fig. 1, clade 1, 37). In this study, we
report three Sporidesmiella species which were collected
from freshwater habitats with one new taxon.
Sporidesmiella aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555659, Facesoffungi
number: FoF 05431, Fig. 27
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1602
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, hairy, yellow
brown to brown. Mycelium partly superficial, partly
immersed, composed of septate, branched, smooth, pale
brown hyphae. Conidiophores 178–228 lm long, 8–10 lm
wide
(
x = 203 9 9 lm,
n = 20),
macronematous,
mononematous, erect, unbranched, septate, straight or
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Fig. 31 Wongia aquatica (MFLU 18–1596, holotype) a–c Conidiophore with conidia. d Conidiogenous cells. e–i Conidia. j Germinating
conidium. Scale bars: a–c 45 lm, d–j 20 lm
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flexuous, cylindrical, yellow–brown, paler towards the
apex, smooth. Conidiogenous cells holoblastic, polyblastic,
sympodial, integrated, terminal, subterminal, subhyaline to
pale brown, with percurrent or sympodial proliferations.
Conidia
51–59 lm
long,
18–22 lm
wide
(
x = 55 9 20 lm, n = 20), acropleurogenous, dry, clavate
or obovoid, rounded at the apex, truncate at the base, 3–4distoseptate, pale brown, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a
freshwater stream, October 2016, Z.L. Luo, S-777 (MFLU
18–1602, holotype), ex-type living culture DLUCC 0777;
Laojunshan Mountain, saprobic on decaying wood submerged in a freshwater stream, August 2017, H.W. Shen,
S-1339 (MFLU 18–2331, paratype), living culture DLUCC
1339.
Notes: Sporidesmiella aquatica resembles S. hyalosperma in having macronematous, mononematous, erect,
unbranched, septate, cylindrical conidiophores, integrated,
terminal conidiogenous cells with percurrent or sympodial
proliferations and acrogenous, dry, clavate or obovoid
conidia (Kirk 1982; Wu and Zhuang 2005). However, S.
aquatica differs from S. hyalosperma in having polyblastic
conidiogenous cells and larger conidia (51–59 9 18–22 vs.
22–25 9 9–12 lm).
Sporidesmiella novae-zelandiae (S. Hughes) Madrid et al.
: Sporidesmium hyalospermum var. novae-zelandiae
S. Hughes, N.Z. Jl Bot. 16(3): 349 (1978)
: Sporidesmiella hyalosperma var. novae-zelandiae (S.
Hughes) P.M. Kirk, Trans. Brit. Mycol. Soc. 79: 479
(1982)
: Exserohilum novae-zelandiae (S. Hughes) H.P.
Upadhyay & Mankau, Mycologia 83: 373 (1991)
Facesoffungi number: FoF 05432, Fig. 28
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, hairy, yellow
brown to brown. Mycelium partly superficial, partly
immersed, composed of septate, branched, smooth, pale
brown hyphae. Conidiophores 132–186 lm long, 5–7 lm
wide
(
x = 159 9 6 lm,
n = 20),
macronematous,
mononematous, erect, unbranched, septate, straight or
flexuous, cylindrical, yellow–brown, paler towards the
apex, smooth. Conidiogenous cells holoblastic, polyblastic,
sympodial, integrated, terminal, subhyaline to pale brown.
Conidia
25–29 lm
long,
12–14 lm
wide
(
x = 27 9 13 lm, n = 20), acrogenous, dry, pale brown,
clavate or obovoid, rounded at the apex, truncate at the
base, 2–4-distoseptate, basal cell cut off by a dark brown
septum, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on decaying wood submerged, October
2016, Z.L. Luo, S-951 (MFLU 18–1604), living culture
527
DLUCC 0951; Laojunshan Mountain, saprobic on decaying wood submerged in a freshwater stream, August 2017,
H.W. Shen, S-1256 (MFLU 18–2332), living culture
DLUCC 1256.
Notes: Holotype PDD 30420, isotype DAOM 159962,
other specimen collected from freshwater habitats: MFLU
18–1604. We provide the ITS, LSU, SSU, RPB2 and
TEF1a sequence data for this species. The species Sporidesmiella novae-zelandiae was originally described as
Sporidesmium hyalospermum var. novae-zelandiae by
Hughes (1978). This species and the type variety were
reallocated to Sporidesmiella by Kirk (1982) and Sporidesmium hyalospermum var. novae-zelandiae was later
transferred to Exserohilum as E. novae-zelandiae by
Upadhyay and Mankau (1991). Hernández-Restrepo et al.
(2018) considered that Exserohilum novaezelandiae should
be retained in Sporidesmiella and combined it as S. novaezelandiae. During an investigation on lignicolous freshwater fungi in China, a Sporidesmiella-like fungus was
collected from Yunnan Province and the morphological
characters of our new collection fits well with S. novaezelandiae (Hughes 1978; Kirk 1982; Wu and Zhuang 2005;
Hernández-Restrepo et al. 2018). Phylogenetic analysis
shows that our isolates cluster with other Sporidesmiella
species with good bootstrap support value and form in a
distinct clade (Fig. 1, clade 1). Based on the morphology
and phylogeny, we identify our isolates as Sporidesmiella
novae-zelandiae.
Sporidesmiella hyalosperma (Corda) P.M. Kirk
: Helminthosporium hyalospermum Corda, Icon. fung.
(Prague) 1: 13 (1837)
Facesoffungi number: FoF 05433, Fig. 29
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, hairy, yellow
brown to brown. Mycelium partly superficial, partly
immersed, composed of septate, branched, smooth, pale
brown hyphae. Conidiophores 116–144 lm long, 5–7 lm
wide
(
x = 130 9 6 lm,
n = 20),
macronematous,
mononematous, erect, unbranched, septate, straight or
flexuous, cylindrical, brown, paler towards the apex,
smooth. Conidiogenous cells holoblastic, polyblastic,
sympodial, integrated, terminal, later becoming subterminal, subhyaline to pale brown, with percurrent or sympodial proliferations. Conidia 17–21 lm long, 8–10 lm wide
(
x = 19 9 9 lm, n = 20), acropleurogenous, dry, clavate
or obovoid, rounded at the apex, truncate at the base, 3–4distoseptate, pale brown, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Jinsha
River, saprobic on decaying wood submerged, April 2015,
Z.L. Luo, S-563 (MFLU 18–1603, HKAS 92987), living
123
528
culture KUMCC 15–0431; Cangshan Mountain, saprobic
on decaying wood submerged in a freshwater stream,
August 2017, H.W. Shen, S-1400, living culture MFLUCC
18–1312; Ibid., saprobic on decaying wood submerged in a
freshwater stream, August 2017, H.W. Shen, S-1518
(MFLU 18–2330), living culture DLUCC 1518.
Notes: Holotype PDD 30420, specimen collected from
freshwater habitats: MFLU 18–1603. We provide the ITS,
LSU, SSU, RPB2 and TEF1a sequence data for this species
in this study. Sporidesmiella hyalosperma is the most
common species in the genus Sporidesmiella. Morphologically, our isolates fit well with the Sporidesmiella hyalosperma in having macronematous, mononematous,
solitary, erect, unbranched, septate, straight or flexuous,
cylindrical, brown, smooth conidiophores paler towards the
apex, holoblastic, integrated, terminal, subhyaline to pale
brown conidiogenous cells with percurrent or sympodial
proliferations and acrogenous, dry, clavate or obovoid,
distoseptate conidia rounded at the apex, truncate at the
base and even the size of conidiophores and conidia are
also similar (Kirk 1982; Wu and Zhuang 2005). Therefore,
we identify our collections as Sporidesmiella hyalosperma,
and it is the first time to report this species from freshwater.
Papulosaceae Winka & O.E. Erikss.
Brunneosporella Ranghoo & K.D. Hyde, Mycol. Res.
105(5): 625 (2001)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Ranghoo et al. (2001).
Type species: Brunneosporella aquatica Ranghoo &
K.D. Hyde, Mycol. Res. 105(5): 625 (2001)
Notes: Ranghoo et al. (2001) introduced the genus
Brunneosporella with single species B. aquatica which was
collected from freshwater habitats. This species is only
known from the type locality.
Brunneosporella aquatica Ranghoo & K.D. Hyde
Distribution: China, Hong Kong, New Territories, Plover Cove Reservoir, on submerged wood (Ranghoo et al.
2001).
Asexual morph: Undetermined
Notes: Holotype IFRD 8717. ITS and LSU sequence
data are available.
Fluminicola Wong et al., Fungal Diversity Res. Ser. 2: 190
(1999)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Wong et al. (1999b) and
Zhang et al. (2017a, b).
Type species: Fluminicola bipolaris Wong et al., Fungal
Divers 2: 190 (1999)
Notes: Wong et al. (1999b) established the genus Fluminicola with F. bipolaris as type species. Zhang et al.
(2017a, b) introduced three species which were collected
from a freshwater River in southern Thailand.
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Fungal Diversity (2019) 99:451–660
Fluminicola aquatica Dong et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged wood in a small River (Zhang et al.
2017a, b).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2710. ITS, LSU and SSU
sequence data are available.
Fluminicola bipolaris Wong et al.
Distribution: Philippines, Mindanao, Bukidnon, Impalutao, Natigbasan Creek, on submerged wood (Wong et al.
1999b).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 3127 (now in IFRD).
Sequence data is not available.
Fluminicola saprophytica Dong et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged bamboo in a small River (Zhang et al.
2017a, b).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2694. ITS, LSU, SSU and
RPB2 sequence data are available.
Fluminicola thailandensis Dong et al.
Facesoffungi number: FoF 03346, Fig. 30
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 138–156 lm high, 100–120 lm diam.,
immersed with neck erumpent through host surface, subglobose to ellipsoid, solitary, dark brown to black. Ostiole
central, with straight upright, black neck. Peridium 12–
18 lm thick, consisting two-layers, outer layer comprising
3–4 layers of dark brown, thick-walled cells of textura
angularis, inner layer comprising 2–3 layers of pale brown
to hyaline, thin-walled cells of textura angularis. Paraphyses 4–6 lm wide, hyaline, unbranched, septate, slightly
constricted at the septum. Asci 135–181 9 11–15 lm
(
x = 158 9 13 lm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, tapering to a point, apically rounded or
slightly obtuse, with an indistinct, refractive, small, discoid
apical ring. Ascospores 19–21 9 7–9 lm (
x = 20 9 8 lm,
n = 20), uniseriate, fusiform, 3-septate, slightly constricted
at the septa, with 2 prominent guttules, with a thin
mucilaginous sheath, hyaline, smooth-walled.
Material examined: THAILAND, Chiang Rai Province,
saprobic on decaying wood submerged in a freshwater
stream, November 2013, Z.L. Luo, ZL-10 (MFLU
15–0085), living culture MFLUCC 14–0037.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged wood in a small River (Zhang et al.
2017a, b), Chiang Rai Province, saprobic on decaying
wood submerged in a freshwater stream (This study).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2704, other specimen collected from freshwater MFLU 15–0085. ITS, LSU, SSU
Fungal Diversity (2019) 99:451–660
and TEF1a sequence data are available. Fluminicola
thailandensis was introduced by Zhang et al. (2017a, b)
based on an isolate collected from southern Thailand.
During our investigation of lignicolous freshwater fungi in
northern Thailand, an Annulatascaceae-like fungus was
collected from Chiang Rai Province. Morphologically, the
new isolate resembles Fluminicola species in having
immersed ascomata with black neck, cylindrical asci with
apical ring and fusiform, septate, guttulate ascospores with
a thin mucilaginous sheath. Phylogenetic analysis based on
LSU, SSU, RPB2 and TEF1a sequence data shows that the
new strain (MFLUCC 14–0037) clusters with Fluminicola
species with good support, and close to F. thailandensis
and F. saprotrophitica. We compared the ITS sequences
and noted that there are 3 nucleotide differences (including
2 gaps) between the ex-type strain of Fluminicola thailandensis (MFLUCC 15–0984) and our strain (MFLUCC
14–0037). Based on morphology and phylogeny, we
identify our species as Fluminicola thailandensis.
Wongia Khemmuk et al., IMA Fungus 7(2): 249 (2016)
Asexual morph Colonies effuse, scattered, hairy, dark
brown. Mycelium partly immersed, composed of branched,
septate, brown, smooth hyphae. Conidiophores macronematous, mononematous, solitary, erect, straight or flexuous, unbranched, septate, dark brown, smooth.
Conidiogenous cells polyblastic, denticulate, integrated,
terminal, subhyaline. Conidia acrogenous, clavate to fusiform, 3-septate, guttulate, dark brown at central two cells,
paler at end cells, rounded and narrow at apex, truncate at
base. Sexual morph Descriptions refer to Khemmuk et al.
(2016).
Type species: Wongia garrettii (P. Wong & M.L.
Dickinson) Khemmuk et al., IMA Fungus 7(2): 250 (2016)
Notes: Khemmuk et al. (2016) re-examined Magnaporthe garrettii and M. griffinii that are pathogenic on roots
of couch and hybrid couch (Wong et al. 2012) and established the genus Wongia. Wongia is the fourth genus to be
placed in Papulosaceae, along with Brunneosporella
(Ranghoo et al. 2001), Fluminicola (Wong et al. 1999a, b)
and Papulosa (Kohlmeyer and Volkmann-Kohlmeyer
1993). In this study, we introduce an asexual Wongia
species which was collected from freshwater habitats in
northwestern Yunnan Province, China.
Wongia aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.
Index Fungorum number: IF 555660, Facesoffungi
number: FoF 05434, Fig. 31
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1596
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, scattered, hairy, dark brown.
Mycelium partly immersed, composed of branched, septate,
529
brown, smooth hyphae. Conidiophores 54–90 lm long, 3–
6 lm wide (
x = 72 9 4 lm, n = 20), macronematous,
mononematous, solitary, erect, straight or flexuous,
unbranched, septate, dark brown, smooth. Conidiogenous
cells polyblastic, sympodial, denticulate, integrated, terminal, subhyaline, with pale brown scar. Conidia 17–
21 lm long, 5–7 lm wide (
x = 19 9 6 lm, n = 20),
acropleurogenous, fusiform, guttulate, 3-septate, dark
brown at central two cells, paler at end cells, rounded and
narrow at apex, truncate at base, smooth-walled. Sexual
morph Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on decaying wood submerged, October
2016, Z.L. Luo, S-912 (MFLU 18–1596, holotype), ex-type
living culture MFLUCC 18–1607.
Notes: There are no asexual morphs reported for Wongia
garrettii and W. griffinii. Wongia aquatica is the first
asexual morph species in the genus Wongia. Phylogenetically, W. aquatica shares a sister relationship to W. griffinii
with high bootstrap support (96% ML) (Fig. 1, clade 11).
Pseudoproboscisporaceae Zhang et al.
Diluviicola Hyde et al., Fungal Diversity Res. Ser. 1: 141
(1998)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Hyde et al. (1998b).
Type species: Diluvicola capensis K.D. Hyde, S.W.
Wong & E.B.G. Jones, Fungal Divers 1: 134 (1998)
Notes: Hyde et al. (1998b) introduced the genus Diluviicola based on a collection from Brunei. Zhang et al.
(2017a, b) introduced the second species in this genus,
Diluviicola aquatica W. Dong, H. Zhang & K.D. Hyde,
based on a collection made from submerged bamboo in a
river in Thailand.
Diluviicola aquatica Dong et al.
Distribution: Thailand, Prachuap Khiri Khan, on submerged bamboo in a small River (Zhang et al. 2017a, b).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2701. ITS, LSU, SSU and
RPB2 sequence data obtained from ex-type culture are
available.
Diluviicola capensis Hyde et al.
Distribution: Brunei, Temburong, Kuala Belalong Field
Studies Centre, Sungai Esu, on submerged wood (Hyde
et al. 1998c).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 3125 (now in IFRD).
Sequence data is unavailable.
Pseudoproboscispora Punith., Kew Bull. 54(1): 234 (1999)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Wong and Hyde (1999a, b).
Type species: Pseudoproboscispora aquatica (S.W.
Wong & K.D. Hyde) Punith., Kew Bull. 54(1): 234 (1999)
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Fungal Diversity (2019) 99:451–660
Fig. 32 Rhodoveronaea aquatica (MFLU 18–1593, holotype). a, b Conidiophores with conidia. c, d Conidiogenous cells with conidia. e–
o Conidia. Scale bars: a–c 50 lm, d–o 10 lm
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Fungal Diversity (2019) 99:451–660
Notes: Index Fungorum (2018) lists three species in the
genus Pseudoproboscispora. Previous studies showed that
the species of Pseudoproboscispora are distributed
worldwide and all of them were reported from freshwater
habitats (Wong and Hyde 1999a, b; Campbell et al. 2003a;
Zhang et al. 2017a, b).
Pseudoproboscispora aquatica (S.W. Wong & K.D. Hyde)
Punith.
: Proboscispora aquatica S.W. Wong & K.D. Hyde,
Mycol. Res. 103: 82 (1999)
Distribution: Australia, North Queensland, near Cairns,
Crystal Cascades, on submerged wood; Philippines,
Negros Occidental, Bacolod, Kaliban River, on submerged
wood (Wong and Hyde 1999a, b). Japan, Koito River, on
submerged wood (Tsui et al. 2001a)
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2792 (now in IFRD).
Sequence data is not available.
Pseudoproboscispora caudae-suis (Ingold) J. Campb. et al.
: Ceriospora caudae-suis Ingold, Trans. Br. Mycol.
Soc. 34: 210 (1951)
Distribution: UK, Westmorland, Lake Windermere, on
submerged, decorticated twigs of Fraxinus; USA, Arkansas, Lee County, Florida, Hamilton County, Illinois,
Johnson County, Lousiana, Caldwell Parish, Mississippi,
Franklin County, New York, Adirondack Park, North
Carolina, Cheoah River, Oregon, Florence County, Pennsylvania, Columbia County, Tennessee, Great Smoky
Mountains National Park, West Virginia, Pocahontas
County, Wisconsin, Adams County, on submerged wood in
lentic or lotic habitats (Campbell et al. 2003a).
Asexual morph: Undetermined
Notes: Holotype K(M) 109308; Sequence data is not
available.
Pseudoproboscispora thailandensis Dong et al.
Distribution: Thailand, Prachuap Khiri Khan, on submerged bamboo in a small River (Zhang et al. 2017a, b).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–2705. ITS, LSU and SSU
sequence data are available.
Rhamphoriaceae Réblová
Rhamphoria Niessl, Verh. nat. Ver. Brünn 14: 204 (1876)
Asexual morph Hyphomycetous (Phaeoisaria-like). Sexual morph Ascomata partly immersed in the wood, black,
single or sometimes in pairs, flask-shaped with an ostiolar
neck or depressed cup shape with the neck arising sideways. Peridium composed of thick-walled, dark brown
polygonal cells which become thin-walled and hyaline
towards interior. Paraphyses filiform, hyaline and numerous. Asci 8-spored, unitunicate, cylindrical, long-stalked.
Ascospores monostichous in the ascus, ovoid to oblong,
hyaline to subhyaline, smooth-walled.
531
Type species: Rhamphoria delicatula Niessl, Verh. nat.
Ver. Brünn 14: 206 (1876)
Notes: 15 Rhamphoria epithes are listed in Index Fungorum (December 2018) and only one species was reported
from freshwater habitats (Ho et al. 2001).
Rhamphoria pyriformis (Pers.) Höhn
: Sphaeria pyriformis Pers., Syn. meth. fung. (Göttingen) 1: 64 (1801)
Distribution: China, Hong Kong, Tai Po Kau Forest, on
submerged wood in a stream (Ho et al. 2001)
Asexual morph: Undetermined
Notes: Herbarium records CBS H-115, CBS H-17785.
ITS, LSU and RPB2 sequence data are available. Ho et al.
(2001) reported that Rhamphoria pyriformis was found
from freshwater habitats in China, but they did not provide
illustrations and descriptions for this species.
Rhodoveronaea Arzanlou et al., Stud. Mycol. 58: 89
(2007)
Asexual morph Colonies velvety, floccose; surface olivaceous-grey to dark olivaceous-green, reverse olivaceousblack. Hyphae pale olivaceous, smooth, thin-walled.
Conidiophores arising vertically from creeping hyphae,
straight or flexuose, simple, thick-walled, red-brown, with
inflated basal cell. Conidiogenous cells polyblastic, terminally integrated, sympodial, smooth, thick-walled, pale
brown, rachis straight, occasionally geniculate, with
crowded, slightly prominent conidium-bearing denticles,
denticles flat-tipped, slightly pigmented. Conidia solitary,
ellipsoidal to obovoidal, aseptate to multiseptate, with a
protruding base and a marginal basal frill, pale brown,
thinor slightly thick-walled, smooth. Conidial secession
schizolytic (Arzanlou et al. 2007). Sexual morph Ascomata nonstromatic, gregarious or solitary, dark brown to
black, venter subglobose to conical, immersed; neck conical, emerging above the substratum, straight or slightly
curved, ostiolum periphysate. Peridium leathery, two-layered. Paraphyses septate, hyaline, tapering towards the tip,
longer than the asci. Asci 8-spored, unitunicate, cylindrical,
broadly rounded at the apex, long-stipitate. Ascospores
fusiform, aseptate to multiseptate, hyaline (Réblová 2009).
Type species: Rhodoveronaea varioseptata Arzanlou,
W. Gams & Crous, Stud. Mycol. 58: 91 (2007)
Notes: Arzanlou et al. (2007) introduced the genus
Rhodoveronaea with single asexual species R. varioseptata. Réblová (2009) provided the sexual morph for this
species. In this study, we introduce the second species R.
aquatica, which was collected from freshwater habitats in
northwestern Yunnan, China.
Rhodoveronaea aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555661, Facesoffungi
number: FoF 05435, Fig. 32
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Fig. 33 Pseudostanjehughesia lignicola (DLU 078, holotype) a, c–f Conidia. b, g Germinating conidium. h, i Culture on PDA after 20 days
(i from below). Scale bars: a–g 20 lm
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1593
123
Saprobic on decaying, submerged wood in freshwater
habitats. Asexual morph Hyphomycetous. Colonies
effuse, inconspicuous, brown, hairy. Conidiophores 182–
310 lm long, 9–13 lm wide (
x = 246 9 11 lm, n = 20),
Fungal Diversity (2019) 99:451–660
macronematous, mononematous, cylindrical, arising vertically from creeping hyphae, straight or flexuose, simple,
thick-walled, septate, red-brown, paler at apex. Conidiogenous cells polyblastic, terminally integrated, sympodial,
smooth, thick-walled, pale brown at the base, paler towards
the apex. Conidia 23–27 lm long, 9–11 lm wide
(
x = 25 9 10 lm, n = 20), acropleurogenous, ellipsoid to
obovoid, apically rounded, with a flat basal scar, 1–3-septate, pale brown, smooth-walled. Conidial secession schizolytic. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on submerged decaying wood in
a freshwater stream, 10 July 2015, X.J. Su, S-718 (MFLU
18–1593, holotype), ex-type living culture MFLUCC
18–1339.
Notes: Rhodoveronaea aquatica resembles R. varioseptata in having macronematous, mononematous,
cylindrical, straight or flexuose, septate, red-brown conidiophores, terminally integrated, polyblastic, sympodial,
smooth conidiogenous cells, ellipsoid to obovoid, pale
brown, septate conidia (Arzanlou et al. 2007; Réblová
2009). However, R. aquatica differs from R. varioseptata
in having longer conidiophores (182–310 vs. 80–125 lm)
and
larger
conidia
(23–27 9 9–11 lm
vs.
8–15 9 3–4 lm). Phylogenetic analysis also shows that
Rhodoveronaea aquatica and R. varioseptata are distinct
species (Fig. 1, clade 8).
Woswasiaceae Zhang et al.
Cyanoannulus Raja et al., Mycotaxon 88: 8 (2003)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Raja et al. (2003).
Type species: Cyanoannulus petersenii Raja, J. Campb.
& Shearer
Notes: The genus Cyanoannulus was introduced by Raja
et al. (2003) with single species Cyanoannulus petersenii
Raja, J. Campb. & Shearer which was collected from
freshwater habitats. And this species is known only from
the type locality.
Cyanoannulus petersenii Raja et al.
Distribution: USA, North Carolina and Tennessee, Great
smoky Moubtains National Park, on submerged wood
(Raja et al. 2003).
Asexual morph: Undetermined
Notes: Holotype ILL, Raja & Hamburger R044-1. ITS
and LSU sequence data are available.
Diaporthomycetidae genera incertae sedis
Aquaticola Ho et al., Fungal Diversity Res. Ser. 3: 88
(1999)
Asexual morph Undetermined. Sexual morph Descriptions and illustration see Ho et al. (1999c).
Type species: Aquaticola hyalomura W.H. Ho, C.K.M.
Tsui, Hodgkiss & K.D. Hyde, Fungal Divers 3: 88 (1999)
533
Notes: The genus Aquaticola was introduced for tropical
freshwater fungi occurring on submerged wood in Asia
with two species and was placed in the family Annulatascaceae (Ho et al. 1999c). Subsequently, Tsui et al. (2003)
described three additional Aquaticola species from tropical
freshwater environments in Asia and Australia. Réblová
et al. (2016a) transferred Aquaticola ellipsoidea to the
genus Atractospora as At. ellipsoidea. Four freshwater
species are accepted in Aquaticola.
Aquaticola hyalomura Ho et al.
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged decaying wood (Ho et al. 1999c).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2969 (now in IFRD). LSU
sequence data is available.
Aquaticola longicolla Tsui et al.
Distribution: Australia, north Queensland, near
Ravenshoe, submerged in creek (Tsui et al. 2003); Brunei,
Tutong River, on submerged wood (Fryar et al. 2004)
Asexual morph: Undetermined
Notes: Holotype IFRD 8682. Sequence data is not
available.
Aquaticola minutiguttulata Tsui et al.
Distribution: China, Hong Kong, Sai Kung, Hang Cho
Shui, on submerged wood (Tsui et al. 2003).
Asexual morph: Undetermined
Notes: Holotype IFRD 8683. Sequence data is not
available.
Aquaticola triseptata Tsui et al.
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Tsui et al. 2003).
Asexual morph: Undetermined
Notes: Holotype IFRD 8684. Sequence data is not
available.
Bullimyces Ferrer et al., Mycologia 104(4): 868 (2012)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Ferrer et al. (2012).
Type species: Bullimyces communis Ferrer et al.,
Mycologia 104(4): 868 (2012)
Notes: Ferrer et al. (2012) introduced the genus Bullimyces with three freshwater species. All these species are
known only from their respective type localities.
Bullimyces aurisporus Ferrer et al.
Distribution: Costa Rica, Alajuela, on submerged wood
(Ferrer et al. 2012).
Asexual morph: Undetermined
Notes: Holotype ILL AF316–1. LSU and SSU sequence
data are available.
Bullimyces communis Ferrer et al.
Distribution: Costa Rica, Alajuela and Heredia, on
submerged wood (Ferrer et al. 2012).
Asexual morph: Undetermined
123
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Fungal Diversity (2019) 99:451–660
Fig. 34 a Cylindrotrichum submersum (MFLU 18–2320, holotype) a Colonies on wood. b Conidiophore. c, d Conidiogenous cells. e–
j Conidium. Colonies on PDA from above (l) and reverse (k). Scale bars: b 40 lm, c, d 20 lm, e–j 10 lm
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Fungal Diversity (2019) 99:451–660
Notes: Holotype ILL AF281–1, other specimens collected from freshwater habitats: ILL AF281–4, ILL
AF281–7, ILL AF281–3. LSU, SSU sequence data are
available.
Bullimyces costaricensis Ferrer et al.
Distribution: Costa Rica, Limón, on submerged wood
(Ferrer et al. 2012).
Asexual morph: Undetermined
Notes: Holotype ILL AF317–1. LSU and SSU sequence
data are available.
Cancellidium Tubaki, Trans. Mycol. Soc. Japan 16(4): 357
(1975)
Asexual morph Colonies on natural substrate effuse,
black, shiny. Mycelium immersed or superficial, composed
of septate, subhyaline to hyaline, smooth-walled hyphae.
Conidiophores micronematous, short. Conidiogenous cells
integrated, terminal, determinate, cylindrical, subhyaline.
Conidia acrogenous, solitary, muriform, dictyosporous,
strongly flattened, fan-shaped, obovate to obcordate, brown
to almost black, shiny. Sexual morph Undetermined.
Type species: Cancellidium applanatum Tubaki, Trans.
Mycol. Soc. Japan 16(4): 358 (1975)
Notes: The genus Cancellidium is typified by C.
applanatum which was collected from wood blocks of
Ochroma pyramidale (Cav. ex Lam.) Urb. in Kobe, Japan.
The genus includes two species (Tubaki 1975; Yeung et al.
2006).
Cancellidium applanatum Tubaki
Distribution: Perú, Cusco, on submerged, decomposing
woody and herbaceous debris in freshwater habitats (Zelski
et al. 2014).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater: ILL
41205, ILL 41206. LSU sequence data is available.
Ceratostomella Sacc., Michelia 1(no. 4): 370 (1878)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Inderbitzin (2000) and Réblová
(2006).
Type species: Ceratostomella vestita Sacc., Michelia
1(no. 4): 370 (1878)
Notes: Index fungorum (2018) lists 112 epithets for the
genus Ceratostomella and only one species was collected
from freshwater habitats (Inderbitzin 2000).
Ceratostomella hyalocoronata Inderb
Distribution: China, Guangdong Province, on submerged wood in a stream (Inderbitzin 2000).
Asexual morph: Undetermined
Notes: Holotype UBC F13874. Sequence data is not
available.
Clohiesia K.D. Hyde, Nova Hedwigia 61(1-2): 125 (1995)
535
Asexual morph: Undetermined. Sexual morph: Descriptions and illustrations see Hyde (1995), Cai & Hyde
(2007).
Type species: Clohiesia corticola K.D. Hyde, Nova
Hedwigia 61(1–2): 126 (1995)
Notes: Clohiesia K.D. Hyde was established by Hyde
(1995) to accommodate the freshwater taxon C. corticola
from tropical Australia. Subsequently, another two species
Clohiesia curvispora and C. lignicola were introduced by
Tsui et al. (1998) and Cai and Hyde (2007) respectively.
All these species were collected from freshwater habitats.
Presently, Clohiesia is placed in the family Annulatascaceae (Maharachchikumbura et al. 2015, 2016;
Wijayawardene et al. 2017). However, Zhang et al. (2016)
showed that Clohiesia corticola (HKUCC 3712) is close to
Chaetosphaeriales based on phylogenetic analyses. In this
study, Clohiesia corticola (HKUCC 3712) is basal to
Phyllachoraceae based on phylogenetic analysis, therefore
we assign this genus to Diaporthomycetidae genera incertae sedis.
Clohiesia corticola K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Hyde 1995b); Japan, Koito River, on submerged
wood (Tsui et al. 2001a).
Asexual morph: Undetermined
Notes: Holotype BRIP 21485. LSU sequence data is
available.
Clohiesia curvispora L. Cai & K.D. Hyde
Distribution: China, Yunnan Province, Jinghong city,
on submerged wood in a small forest stream (Cai and Hyde
2007).
Asexual morph: Undetermined
Notes: Holotype HKU(M) 10854 (now in IFRD).
Sequence data is not available.
Clohiesia lignicola Tsui et al.
Distribution: China, Hong Kong, on submerged wood
(Tsui et al. 1998a)
Asexual morph: Undetermined
Notes: Holotype HKU(M) 5533 (now in IFRD).
Sequence data is not available.
Hyalorostratum Raja & Shearer, Mycosphere 1(4): 281
(2010)
Asexual morph Undetermined. Sexual morph Description
and illustrations refer to Raja et al. (2010).
Type species: Hyalorostratum brunneisporum Raja &
Shearer, Mycosphere 1(4): 281 (2010)
Notes: Hyalorostratum was introduced by Raja et al.
(2010) with Hyalorostratum brunneisporum Raja &
Shearer as type species which was collected from freshwater habitat in north America. Raja et al. (2010) suggested
that Hyalorostratum belongs to the order Diaporthales
based on results of their phylogenetic analyses, but they did
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536
not assign it to any of families in Diaporthales and referred
it as Diaporthales genera incertae sedis. In our phylogenetic analysis, two strains of Hyalorostratum brunneisporum formed a distinct subclade and basal to Jobellisiales,
we also assign this genus to Diaporthomycetidae genera
incertae sedis. Presently, a single species is accepted in this
genus.
Hyalorostratum brunneisporum Raja & Shearer
Distribution: USA, Alaska, Headquarters, Lake Kenai
Wildlife Refuge; New Hampshire, Hubbard Brook Forest
stream, on submerged woody debris, on submerged woody
debris (Raja et al. 2010).
Asexual morph: Undetermined
Notes: Holotype ILL 40792. LSU and SSU sequence
data are available.
Pseudostanjehughesia J. Yang & K.D. Hyde, Mycol Prog
17: 609 (2018)
Asexual morph Description and illustrations see Yang
et al. (2018). Sexual morph Undetermined.
Type species: Pseudostanjehughesia aquitropica J.
Yang & K.D. Hyde, Mycol. Progr. 17(5): 610 (2017)
Notes: The genus Pseudostanjehughesia was introduced
by Yang et al. (a, b) with single species which was collected from freshwater habitats in Thailand. Pseudostanjehughesia resembles Stanjehughesia and Linkosia due to
the absence or reduced conidiophores, and brown and
obclavate conidia. Linkosia is easily distinguished from
Pseudostanjehughesia by the lageniform or ampulliform
conidiogenous cells and distoseptate conidia. It is difficult
to separate Pseudostanjehughesia from Stanjehughesia as
both genera share similar morphological characters of
conidiophores, conidiogenous cells and conidia. However,
they are phylogenetically distinct.
Pseudostanjehughesia aquitropica J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Yang
et al. 2018a).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0857, isotype GZAAS
17–0010. ITS, LSU, SSU and TEF1a sequence data are
available.
Pseudostanjehughesia lignicola Z.L. Luo, K.D. Hyde &
H.Y. Su, sp. nov.
Index Fungorum number: IF 555662, Facesoffungi
number: FoF 05436, Fig. 33
Etymology: Referring to this fungus dwelling on wood
Holotype: DLU 078
Saprobic on decaying, submerged wood in freshwater
habitats. Asexual morph Hyphomycetous. Colonies
effuse, dark brown, scattered, glistening. Mycelium partly
superficial, composed of brown, septate, branched hyphae.
Conidiophores indistinct. Conidiogenous cells holoblastic,
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monoblastic, integrated, terminal, cylindrical. Conidia 51–
62 lm long, 16–22 lm wide (
x = 57 9 19 lm, n = 20),
acrogenous, solitary or group in 2–3 conidia, ovoid,
slightly curved, multi-euseptate, verrucose, dark brown at
base, tapering and becoming pale brown towards the apex,
truncate at the base. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, saprobic
on decaying wood submerged in Heilong stream in Cangshan Mountain. 15 March 2014, Z.L. Luo, HLXM 7-4
(DLU 078, holotype) ex-type living culture MFLUCC
15–0352.
Notes: Pseudostanjehughesia lignicola morphologically
resembles P. aquitropica in having terminal, monoblastic
conidiogenous cells, acrogenous, multi-euseptate, verrucose conidia dark brown at base, tapering and becoming
pale brown towards the apex (Yang et al. 2018a). However,
P. lignicola differs from P. aquitropica in having ovoid,
shorter conidia (51–62 lm vs. 55–120 lm), while P.
aquitropica have rostrate conidia which are oval or ellipsoidal when young, fusiform or obclavate when mature.
Phylogenetic analysis shows that Pseudostanjehughesia
lignicola clusters with P. aquitropica with strong support
(100% ML) (Fig. 1, clade 10). Following Jeewon and
Hyde’s (2016) recommendations on species delimitation
for new species, we delved into pairwise dissimilarities of
DNA sequences and noted that there are indeed differences
in the ribosomal ITS sequences, 68 noticeable nucleotide
differences including 11 gaps among the 485 nucleotides
analysed between Pseudostanjehughesia lignicola and P.
aquitropica. Pseudostanjehughesia lignicola is the second
species for Pseudostanjehughesia and both species are
collected from freshwater habitats.
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Coronophorales Nannf. (=Melanosporales N. Zhang &
M. Blackw.)
Bertiaceae Smyk
Bertia De Not., G. bot. ital. 1(1): 334 (1844)
Asexual morph Undetermined. Sexual morph Updated
descriptions and illustrations see Maharachchikumbura
et al. (2016).
Type species: Bertia moriformis (Tode) De Not., G. bot.
ital. 1(1): 335 (1844)
Notes: The genus Bertia was introduced by De Notaris
(1844) and 48 epithets are listed in Index Fungorum (December 2018). Among these species, only Bertia convolutispora K.D. Hyde was collected from freshwater habitats
(Hyde 1995c).
Bertia convolutispora K.D. Hyde
Distribution: Australia, north Queensland, on wood
submerged in stream (Hyde 1995c).
Asexual morph: Undetermined
Fungal Diversity (2019) 99:451–660
Notes: Holotype BRIP 22512. Sequence data is not
available.
Glomerellales Chadef. ex Réblová et al.
Reticulascaceae Réblová & W. Gams
Cylindrotrichum Bonord., Handbuch der allgemeinen
Mykologie: 88 (1851)
Asexual morph Hyphomycetous. Colonies in vivo brown
to black, hairy, effuse. Setae absent. Conidiophores
macronematous, mononematous, cylindrical, straight.
Conidiogenous cells usually monophialidic, rarely
polyphialidic with up to two lateral enteroblastic openings,
collarette hyaline to subhyaline. Conidia cylindrical,
slightly tapering, rounded at apex, obtuse at base, 1-septate,
not constricted at septum, hyaline, guttulate, smooth.
Sexual morph Stroma absent. Ascomata superficial, solitary, or gregarious, brown, venter subglobose to conical.
Ostiolum periphysate. Peridium fragile, 2-layered. Paraphyses septate, hyaline, filiform, forming a branching and
anastomosing ‘‘network’’. Asci 8-spored, unitunicate,
cylindrical-clavate, short-stipitate. Ascospores ellipsoidal
to fusiform, septate, hyaline.
Type species: Cylindrotrichum oligospermum (Corda)
Bonord., Handb. Allgem. mykol. (Stuttgart): 88 (1851)
Notes: Réblová et al. (2011) introduced a new genus
Reticulascus including two species. Reticulascus clavatus
with its asexual morph Cylindrotrichum clavatum is a
common dweller of submerged wood in lotic sites in
France. Cylindrotrichum includes 23 names (Rambelli and
Onofri 1987) while Reticulascus includes only two names
(Réblová et al. 2011). Given its widespread use, priority,
and greater number of names, Réblová et al. (2016b) recommend the use of Cylindrotrichum over Reticulascus.
Cylindrotrichum aquaticum (Luo et al.) Z.L. Luo & K.D.
Hyde comb. nov.
: Blastophorum aquaticum Luo et al., Fungal Divers
80:177 (2016)
Index Fungorum: IF 555663; Facesoffungi number: FoF
05437
Description: For a complete description of this taxon see
Hyde et al. (2016) as Blastophorum aquaticum.
Notes: Holotype DLU 084. ITS and LSU sequence data
are available. This fungus was introduced as Blastophorum
aquaticum by Hyde et al. (2016). It resembles Cylindrotrichum species in morphology of conidiophores,
conidiogenous cells and conidia. In this study, the phylogenetic analysis shows that the isolate from type clusters
within the genus Cylindrotrichum with strong support
(Fig. 1, clade 42). Based on the morphology and phylogeny, we synonymized Blastophorum aquaticum under
Cylindrotrichum aquaticum.
Cylindrotrichum clavatum W. Gams & Hol.-Jech., Studies
in Mycology 43: 54 (1976).
537
Distribution: China, Yunnan Province, on submerged
wood (Maharachchikumbura et al. 2018); France, on
submerged wood of Alnus glutinosa, Fraxinus sp., Platanus sp. (Réblová et al. 2011).
Asexual morph: see Réblová et al. (2011).
Notes: Holotype CBS 128.76, other specimens collected
from freshwater habitats: PRM 915717, PRM 915719,
DLU 0575, DLU 0572. ITS, LSU, SSU and RPB2
sequence data are available. Cylindrotrichum clavatum was
found on submerged wood in freshwater habitats in France
and China respectively (Réblová et al. 2011;
Maharachchikumbura et al. 2018).
Cylindrotrichum gorii Lunghini
Distribution: China, Yunnan Province, Dali, on submerged in a stream in Jizu Mountain (Maharachchikumbura et al. 2018).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater habitats:
MFLU 17–1965. ITS, LSU and RPB2 sequence data are
available.
Cylindrotrichum submersum Z.L. Luo, H.Y. Su & K.D.
Hyde sp. nov.
Index Fungorum number: IF 555664, Facesoffungi
number: FoF 05439, Fig. 34
Etymology: Referring to the submerged habitats of this
fungus
Holotype: MFLU 18–2320
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate,
effuse, superficial, brown to black, hairy, in groups.
Conidiophores 77–135(–148) lm long, 4–6 lm wide
(
x = 106 9 5 lm, n = 20), macronematous, mononematous, solitary, cylindrical, erect, straight or slightly flexuous, unbranched, septate, dark brown, becoming pale
brown to subhyaline towards the apex, smooth. Conidiogenous cells monophialidic, integrated, terminal, without a
conspicuous collarette. Conidia 15–17 lm long, 4–6 lm
wide (
x = 16 9 5 lm, n = 20), acrogenous, cylindrical or
clavate, rounded at apex, slightly tapering, guttulate,
aseptate, hyaline, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Jizu
Mountain, saprobic on decaying wood submerged in a
freshwater stream, October 2016, Z.L. Luo, S-988 (MFLU
18–2320, holotype), ex-type living culture DLUCC 0988.
Notes: Cylindrotrichum submersum resembles C.
clavatum in having macronematous, solitary, cylindrical,
erect, straight or slightly flexuous, unbranched, septate
conidiophores of similar size and cylindrical or clavate,
hyaline, smooth conidia. However, Cylindrotrichum submersum differs from C. clavatum in having monophialidic
conidiogenous cells, aseptate, guttulate, longer (15–17 vs.
8.5–13 lm) conidia, while the later one has monophialidic
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b Fig. 35 Cosmospora aquatica (MFLU 15–0072, holotype) a, b As-
comata on natural substrate. c Breaked ascomata. d Section of
ascomata. e Structure of papillate on ascomata. f Structure of
peridium. g, h Asci. i–k Ascospores. l Germinating ascospore. Scale
bars: b 200 lm, c, d 100 lm, e, g, h 30 lm, l 20 lm, i–k 10 lm
or polyphialidic conidiogenous cells, 1–2-septate conidia
(Gams and Holubova-Jechova 1976; Maharachchikumbura
et al. 2018). Furthermore, they are phylogenetically distinct
species (Fig. 1, clade 42). To further support Cylindrotrichum submersum as a new species, we compared
nucleotide differences with C. aquaticum (MFLUCC
15–0264) following the guidelines of Jeewon and Hyde
(2016). Comparison of the 543 nucleotides across the ITS
region reveals 36 bp differences including 6 gaps. Based
on the differences of morphology and DNA nucleotide, we
introduce our isolate as new species in Cylindrotrichum.
Kylindria DiCosmo et al., Mycologia 75 (6): 970 (1983)
Asexual morph Hyphomycetous. Colonies effuse, hairy,
dark. Conidiophores macronematous, mononematous,
setiform, solitary, erect, sometimes curved, unbranched,
often in fascicles, multiseptate, thick-walled, dark brown
below, paler above, smooth-walled. Conidia blastic-phialidic, 1-several-septate, oblong-ellipsoid or cylindrical,
apex rounded, base tapered and truncate, hyaline, smoothwalled. Sexual morph Undetermined.
Type species: Kylindria triseptata (Matsush.) DiCosmo
et al., Mycologia 75(6): 971 (1983)
Notes: DiCosmo et al. (1983) introduced the genus
Kylindria based on Cylindrotrichum triseptatum Matsush.
(Matsushima 1975). Presently, there are 21 species included in this genus (Mycobank 2019). Three species of
Kylindria have been reported from freshwater habitats in
China and UK (Hyde and Goh 1999; Maharachchikumbura
et al. 2018).
Kylindria aquatica Luo et al.
Distribution: China, Yunnan Province, on submerged
wood (Maharachchikumbura et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–1967, paratype MFLU
17–1966. ITS, LSU and RPB2 sequence data are available.
Kylindria chinensis Maharachch. et al.
Distribution: China, Yunnan Province, on submerged
wood (Maharachchikumbura et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–1964. ITS, LSU and RPB2
sequence data are available.
Hypocreales Lindau
Bionectriaceae Samuels & Rossman
Didymostilbe Henn., Hedwigia 41: 148 (1902)
539
Asexual morph Conidiomata synnemayous, solitary,
errect, straight, cylindrical to clavate, or subulte-capitate,
Conidiophores macronematous, unbranched. Conidiogenous cells enteroblastic phialidic, integrated or discrete,
terminal or lateral, hyaline, smooth-walled, cylindrical.
Conidia acrogenous, hyaline, 0–1-septate, sometimes
slightly constricted at septum, cylindrical to slight clavate,
broadly rounded at the apex, subtruncate to obconically
truncate at the base, thick-walled, smooth. Sexual morph
Undetermined.
Type species: Didymostilbe coffeae Henn., Hedwigia
41: 148 (1902)
Notes: The genus Didymostilbe was established by
Hennings (1902) with D. coffeae as the type species. Index
Fungorum (2019) list 17 epithets for Didymostilbe. Only
Didymostilbe australiensis Goh & K.D. Hyde is collected
from freshwater habitats from Queensland, Australia (Hyde
and Goh 1997).
Didymostilbe australiensis Goh & K.D. Hyde
Distribution: Australia, Queensland, Cape Tribulation,
Mountain Lewis, on decaying wood submerged in a stream
(Hyde and Goh 1997).
Sexual morph: Undetermined
Notes: Holotype ML 28 (BRIP). Sequence data is not
available. Didymostilbe australiensis was introduced by
Hyde and Goh (1997) based on their collection from
freshwater habitats in Australia, and it is known only from
the type locality.
Hypocreaceae De Not.
Trichoderma Pers., Neues Mag. Bot. 1: 92 (1794)
Asexual morph Hyphomycetous. Colonies usually growing rapidly, mycelium initially submerged, eventually with
variably matted, floccose, woolly or arachnoid hyaline
aerial mycelium. Conidiophores in most species with a
broad main axis branched at regular intervals, usually with
successive branches apically and distally progressively
shorter and narrower, branches more or less divergent,
solitary, paired or in verticils. Conidiogenous cells phialidic, typically disposed in divergent verticels terminally on
branches of the conidiophore, or in whorls directly beneath
septa along the conidiophore and branches, otherwise
paired or solitary and irregularly disposed; cylindrical,
subulate, lageniform, ampulliform or subglobose; usually
attenuated to a narrow, short-cylindrical, conidium bearing
neck. Conidia one-celled, typically green, or otherwise
colourless, greyish, or brownish; smooth-walled to distinctly roughened, or with sinuate, bullate or winglike
projections from the outer wall, subglobose, obovoid,
ellipsoid, oblong or short-cylindrical; accumulating in
gloeoid heads, sometimes enclosed in a sac-like sheath
visible at high magnifications. Sexual morph Stromata
solitary or aggregated, rounded or elongate, yellowishbrown to brown with white-yellow margin, becoming
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b Fig. 36 Phylogram generated from maximum likelihood analysis
based on combined ITS and LSU sequence data for species of
Cosmospora (with Cosmosporella olivacea as outgroup). The best
scoring RAxML tree with a final likelihood value of - 2406.322240
is presented. RAxML bootstrap support values equal to or greater than
75% are given before the forward slash (black). Maximum parsimony
bootstrap support values equal to or greater than 75% are given after
the forward slash (red). Hyphen (‘-’) indicates a value lower than 75%
for RAxML and maximum parsimony. Newly generated sequences
are in red
pulvinate, more rarely turbinate or discoid, circular to
irregular in outline, surface smooth to slightly uneven to
granular, broadly attached, margin often becoming free and
concolorous with stroma surface. Ostiolar openings visible
as slightly raised, purple-brown spots. Ascomata immersed
in stroma, densely disposed, globose to subglobose.
Peridium colourless, consisting of laterally strongly compressed thin hyphae, basally and apically pseudoparenchymatous, indistinct, scarcely differentiated from
and merging with the surrounding tissue, apical part
flanking the ostioles conspicuously thickened. Asci cylindrical, with short stipe, sometimes with a knob-like base,
with apical minute pore. Ascospores 1-septate, dimorphic,
distal part subglobose to oval, sometimes slightly tapered
towards the upper end, proximal part oblong to wedgeshaped, with lower end broadly rounded, sometimes verrucose, hyaline.
Type species: Trichoderma viride Pers., Neues Mag.
Bot. 1: 92 (1794)
Notes: The genus Trichoderma is cosmopolitan in soils
and on decaying wood and vegetable matter. Species of
Trichoderma are frequently from dominant components of
the soil microflora in widely varying habitats (Kubicek and
Harman 1998). Au et al. (1992) reported T. glaucum
Abboott from freshwater habitats in Hong Kong, China.
Trichoderma glaucum Abboott
Distribution: China, Hong Kong, on submerged leaves
(Au et al. 1992).
Sexual morph: Undetermined.
Notes: ITS sequence data is available in GenBank.
Nectriaceae Tul. & C. Tul.
Aquanectria L. Lombard & Crous, Stud. Mycol. 80: 207
(2015)
Holomorph Descriptions and illustrations refer to Lombard et al. (2015).
Type species: Aquanectria penicillioides (Ingold) L.
Lombard & Crous, Stud. Mycol. 80: 207 (2015)
Notes: The aquatic genus Aquanectria was established
by Lombard et al. (2015) to accommodate two fungal
species previously treated as members of the genera
Flagellospora and Heliscus (Ingold 1942; Ranzoni
1956a, b; Hudson 1961). Three species are included in this
541
genus and all were collected from freshwater habitats. In
this study, description and illustrations for Aquanectria
penicillioides are provided and it is new record for China.
Aquanectria jacinthicolor Huang et al.
Distribution: China, Yunnan Province, Baoshan City,
on submerged wood in a stream along the roadside (Huang
et al. 2018a, b).
Asexual morph: Undetermined.
Notes: Holotype HKAS 99551, other specimen collected
from freshwater habitat: HKAS 92802. ITS and LSU
sequence data are available.
Aquanectria penicillioides (Ingold) L. Lombard & Crous
Distribution: China, Yunnan Province, Dali, Erhai lake,
on submerged decaying wood (This study). USA, California, Napa County, Green Valley Falls, on decaying leaves
of Acer sp. submerged in a stream (Lombard et al. 2015).
Sexual morph: Refer to Ranzoni (1956a, b).
Notes: Specimen collected from freshwater habitats:
HKAS 92560. Acl1, act, cmdA, ITS, LSU, RPB1, RPB2
and TEF1a sequence data are available.
Aquanectria submersa (H.J. Huds.) L. Lombard & Crous
: Heliscus submersus H.J. Huds., Trans. Br. mycol.
Soc. 44(1): 91 (1961)
Distribution: Jamaica, St. Andrew, Hardwar Gap, on
decaying leaves submerged in a stream (Hudson 1961).
Sexual morph: Undetermined
Notes: Holotype IMI 76792. acl1, act, his3, ITS, LSU,
RPB1, RPB2 and TEF1a sequence data are available.
Baipadisphaeria Pinruan, Mycosphere 1: 58 (2011)
Asexual morph Undetermined. Sexual morph Ascomata
immersed, or semi-immersed, coriaceous, ostiolate, solitary. Asci 8-spored, unitunicate, clavate to ovoid, apedicellate, apically narrow and rounded, lacking any apical
structure. Paraphyses elongate, unbranched, hyphal-like.
Ascospores 3–4-seriate, fusiform to cylindrical, straight or
curved, aseptate, hyaline to pale brown, smooth-walled.
Type species: Baipadisphaeria spathulospora Pinruan,
Mycosphere 1: 58 (2011)
Notes: The genus Baipadisphaeria was introduced by
Pinruan et al. (2010) with single species which was collected from freshwater habitats in Thailand.
Baipadisphaeria spathulospora Pinruan
Distribution: Thailand, Narathiwat Province, Sirindhorn Peat Swamp Forest, on submerged trunk of Licuala
longicalycata (Pinruan et al. 2010).
Asexual morph: Undetermined
Notes: Holotype BBH, Pinruan Wah32A; LSU, SSU
sequence data are available.
Chaetopsina Rambelli, Atti Accad. Sci. Ist. Bologna, Cl.
Sci. Fis. Rendiconti 3: 5 (1956)
: Chaetopsinectria J. Luo & W.Y. Zhuang, Mycologia
102: 979 (2010).
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542
Asexual morph Descriptions and illustrations refer to
Rambelli (1956). Sexual morph Descriptions and illustrations refer to Luo and Zhuang (2010).
Type species: Chaetopsina fulva Rambelli, Diagn. IV 3:
5 (1956)
Notes: The dematiaceous hyphomycete genus Chaetopsina was established by Rambelli (1956) with C. fulva
as the type species. Luo and Zhuang (2010) established a
sexual genus Chaetopsinectria based on Cosmospora
chaetopsinae (Samuels 1985), for a group of fungi having
Chaetopsina asexual morphs. Lombard et al. (2015) proposed that the sexual genus Chaetopsinectria (2010) be
suppressed in favour of asexual genus Chaetopsina (1956),
which has priority by date and would require no new
combinations. Chaetopsina hongkongensis was introduced
by Goh and Hyde (1997a) on decaying rachids of Phoenix
roebelenii. Ho et al. (2002b) reported this species from
freshwater habitats in Hong Kong, China. We provide the
DNA sequence data for Chaetopsina beijingensis based on
the specimen collected from decaying wood submerged in
a stream in Dali, Yunnan Province, China.
Chaetopsina hongkongensis Goh & K.D. Hyde
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
Notes: Holotype IFRD 8729. Sequence data is not
available.
Chaetopsina beijingensis Crous & Y. Zhang ter
Distribution: China, Yunnan Province, Jizu Mountain,
saprobic on decaying wood submerged in a freshwater
stream.
Sexual morph: Undetermined
Notes: Holotype CBS H-21718, specimen collected
from freshwater MFLU 18–2327. ITS, LSU, SSU and
TEF1a sequence data are available. Chaetopsina beijingensis was introduced by Crous et al. (2014a) based on a
collection obtained from needles of Pinus tabulaeformis in
Beijing, China.
Cosmospora Rabenh., Hedwigia 2: 59 (1862)
Holomorph Descriptions and illustrations refer to Lombard et al. (2015).
Type species: Cosmospora coccinea Rabenh., Hedwigia
2: 59 (1862)
Notes: Cosmospora was established in 1862 and is
typified with C. coccinea Rabenh. (Rossman et al. 1999).
About 73 species have been attributed to Cosmospora
sensu Rossman (Gräfenhan et al. 2011; Lombard et al.
2015; Zeng and Zhuang 2016). In this study, we introduce
a new Cosmospora species based on the isolates obtained
from submerged wood in freshwater habitats.
Cosmospora aquatica Z.L. Luo, H.Y. Su & K.D. Hyde, sp.
nov.
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Fungal Diversity (2019) 99:451–660
Index Fungorum number: IF 555665, Facesoffungi
number: FoF 05442, Fig. 35
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 15–0072
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 210–318 lm high, 176–296 lm diam., perithecial, solitary, rarely gregarious, nonstromatic, superficial,
globose to subglobose, papillate, surface smooth to slightly
roughened, orange red. Peridium 20–27 lm, composed of
irregular, thick walled cells. Paraphyses subhyaline, septate, constricted at septa. Asci 65–79 9 7–9 lm
(
x = 72 9 8 lm, n = 10), 8-spored, unitunicate, cylindrical, subhyaline, with apical ring. Ascospores 9–11 9 3.5–
4.5 lm (
x = 10 9 4 lm, n = 20), uniseriate, oval or
oblong, guttulate, uniseptate, slightly constricted at septa,
hyaline or subhyaline, smooth-walled.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on submerged decaying wood in a
freshwater stream, September 2013, Z.L. Luo, ZL-32
(MFLU 15–0072, holotype; HKAS 86447, isotype), extype living culture MFLUCC 13–0884; Ibid., on submerged wood in a freshwater stream, March 2014, H.Y. Su,
S-026 (HKAS 83987, paratype), living culture MFLUCC
15–0244.
Notes: Cosmospora aquatica mostly resembles C. lavitskiae in having perithecial, superficial, globose to subglobose, orange red ascomata, unitunicate asci and
uniseptate, hyaline or subhyaline, uniseriate ascospores
(Zeng and Zhuang 2016). However, Cosmospora aquatica
differs from C. lavitskiae in having larger asci
(65–79 9 7–9 vs. 40–52.5 9 2.5–4.5 lm) with apical ring,
while the asci of C. lavitskiae without apical ring, and
guttulate, longer ascospores (9–11 vs. 4–6 lm). Phylogenetic analysis shows that Cosmospora aquatica is distinct
from other Cosmospora species (Fig. 36).
Cosmosporella Huang et al., Cryptog. Mycol. 39(2): 179
(2018)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Huang et al. (2018b).
Type species: Cosmosporella olivacea S.K. Huang, R.
Jeewon & K.D. Hyde, Cryptog. Mycol. 39(2): 181 (2018)
Notes: The genus Cosmosporella was introduced by
Huang et al. (2018b) with Cosmosporella olivacea as type
species which was collected from freshwater habitats in
China.
Cosmosporella olivacea Huang et al.
Distribution: China, Xinjiang Province, a lake near by
the snow mountains, on dead wood (Huang et al. 2018b).
Asexual morph: Undetermined
Fungal Diversity (2019) 99:451–660
Notes: Holotype HKAS99607. ITS, LSU and b-tubulin
sequence data are available.
Fusicolla Bonord., Handb. Allgem. mykol. (Stuttgart): 150
(1851)
Holomorph Descriptions and illustrations refer to
Gräfenhan et al. (2011).
Type species: Fusicolla betae Bonord., Handb. Allgem.
mykol. (Stuttgart): 150 (1851)
Notes: Fusicolla belongs to the Nectriaceae and comprises an estimated 8 species (Wijayawardene et al. 2017).
Fusicolla has generally been considered a synonym of
Fusarium. Gräfenhan et al. (2011) resurrected this genus
typified by F. betae. Huang et al. (2018a, b) reported Fusicolla aquaeductuum from freshwater habitats in China.
Fusicolla aquaeductuum (Radlk. & Rabenh.) Gräfenhan
et al.
Distribution: China, Xinjiang Province, a lake near by
the snow mountains, on dead wood (Huang et al. 2018a, b).
Asexual morph: Undetermined
Notes:
Specimen
collected
from
freshwater
HKAS99608. ITS, LSU and b-tubulin sequence data are
available.
Gliocladiopsis S.B. Saksena, Mycologia 46: 662 (1954)
Holomorph Descriptions and illustrations refer to Lombard et al. (2015).
Type species: Gliocladiopsis sagariensis S.B. Saksena,
Mycologia 46: 662 (1954)
Notes: The genus Gliocladiopsis was introduced by
Saksena (1954) to accommodate a soil-born species G.
sagariensis. Liu and Cai (2013) introduced a new Gliocladiopsis species, G. guangdongensis, which was the first
isolation from freshwater habitats in China. Hyde et al.
(2018) introduced the second Gliocladiopsis taxon from
freshwater habitats in Thailand.
Gliocladiopsis aquaticus Lu et al., Mycosphere 9: 387
(2018)
Distribution: Thailand, Chiang Rai Province, Mae Fah
Luang University, on submerged decaying wood in a
freshwater lake in campus (Hyde et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–1976, paratype MFLU
17–1977. ITS, LSU, SSU, his3 and b-tubulin sequence data
are available.
Gliocladiopsis guangdongensis F. Liu & L. Cai, Cryptog.
Mycol. 34: 235 (2013)
Distribution: China, Guangdong Province, Zhaoqing,
Dinghu Mountain, on submerged wood in a stream (Liu
and Cai 2013).
Sexual morph: Undetermined
Notes: Holotype HMAS 244829, other specimen collected from freshwater habitats HMAS 244830. ITS, LSU,
TEF1a, his3 and b-tubulin sequence data are available.
543
Mariannaea G. Arnaud, Bull. trimest. Soc. mycol. Fr. 68:
196 (1952)
Holomorph Descriptions and illustrations refer to Lombard et al. (2015) and Hu et al. (2017).
Type species: Mariannaea elegans G. Arnaud, Bull.
trimest. Soc. mycol. Fr. 68: 196 (1952)
Notes: The genus Mariannaea G. Arnaud ex Samson
was established by Samson (1974) with M. elegans (Corda)
Samson as type species. There are 21 records of Mariannaea in Index Fungorum (December 2018). Hu et al.
(2017) revisited the genus Mariannaea and accepted 15
species in this genus based on morphology and molecular
phylogeny. Species of Mariannaea were mostly reported
from terrestrial habitats (Samson 1974; Samson and Bigg
1988; Matsushima 1989; Samuels 1989; Samuels and
Seifert 1991; Tokumasu et al. 1994; Gräfenhan et al. 2011;
Zeng and Zhuang 2014; Lombard et al. 2015; Nonaka et al.
2015). Cai et al. (2010) firstly reported a Mariannaea
species from freshwater stream in northern Thailand. Hu
et al. (2017) introduced four new species for Mariannaea
and all of them were collected from freshwater habitats in
China.
Mariannaea aquaticola Kurniawati et al.
Distribution: Thailand, Chiang Rai Province, Amphoe
Mae Chan, on wood submerged in a freshwater stream (Cai
et al. 2010).
Sexual morph: Undetermined
Notes: Holotype MFLU 09–0223, other specimens collected from freshwater habitats: MFLU 09–0224, MFLU
09–0225. ITS and LSU sequence data are available.
Mariannaea chlamydospora D.M. Hu & L. Cai
Distribution: China, Hubei Province, Shennongjia Scenic Area, Yakou Stream, on submerged wood (Hu et al.
2017).
Sexual morph: Undetermined
Notes: Holotype HMAS 245222. ITS, LSU and btubulin sequence data are available.
Mariannaea cinerea D.M. Hu & L. Cai
Distribution: China, Yunnan Province, Mengla county,
Dashaba reservoir, on submerged wood (Hu et al. 2017).
Sexual morph: Undetermined
Notes: Holotype HMAS 245224. ITS, LSU and btubulin sequence data are available.
Mariannaea fusiformis D.M. Hu & L. Cai
Distribution: China, Hubei Province, Shennongjia Scenic Area, Yakou Stream, on submerged wood (Hu et al.
2017).
Sexual morph: Undetermined
Notes: Holotype HMAS 245223. ITS, LSU and btubulin sequence data are available.
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Fungal Diversity (2019) 99:451–660
Fig. 37 Stachybotrys chartarum (MFLU 18–1637) a, b Colonies on wood. c–f Conidiophores with conidia. g, h Conidiogenous cells. i–
o Conidia. p Germinating conidium. q, r Culture on MEA from above and reverse. Scale bars: c 100 lm, d–g 50 lm, h, i, p 10 lm, j–o 5 lm
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Mariannaea lignicola D.M. Hu & L. Cai
Distribution: China, Jiangxi Province, Longnan County,
Jiulianshan Mountain Natural Reserve, Xiagongtang
Stream, on submerged wood (Hu et al. 2017).
Sexual morph: Undetermined
Notes: Holotype HMAS 245225. ITS, LSU and btubulin sequence data are available.
Mariannaea samuelsii Seifert & Bissett
Distribution: China, Yunnan Province, Dali, on submerged wood in a stream (This study)
Sexual morph: Undetermined
Notes: Holotype DAOM 235814, other specimen collected from freshwater habitats: MFLU 18–2328. ITS,
LSU, SSU, RPB2 and TEF1a sequence data are available.
Mariannaea superimposita (Matsush.) Samuels
Distribution: China, Yunnan Province, Jinsha river, on
submerged wood (This study)
Sexual morph: Undetermined
Notes: Holotype Cultura in b/c desiccata, MFC-1409,
specimen collected from freshwater MFLU 18–1632
(HKAS 92903). ITS and LSU sequence data are available.
Nectria (Fr.) Fr., Summa veg. Scand., Sectio Post.
(Stockholm): 387 (1849)
Holomorph
Descriptions
and
illustrations
see
Maharachchikumbura et al. (2016).
Type species: Nectria cinnabarina (Tode) Fr., Summa
veg. Scand., Summa veg. Scand., Sectio Post. (Stockholm):
388 (1849)
Notes: Fries (1849) introduced the genus Nectria based on
Hypocrea sect. Nectria Fr. Since then many species have been
added to the genus. Based on morphological characteristics
such as structure of ascomata and asexual morphs (Hirooka
et al. 2010; Rossman et al. 1999) and LSU sequence data
(Rehner and Samuels 1995). Nectria sensu lato was separated
into a number of genera. Hirooka et al. (2012) recently revised
Nectria, recognising 29 species within the genus.
Nectria chaetopsinae Samuels
Distribution: Thailand, Nakorn Ratchassima Province,
on submerged Xylia dolabriformis baits in a stream (Sivichai et al. 2000a).
Asexual morph: Undetermined
Notes: Holotype PDD 44237. Sequence data is not
available. Sivichai et al. (2000a) found Nectria chaetopsinae from freshwater in Thailand, but did not provide
description and illustration for this species.
Nectria curta J. Webster
Distribution: UK, on submerged wood of Fraxinus
excelsior (Webster 1993).
Asexual morph: Flagellospora sp. (Webster 1993).
Notes: Holotype IMI 353882. ITS and SSU sequence
data are available. This species is originally collected from
freshwater in UK (Webster 1993).
545
Nectria discophora Mont
Distribution: UK, on submerged wood in River Teign
(Shearer and Webster 1991).
Asexual morph: Undetermined
Notes: ITS, LSU, SSU and RPB2 sequence data are
available. Shearer and Webster (1991) found this species
from a freshwater River in UK, but they did not provide
descriptions and illustrations.
Nectria lugdunensis J. Webster
Distribution: UK, on submerged wood in River Teign
(Shearer and Webster 1991).
Sexual morph: Undetermined
Notes: Holotype IMI 74951. ITS, LSU and SSU
sequence data are available. Shearer and Webster (1991)
first found this species from freshwater habitats.
Neocosmospora E.F. Sm., U.S.D.A. Div. Veg. Pathol. Bull.
17: 45 (1899)
Holomorph Descriptions and illustrations refer to Lombard et al. (2015).
Type species: Neocosmospora vasinfecta E.F. Sm., Bull.
U.S. Department of Agriculture 17: 45 (1899)
Notes: The genera Haematonectria (1999), Lachnidium
(1891) and Neocosmospora (1899) could be applied to this
group of fungi (Rossman et al. 1999; Summerbell and
Schroers 2002; Lombard et al. 2015). The generic name
Neocosmospora was recommendated for these fungi by
Lombard et al. (2015) as the generic name Lachnidium is
based on a nomen confusum (Madelin 1966; Kendrick
1974), and can therefore not be used and in addition, the
generic name Neocosmospora (1899) is older than the
generic name Haematonectria (1999).
Neocosmospora haematococca (Berk. & Broome)
Samuels, Nalim & Geiser
Distribution: China, Hong Kong, on submerged wood
(Tsui et al. 2001b, Tsui and Hyde 2004)
Asexual morph: Undetermined
Notes: ITS, LSU, SSU and RPB2 are available. Neocosmospora haematococca was found from freshwater in
China, but without descriptions and illustration (Tsui et al.
2001b; Tsui and Hyde 2004).
Paracremonium L. Lombard & Crous, Stud. Mycol. 80:
233 (2015)
Asexual morph Descriptions and illustrations see Lombard et al. (2015) Sexual morph Undetermined.
Type species: Paracremonium inflatum L. Lombard &
Crous, Stud. Mycol. 80: 233 (2015)
Notes: The genus Paracremonium was recently established for different strains from a group of fungi previously
treated as Acremonium recifei (Lombard et al. 2015). Five
species are currently accepted in Paracremonium; P. binnewijzendii, P. contagium, P. inflatum, P. pembeum, and P.
variiforme. Paracremonium binnewijzendii was isolated
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Fungal Diversity (2019) 99:451–660
Fig. 38 a Stachybotrys chlorohalonata (MFLU 18–1638) a, b Colonies on wood. c, d Conidiophores with conidia and conidiogenous cells. e–
m Conidia. n Germinating conidium. o, p Culture on MEA from above and reverse. Scale bars: c, d 50 lm, e–n 15 lm
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from stream embankments (The Netherlands), while P.
contagium (Canada) and P. inflatum (India, Colombia)
were associated with human infections and P. pembeum
with trees (Acer negundo, Persea americana, Platanus
racemosa, Ricinus communis) and heads of Euwallacea sp.
(California, USA), P. variiforme was collected from water
in an unnamed karst cave (China) (Lombard et al. 2015;
Lynch et al. 2016; Crous et al. 2017; Zhang et al. 2017a, b).
In this study, we introduce a new species P. aquaticum and
a new record of P. binnewijzendii for China, both of them
are collected from freshwater habitats.
Paracremonium binnewijzendii Houbraken, van der Kleij
& L. Lombard
Distribution: China, Yunnan Province, Erhai Lake, on
submerged wood (This study)
Sexual morph: Undetermined
Notes: Holotype CBS H-23246, specimen collected
from freshwater habitats: MFLU 18–1636. ITS, LSU and
b-tubulin are available.
Payosphaeria W.F. Leong, Bot. Mar. 33: 511 (1990)
Asexual morph Undetermined. Sexual morph Ascomata
globose to pyriform, superficial, papillate, ostiolate, lacking periphyses, membranous, hyaline, solitary to gregarious. Neck cylindrical, stout, hyaline. Peridium composed
of a single layer of thin-walled elongate cells. Paraphyses
branched, septate and hyaline. Asci unitunicate, 8-spored,
long-cylindrical, short pedicellate, persistent, thin-walled
and developing at the base of the ascoma. Ascospores
uniseriate, round to oval, aseptate, hyaline, thin-walled,
smooth.
Type species: Payosphaeria minuta H.Y.M. Leung, Bot.
Mar. 33: 511 (1990)
Notes: The monotypic genus Payosphaeria was introduced by Leong et al. (1990) with the type species P.
minuta Leong. Payosphaeria minuta is a common mangrove lignicolous species reported worldwide (Leong et al.
1990; Chinnaraj 1993; Ravikumar and Vittal 1996). The
only report of P. minuta from freshwater habitats was
provided by Cai et al. (2006a).
Payosphaeria minuta H.Y.M. Leung
Distribution: China, Yunnan Province, on submerged
bamboo culm (Cai et al. 2006a).
Asexual morph: Undetermined.
Notes: Holotype IMI 327472. Sequence data is not
available. The species Payosphaeria minuta was originally
collected from mangrove wood in Singapore. Cai et al.
(2006a) reported this species from freshwater in China
without descriptions and illustration.
Varicosporella Lechat & J. Fourn., Ascomycete.org 7(1): 2
(2015)
Asexual morph Fusarium like, macroconidia cylindrical,
slightly curved, acute at tip, truncate to rounded at base,
547
septate. Sexual morph Ascomata nonstromatic, solitary,
superficial with base slightly immersed in substratum, softtextured, greyish yellow to pale orange, becoming pale
yellow in 3% KOH and in lactic acid, obpyriform, laterally
collapsing when dry, uniloculate, translucent, with a
broadly conical to rounded apex, composed of cylindrical
yellow cells, thick-walled, septate, clavate at top. Ascomatal wall composed of subglobose to angular thick-walled
cells, becoming more flattened inwardly. Perithecial surface cells forming a textura angularis in surface view.
Basal hyphae sparse and short, thick-walled, hyaline. Asci
8-spored, unitunicate, cylindrical, short-stipitate, apically
truncate to slightly rounded with a conspicuous, refractive
apical apparatus. Periphyses copious, embedded in gel
matrix, simple or branched, septate. Ascospores obliquely
uniseriate, ellipsoid with narrowly to broadly rounded ends,
equally two-celled, slightly constricted at septum, hyaline
to pale yellowish brown, with two large guttules in each
cell, wall roughened by short, sinuous, brown, thick ribs,
sometimes anastomosed. Hyaline ascospores germinate
more often than pigmented ones.
Type species: Varicosporella aquatica Lechat & J.
Fourn., Ascomycete.org 7(1): 2 (2015)
Notes: The genus Varicosporella was introduced by
Lechat and Fournier (2015) with single species Varicosporella aquatica which was collected from freshwater
habitats in southern France.
Varicosporella aquatica Lechat & J. Fourn
Distribution: France, Ariège and Lozère, on submerged
twig of Buxus sempervirens, Salix sp. and Alnus glutinosa
(Lechat and Fournier 2015).
Asexual morph: see Lechat and Fournier (2015).
Notes: Holotype LIP, Fournier 09197. ITS and LSU
sequence data are available.
Niessliaceae Kirschst.
Paraniesslia Tsui et al., Mycologia 93(5): 1002 (2001)
Asexual morph Undetermined. Sexual morph Ascomata
superficial, pyriform to subglobose, papillate, with setae,
ostiolate, periphysate, solitary to gregarious. Setae acute,
straight, unbranched, septate, black. Peridium membranous, textura angularis in longitudinal section, textura
epidermoidea in surface view. Paraphyses septate, thinwalled, deliquescent. Asci 8-spored, unitunicate, clavate,
pedicellate, with a nonamyloid discoid refractive apical
ring. Ascospores overlapping uniseriate to biseriate, aseptate, verrucose, with or without a mucilaginous sheath.
Type species: Paraniesslia tuberculata Tsui et al.,
Mycologia 93(5): 1002 (2001)
Notes: Paraniesslia was established to accommodate a
fungus collected from freshwater habitats in Hong Kong,
China (Tsui et al. 2001d). Cai and Hyde (2007) introduced
123
548
the second species Paraniesslia aquatica which was collected from freshwater habitats.
Paraniesslia aquatica L. Cai & K.D. Hyde
Distribution: China, Yunnan Province, Kunming city,
Qinglongxia, submerged wood in a small stream (Cai and
Hyde 2007).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 10856 (now in IFRD).
Sequence data is not available. Paraniesslia aquatica was
introduced by Cai and Hyde (2007) and only known from
the type locality.
Paraniesslia tuberculata Tsui et al.
Distribution: China, Hong Kong, on submerged wood
(Tsui et al. 2001d).
Asexual morph: Undetermined
Notes: Holotype IFRD 8827; Sequence data is not
available.
Stachybotryaceae L. Lombard & Crous
Koorchalomella Chona et al., Indian Phytopath. 11: 130
(1958)
Asexual morph Hyphomycetous. Conidiomata sporodochial, scattered or gregarious, superficial, raised, convex to
pulvinate, circular, oval to elongate in outline. Conidiophores unbranched, aseptate, hyaline to apricot in color,
smooth-walled, terminating in one conidiogenous cell.
Conidiogenous cells phialides, arranged in a dense palisade, clavate with truncate base, hyaline to apricot in
color. Conidia fusiform, unicellular, rarely one-septate,
hyaline to apricot to salmon in color, smooth-walled.
Sexual morph Undetermined.
Type species: Koorchalomella oryzae Chona et al.,
Indian Phytopath. 11: 130 (1958)
Notes: Koorchalomella is a monotypic genus established to accommodate K. oryzae which was collected from
dead culms of Oryzae sativa L. from damp ground in
Bangalore, India. Subsequently, two species were introduced in this genus and one of them collected from
freshwater habitats (Hyde et al. 2017).
Koorchalomella salmonispora Abdel-Aziz & AbdelWahab
Distribution: Egypt, Sohag City, on submerged decayed
stem of Phragmites australis (Hyde et al. 2017).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1512. LSU sequence data is
available.
Stachybotrys Corda, Icon. fung. (Prague) 1: 21 (1837)
Holomorph Description see Lombard et al. (2016).
Type species: Stachybotrys chartarum (Ehrenb.) S.
Hughes, Can. J. Bot. 36: 812 (1958)
Notes: The genus Stachybotrys was established by
Corda (1837) and is the type genus of the family Stachybotryaceae. Lombard et al. (2016) revised all the
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Stachybotrys-like taxa by combining morphology and
multi-locus phylogenetic analyses using cmdA, ITS, RPB2,
TEF1a and b-tubulin dataset, and 33 genera were accepted
in the family Stachybotryaceae. The genus Melanopsamma
was resurrected by Lombard et al. (2016) which was synonymised under Stachybotrys by Wang et al. (2015).
Lombard et al. (2016) excluded S. queenslandica from the
genus Stachybotrys due to the lacking of living type
material to determine its phylogenetic placement, and
noted that it might belong to the genus Melanopsamma.
Stachybotrys chartarum (Ehrenb.) S. Hughes
: Stilbospora chartarum Ehrenb., Sylv. mycol. berol.
(Berlin): 21 (1818)
Facesoffungi number: FoF 01247, Fig. 37
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, dark brown to
black, in groups. Conidiophores 41–65 lm long, 3–5 lm
wide (
x = 53 9 4 lm, n = 20), macronematous, mononematous, single or in groups, thin-walled, branched, erect,
straight to slightly flexuous, subhyaline to olivaceous
brown, septate, smooth, bearing 3–6 conidiogenous cells.
Conidiogenous cells phialidic, clavate to subclavate, subhyaline to olivaceous brown, smooth, with conspicuous
collarettes. Conidia 8–10 lm long, 5–7 lm wide
(
x = 9 9 6 lm, n = 20), acrogenous, ellipsoidal to subcylindrical, verrucose, aseptate, thick-walled, olivaceous
brown to dark brown, rounded at both ends. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Erhai
Lake, saprobic on submerged decaying wood, June 2015,
L.W. Wang, S-549 (MFLU 18–1637, ibid HKAS 92619),
living culture MFLUCC 16–0966; Jinsha River, saprobic
on submerged decaying wood, April 2015, Z.L. Luo,
S-370, living culture MFLUCC 16–1337; Cangshan
Mountain, saprobic on decaying wood submerged in a
freshwater stream, September 2014, X.J. Su, S-223, living
culture MFLUCC 16–1302; Ibid., saprobic on decaying
wood submerged in a freshwater stream, April 2016, X.J.
Su, S-761, living culture MFLUCC 17–1322.
Distribution: China, Yunnan Province, on submerged
wood in Rivers and streams (This study)
Sexual morph: Undetermined
Notes: Epitype CBS H-18496, lectotype DAOM 51026,
specimens collected from freshwater habitats: MFLU
18–1637, HKAS 92619. cmdA, ITS, LSU, SSU, RPB2,
TEF1a and b-tubulin sequence data are available. During
our studies on liginicolous freshwater fungi of northwestern Yunnan Province, three isolates were obtained from
submerged wood in freshwater. The morphology of our
fungus fits well with Stachybotrys species. Phylogenetic
analysis shows that our isolates cluster with Stachybotrys
chartarum with good bootstrap support (Fig. 1, clade 39).
Based on the morphology and phylogeny, we identify our
Fungal Diversity (2019) 99:451–660
549
123
550
b Fig. 39 Ascosacculus fusiformis (MFLU 15–0068, holotype) a Spec-
imen. b, c Appearance of necks on substrate. d Horizontal section
through ascomata. e Section through ascomata. f Structure of
peridium. g Paraphyses. h–k Asci. l–p Ascospore. q Germinating
ascospore. Scale bars: c 300 lm, d 200 lm, e 50 lm, h–k 30 lm, g,
f 20 lm, l–q 10 lm
fungus as S. chartarum. Descriptions and illustration are
provided and this is the first report of S. chartarum from
freshwater habitats.
Stachybotrys chlorohalonata B. Andersen & Thrane
Facesoffungi number: FoF 05447, Fig. 38
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, dark brown to
black, in groups. Conidiophores 117–264 lm long, 3.5–
4.5 lm wide (
x = 175 9 4 lm, n = 20), macronematous,
mononematous, single or in groups, thin-walled, branched,
erect, straight to slightly flexuous, subhyaline at base, often
olivaceous brown toward the apex, and the upper portion is
sometimes irregularly verrucose, septate, bearing 3–6
conidiogenous cells. Conidiogenous cells phialidic, clavate
to subclavate, subhyaline to olivaceous brown, smooth,
with conspicuous collarettes. Conidia 8–10 lm long, 7–
9 lm wide (
x = 9 9 8 lm, n = 20), acrogenous, ellipsoidal to obovoid, or subglobose, aseptate, blackish green
or dark brown, opaque and verrucose. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Jinsha
River, saprobic on submerged decaying wood, April 2015,
Q. Dai, S-488 (MFLU 18–1638, HKAS 93029), living
culture MFLUCC 16–1338.
Distribution: China, Yunnan Province, on submerged
wood in Jinsha River (This study).
Sexual morph: Undetermined
Notes: Holotype C 60160, specimens collected from
freshwater habitats: MFLU 18–1638, HKAS 93029. cmdA,
ITS, LSU, SSU, RPB2, TEF1a and b-tubulin sequence data
are available. Morphological characters and phylogenetic
analysis evidence identify our isolate as Stachybotrys
chlorohalonata and it is the first report from freshwater
habitats.
Hypocreales genera incertae sedis
Emericellopsis J.F.H. Beyma, Antonie van Leeuwenhoek
6: 264 (1940) [1939]
Asexual morph Acremonium-like. Conidiophores mostly
simple orthotropic. Conidiogenous cells phialidic, terminal
or lateral, smooth-walled, hyaline. Conidia narrowly
ellipsoid, hyaline, smooth, adhering in slimy heads. Chlamydospores absent or mostly intercalary and hyaline.
Sexual morph Ascomata superficial on the substratum,
globose, non-ostiolate, dark brown. Peridium multi-
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layered, pseudoparenchymatous, composed of several layers of compressed cells. Asci unitunicate, saccate, with thin
deliquescent wall, soon dissolving, scattered irregularly in
the ascocarp. Ascospores ellipsoid, pale brown, with
uneven surfaces, surrounded by 3 or 5 longitudinal, subhyaline or hyaline appendages.
Type species: Emericellopsis terricola J.F.H. Beyma,
Antonie van Leeuwenhoek 6: 265 (1940) [1939]
Notes: The genus Emericellopsis was introduced by
Beyma (1940) and placed in Eurotiaceae based on the type
species E. terricola Beyma. Grum-Grzhimaylo et al. (2013)
placed this genus in Bionectriaceae, but later
Maharachchikumbura et al. (2016) treated it as Hypocreales genera incertae sedis. The known Emericellopsis
species were reported from soil, marine, soda lake and
terrestrial habitats (Stolk 1955; Luppi-Mosca 1960; Borut
& Johnson 1962; Park 1972; Grum-Grzhimaylo et al.
2013). Among these Emericellopsis species, only E. terricola has been found in freshwater in Ireland and Japan
(Park 1972; Tubaki 1973).
Emericellopsis terricola J.F.H. Beyma
Distribution: Ireland, on submerged organic debris
(Park 1972); Japan, in freshwater muds of Lake Sengari
(Tubaki 1973).
Asexual morph: Cephalosporium sp.
Notes: ITS, LSU, SSU, RPB2 and b-tubulin sequence
data are available.
Microascales Luttr. ex Benny & Kimbr.
Halosphaeriaceae E. Müll. & Arx, ex Kohlm
Aniptodera Shearer & M.A. Mill., Mycologia 69(5): 893
(1977)
Asexual morph Undetermined. Sexual morph Ascomata
globose or subglobose, immersed or superficial, ostiolate,papillate, membranaceous, hyaline to light brown.
Necks cylindrical, periphysate, brown at the tip. Paraphyses thin-walled cells filling venter of young ascomata,
breaking up into catenophyses. Asci 8-spored, unitunicate,
clavate, short pendunculate, thin-walled except for a thickwalled area below the apex, flattened and refractive at the
tip, provided with a simple pore, slightly constricted below
the apex, relatively persistent even af developing at the
base of the ascomata venter. Ascospores ellipsoidal, 1–3septate, hyaline, thick without appendages.
Type species: Aniptodera chesapeakensis Shearer &
M.A. Miller, Mycologia 69(5): 894 (1977)
Notes: The genus Aniptodera was established by
Shearer and Miller (1977) with A. chesapeakensis Shearer
& M.A. Miller as the type species. The genus is charactized
by having hyaline or light coloured ascomata, catenophyses, apically thickened persistent asci with a distinct pore
and subapical retraction of cytoplasm, and hyaline, thick-
Fungal Diversity (2019) 99:451–660
walled, 1-septate ascospores with or without appendages
(Shearer and Miller 1977; Raja and Shearer 2008).
Aniptodera aquadulcis (Hsieh et al.) Campb. et al.
Distribution: China, Taiwan, Taipei, on decayed wood
submerged in freshwater (Hsieh et al. 1995); China, Hong
Kong, Tai Po, Lam Tsuen River, on submerged wood
(Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype WL0213 (Academia Sinica herbarium).
Sequence data is not available.
Aniptodera aquibella J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, Kaeng Krachan, near Pala-U Waterfall, on submerged wood in a stream (Li et al. 2016a).
Asexual morph: Undetermined
Notes: Holotype MFLU 15–1140. LSU and SSU
sequence data are available. This species was introduced by
Li et al. (2016a, b) and only known from the type locality.
Aniptodera chesapeakensis Shearer & M.A. Mill.
Distribution: Australia, north Queensland, Crystal
Cascades, on submerged wood (Hyde et al. 1999a); Brazil,
Bahia, Santa Terezinha, on submerged twig (Barbosa et al.
2013); Brunei, Tutong River, on submerged wood (Fryar
et al. 2004); China, Hong Kong, Tai Po Kau forest stream,
on submerged wood (Hyde et al. 1999a); India, north
Maharashtra, on submerged wood (Patil and Borse 2012);
Iraq, Shatt al-arab River, on submerged dead stem of
Arundo donax and Phragmites australis (Al-Saadoon and
Al-Dossary 2014); Japan, on submerged wood (Minoura
and Muroi 1978); Mauritius, near Tamarin, Black River,
on submerged wood (Hyde et al. 1999a); South Africa,
Durban, Palmiet River, on submerged wood (Hyde et al.
1998b); USA, Maryland, on submerged wood; Illinois, on
submerged wood (Shearer and Miller 1977; Shearer and
Crane 1986; Shearer 1989b).
Asexual morph: Undetermined
Notes: Holotype ILLS 36523, other specimens collected
from freshwater habitats ILLS 36506–36514, ILLS 31524.
ITS, LSU, SSU, RPB1, RPB2 and TEF1a sequence data
are available. Shearer and Miller (1977) firstly collected
this species from submerged wood in freshwater. This
species have been found from freshwater habitats
worldwide.
Aniptodera fusiformis Shearer
Distribution: Iraq, Al-Kahlaa River, on submerged stem
(Al-Saadoon and Al-Dossary 2014); USA, New York, on
cottonwood twigs (Populus deltoides) submerged in Jordan
Creek (Shearer 1989b).
Asexual morph: Undetermined
Notes: Holotype NY-00966728, isotype: NY-00966729.
Sequence data is not available.
Aniptodera inflatiascigera Tsui et al.
551
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Tsui et al. 1997; Hyde et al.
1999a); Philippines, Negros Occidentalis, Bario Alegria,
Lupit River, on submerged wood (Tsui et al. 1997; Hyde
et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype (HKU(M) 4682 (now in IFRD).
Sequence data is not available.
Aniptodera lignatilis K.D. Hyde
Distribution: Australia, on submerged log in freshwater
(Hyde 1992c); China, Hong Kong, on submerged wood in
Tai Po Kau forest stream (Ho et al. 2001); Philippines,
Luzon, Laguna, Los BaSos, Mt Maquiling, on wood submerged in a freshwater stream (Hyde et al. 1999a); Seychelles, Mahe, Riveire St Mare Louise, on submerged
wood (Hyde et al. 1999a); South Africa, Durban, Palmiet
River, on submerged wood (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype BRIP 17156. LSU sequence data is
available.
Aniptodera lignicola Hyde et al.
Distribution: Australia, Crystal Cascades, on submerged wood (Hyde et al. 1999a); Brunei, Temburong,
Kuala Belalong Field Studies Centre, Sungai Esu, on wood
submerged in a stream (Hyde et al. 1999a); India, north
Maharashtra, on submerged wood (Patil and Borse 2012);
Malaysia, State Negara, Lipur Lentang Nature Reserve, on
submerged wood (Hyde et al. 1999a); Philippines, Luzon,
Laguna, Los Bados, Mt. Maquiling, on wood submerged in
a freshwater stream (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2863 (now in IFRD).
Sequence data is not available.
Aniptodera limnetica Shearer
Distribution: USA, Illinois, on submerged wood in Wolf
Lake (Shearer 1989b).
Asexual morph: Undetermined
Notes: Holotype NY-00966731, isotype: NY-00966732.
Sequence data is not available. This species is known only
from the type locality.
Aniptodera margarition Shearer
Distribution: UK, Cornwall, on unidentified woody
debris submerged in a small stream (Shearer 1989b); USA,
New York/Illinois, on a twig collected from the Sangamon
River (Shearer 1989b).
Asexual morph: Undetermined
Notes: Holotype NY-00966730, other specimens collected from freshwater habitats: NY-2946622, NY2946629. Sequence data is not available.
Aniptodera mauritaniensis Hyde et al.
Distribution: Iraq, Shatt Al-Arab River and Al-Kahlaa
River, on submerged wood (Al-Saadoon and Al-Dossary
123
552
2014); Mauritius, near Tamarin, Black River, on submerged wood (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype IFRD 8646. Sequence data is not
available. This species is only known from the type
locality.
Aniptodera megaloascocarpa Raja & Shearer
Distribution: USA, Florida, Ocala National forest, on
submerged decorticated woody debris (Raja and Shearer
2008).
Asexual morph: Undetermined
Notes: Holotype ILL 40109. Sequence data is not
available. This species is only known from the type
locality.
Aniptodera megalospora Hyde et al.
Distribution: Brunei, Temburong, Kuala Belalong Field
Studies Centre, Sungai Esu, on wood submerged in a
stream (Hyde et al. 1999a); China, Hong Kong, on submerged wood (Hyde et al. 1999a); Malaysia, State Negara,
Lipur Lentang Nature Reserve, on submerged wood (Hyde
et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2885 (now in IFRD).
Sequence data is not available.
Aniptodera palmicola Hyde et al.
Distribution: Iraq, Shatt Al-Arab River and Al-Kahlaa
River, on submerged wood (Al-Saadoon and Al-Dossary
2014); South Africa, Kwa Zulu-Natal, Mt Unzini, The
National Monument, on submerged rachis of Raphia australia (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype IFRD 8647. Sequence data is not
available. This species is only known from the type
locality.
Ascosacculus Campb. et al., Mycologia 95(3): 545 (2003)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell et al. (2003b).
Type species: Ascosacculus aquaticus (K.D. Hyde)
Campb. et al., Mycologia 95(3): 545 (2003)
Notes: The genus Ascosacculus was introduced by
Campbell et al. (2003b) to accommodate two Halosarpheia
species which were originally introduced as Halosarpheia
aquatica and H. heteroguttulata. Both species of Ascosacculus have been reported from freshwater habitats
worldwide (Hyde et al. 1999a; Campbell et al. 2003b).
Ascosacculus aquaticus (K.D. Hyde) Campb. et al.
: Halosarpheia aquatica K.D. Hyde, Aust. Syst. Bot.
5(4): 407 (1992)
Distribution: Australia, north Queensland, Atherton
Tablelands, Clohesy River, on submerged wood (Hyde
et al. 1999a).
Asexual morph: Undetermined
123
Fungal Diversity (2019) 99:451–660
Notes: Holotype BRIP 19331, other specimen collected
from freshwater habitats: BRIP 19384. LSU sequence data
is available. This species is only known from the type
locality.
Ascosacculus heteroguttulatus (Wong et al.) Campb. et al.
: Halosarpheia heteroguttulata Wong et al., Can.
J. Bot. 76(11): 1858 (1999) [1998]
Distribution: Australia, north Queensland, Crystal
Cascades, on submerged wood (Hyde et al. 1999a); Brunei, Temburong, Kuala Belalong Field Studies Centre,
Sungai Esu, on wood submerged in a stream (Hyde et al.
1999a); China, Hong Kong, New Territories, Plover Cove
Reservoir, on submerged wood (Hyde et al. 1999a);
Mauritius, Tamarin, Black River, on submerged wood
(Hyde et al. 1999a); Philippines, Mindanao, Bukidnon,
Impalatao, Natigbasan Creek, on submerged wood (Hyde
et al. 1999a); South Africa, Durban, Palmiet River, on
submerged wood (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype IFRD 8762. LSU and SSU sequence
data are available.
Ascosacculus fusiformis Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555667, Facesoffungi
number: FoF 05448, Fig. 39
Etymology: Referring to the fusiform ascospores of this
fungus
Holotype: MFLU 15–0068
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Ascomata 175–205 lm high, 85–105 lm diam., immersed
with neck erumpent through host surface, subglobose to
ellipsoid, solitary, dark brown to black. Ostiole central,
with straight upright, black, long neck, periphysate.
Peridium 7–11 lm thick, composed of 2–3 layers of
brown, thin-walled, angular cells, which are hyaline
inwardly. Paraphyses 4–6 lm wide, hyaline, unbranched,
septate, slightly constricted at the septum. Asci 127–
159 9 10–14 lm (
x = 143 9 12 lm, n = 10), 8-spored,
unitunicate, cylindrical, apically rounded, pedicellate, with
a bilateral apical ring. Ascospores 21–23 9 8–10 lm
(
x = 22 9 9 lm, n = 20), uni-seriate, fusiform, 3-septate,
slightly constricted at the septa, with 2 prominent guttules,
hyaline, smooth-walled, with a thin mucilaginous sheath.
Material examined: THAILAND, Chiang Mai Province,
saprobic on decaying wood submerged in a freshwater
stream, November 2013, Z.L. Luo, ZL-9 (MFLU 15–0068,
holotype), ex-type living culture MFLUCC 14–0036.
Notes: Ascosacculus fusiformis resembles A. aquaticus
and A. heteroguttulatus in having immersed, solitary, dark
brown ascomata, central ostiole with long, black neck, thinwalled peridium of textura angularis in surface view and
hyaline, fusiform, guttulate ascospores (Hyde 1992b).
Fungal Diversity (2019) 99:451–660
However, A. fusiformis differs from A. aquaticus in having
cylindrical, pedicellate asci with a bilateral apical ring and
shorter (21–23 lm vs. 33.5–64 lm), 3-septate ascospores
with a thin mucilaginous sheath (Hyde 1992b). A. fusiformis differs from A. heteroguttulatus in having thinner
peridium
(7–11 lm
vs.
20–28 lm),
smaller
(21–23 9 8–10 lm vs. 27–37 9 9–17.5 lm), 3-septate
ascospores with a thin mucilaginous sheath. Phylogenetically, Ascosacculus fusiformis clusters with other Ascosacculus species in good support with separate clade
(Fig. 1, clade 40). Ascosacculus fusiformis also shares
similar morphology with Annulatascus species in having
cylindrical, pedicellate asci with a bilateral apical ring and
hyaline, fusiform, septate, guttulate ascospores with a thin
mucilaginous sheath. However, phylogenetic analysis
based on LSU, SSU, RPB2 and TEF1a sequence data
shows that Ascosacculus belongs in Halosphaeriaceae
(Microascales), while Annulatascus belongs to Annulatascaceae (Annulatascales) (Fig. 1, clade 17, 40).
Fluviatispora K.D. Hyde, Mycol. Res. 98(7): 720 (1994)
Asexual morph Undetermined. Sexual morph Ascomata
globose, immersed in wood, subglobose or ellipsoidal,
light-brown, ostiolate, membraneous and solitary. Asci
8-spored, unitunicate, clavate or saccate, thin-walled,
pedunculate and deliquescing early. Ascospores 2–3-seriate, unicellular, ellipsoidal, hyaline. with a mucilaginous
sheath.
Type species: Fluviatispora tunicata K.D. Hyde, Mycol.
Res. 98(7): 722 (1994)
Notes: Fluviatispora was introduced by Hyde (1994) to
accommodate two taxa, F. reticulata and F. tunicata
(Halosphaeriaceae) which were collected from freshwater
habitats. The genus is characterised by hyaline, unicellular
ascospores surrounded by a mucilaginous sheath, thinwalled and clavate to saccate, early deliquescing unitunicate asci and immersed thin-walled ascomata (Hyde 1994).
Three species are accepted in this genus and two of them
have been reported from freshwater habitats.
Fluviatispora reticulata K.D. Hyde
Distribution: Papua New Guinea, Western Province,
Bensbach, Bensbach River, on frons submerged Livistona
sp. (Hyde 1994).
Asexual morph: Undetermined
Notes: Holotype BRIP 21392; Sequence data is not
available. This species is only known from the type
locality.
Fluviatispora tunicata K.D. Hyde
Distribution: Papua New Guinea, Western Province,
Bensbach, Bensbach River, on frons submerged Livistona
sp. (Hyde 1994).
Asexual morph: Undetermined
553
Notes: Holotype BRIP 21391. Sequence data is not
available. This species is only known from the type
locality.
Luttrellia Shearer, Mycologia 70(3): 692 (1978)
Asexual morph Undetermined. Sexual morph Ascomata
partially immersed to superficial, globose to subglobose,
membranous. Paraphyses with catenophyses, septate,
hyaline. Asci clavate to cylindrical, early deliquescent,
thin-walled. Ascospores multiseptate, lacking appendages
accumulating in a mass at tip of neck after discharge,
hyaline.
Type species: Luttrellia estuarina Shearer, Mycologia
70(3): 693 (1978)
Notes: The genus Luttrellia (Halosphaeriaceae) was
established by Shearer (1978) to accommodate
a single species, L. estuarina. Ferrer and Shearer (2007)
introduced three Luttrellia species from temperate and
tropical freshwater habitats. Index Fungorum (2018) lists
eight epithets of Luttrellia and four of them have been
reported from freshwater habitats.
Luttrellia estuarina Shearer
Distribution: Canada, Manitoba, Rennie River, on
submerged decorticated wood (Ferrer and Shearer 2007);
USA, Maryland, Patuxent River (Shearer 1978); Corolado,
Florida, Illinois, Maryland, Minnesota, North Carolina, on
submerged decorticated wood in a small stream (Ferrer and
Shearer 2007).
Asexual morph: Undetermined
Notes: Holotype ILLS 36979. Sequence data is not
available.
Luttrellia guttulata A. Ferrer & Shearer
Distribution: Costa Rica, Heredia, on submerged
decorticated wood (Ferrer and Shearer 2007); Panama,
Barro Colorado National Monument, on submerged
decorticated wood in Wheeler Stream (Ferrer and Shearer
2007).
Asexual morph: Undetermined
Notes: Holotype ILL, AF181–1. Sequence data is not
available.
Luttrellia halonata A. Ferrer & Shearer
Distribution: Ecuador, Yasuni National Park, Tiputini
River, on submerged decorticated wood (Ferrer and
Shearer 2007).
Asexual morph: Undetermined
Notes: Holotype ILL, AF134–1. Sequence data is not
available.
Luttrellia parvulospora A. Ferrer & Shearer
Distribution: French Guiana, Degrad Eskol, Crique
Gabrielle, Commune Noura, on submerged decorticated
wood (Ferrer and Shearer 2007); USA, Mississippi/Lousiana, on submerged decorticated wood (Ferrer and Shearer
2007).
123
554
Asexual morph: Undetermined
Notes: Holotype ILL, Robertson A-276–5. Sequence
data is not available.
Magnisphaera Campb. et al., Mycologia 95(3): 546 (2003)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell et al. (2003b).
Type species: Magnisphaera spartinae (E.B.G. Jones)
Campb. et al., Mycologia 95(3): 547 (2003)
Notes: The genus Magnisphaera was introduced by
Campbell et al. (2003b) to accommodate Halosarpheia
spartinae E.B.G. Jones and two undescribed isolates based
on molecular and morphological data. The genus Magnisphaera is characterized by large, black, globose to flattened
globose ascomata with a central neck that is short in proportion to the length of the ascoma. two-layered peridium
composed of about 8–10 large cells. The asci are ellipsoid,
thin-walled throughout, early deliquescent and lack an
apical pore and apical apparatus. The ascospores are
broadly acerose, phragmoseptate, constricted at the septa,
with a large guttule in each of the central cells (Campbell
et al. 2003b). Two species are accepted in this genus and
only Magnisphaera stevemossago was collected from
freshwater habitats.
Magnisphaera stevemossago J. Campb., J.L. Anderson &
Shearer
Distribution: USA, Colorado, Alaska, on decorticated
wood submerged in creek, pond and River (Campbell et al.
2003b).
Asexual morph: Undetermined
Notes: Holotype ILL, Shearer A409–1, other specimens
collected from freshwater habitats: ILL A409–3, ILL
A409–4, ILL A409–5. LSU, SSU and RPB2 sequence data
are available.
Nais Kohlm., Nova Hedwigia 4: 409 (1962)
Asexual morph Undetermined. Sexual morph Ascomata
immersed or semi-immersed, membranous, globose to
ampulliform, ostiole central, beaked, black, solitary or
gregarious. Peridium thin. Paraphyses absent. Catenophyses numerous. Asci 8-spored, saccate, deliquescent, thinwalled, pedunculate. Ascospores ellipsoid, bi-celled, not
constricted at the septa, with a band of refringent globules
around the equator, mostly with appendages, hyaline.
Type species: Nais inornata Kohlm, Nova Hedwigia 4:
409 (1962)
Notes: The genus Nais Kohlm was introduced by
Kohlmeyer (1962) with Nais inornata Kohlm as type
species. Subsequnetly, another two species N. glitra and N.
aquatica were introduced by Crane and Shearer (1986) and
Hyde (1992a) respectively. Pang et al. (2003) transferred
N. glitra to the genus Saagaromyces as Saagaromyces
glitra based on morphology and phylogenetic analysis.
There are two species accepted in Nais and both of them
123
Fungal Diversity (2019) 99:451–660
have been reported from freshwater habitats (Hyde et al.
1999a; Raja et al. 2009b).
Nais aquatica K.D. Hyde
Distribution: Australia, north Queensland, Millaa Millaa Falls, on submerged wood (Hyde et al. 1999a); China,
Hong Kong, New Territories, Plover Cove Reservoir, on
submerged wood in a stream (Hyde et al. 1999a); South
Africa, Durban, Palmiet River, on submerged wood (Hyde
et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype BRIP 17378. Sequence data is not
available.
Nais inornata Kohlm.
Distribution: USA, Florida, on submerged decaying
wood (Raja et al. 2009b).
Asexual morph: Undetermined
Notes: LSU sequence is available. Nais inornata was
introduced by Kohlmeyer (1962) and collected from rotten
wood in salt water. Raja et al. (2009b) reported this species
from freshwater habitats.
Natantispora Campb. et al., Mycologia 95(3): 543 (2003)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell et al. (2003b).
Type species: Natantispora retorquens (Shearer & J.L.
Crane) Campb. et al., Mycologia 95(3): 543 (2003)
Notes: The genus Natantispora was introduced by
Campbell et al. (2003b) to accommodate two species which
were originally introduced as Halosarpheia lotica and H.
retorquens. Both species of Natantispora have been
reported from freshwater habitats worldwide (Hyde et al.
1999a; Campbell et al. 2003b).
Natantispora lotica (Shearer) Campb. et al.
: Halosarpheia lotica Shearer, Mycotaxon 20(2): 505
(1984)
Distribution: Australia, north Queensland, Atherton
Tablelands, Clohesy River, on submerged wood (Hyde
et al. 1999a); China, Hong Kong, New Territories, Plover
Cove Reservoir, on submerged wood in a stream (Hyde
et al. 1999a); Mauritius, near Tamarin, Black River, on
submerged wood (Hyde et al. 1999a); Philippines, Negros
Occidental, Bario Alagria, Lupit River, on submerged
wood (Hyde et al. 1999a); South Africa, Durban, Palmier
River, on submerged wood (Hyde et al. 1999a); USA,
Wisconsin, Tomahawk River, on submerged decorticated
wood (Campbell et al. 2003b).
Asexual morph: Undetermined
Notes: Holotype NY-00966744, other specimen collected from freshwater ILL A333-1. LSU, SSU sequence
data is available.
Natantispora retorquens (Shearer & J.L. Crane) Campb.
et al.
Fungal Diversity (2019) 99:451–660
: Halosarpheia retorquens Shearer & J.L. Crane, Bot.
Mar. 23: 608 (1981)
Distribution: China, Yunnan Province, on submerged
wood (Cai et al. 2002a); Mauritius, near Tamarin, Black
River, on submerged wood (Hyde et al. 1999a); Philippines, Luzon, Laguna, Los Bados, Mt. Maquiling, on wood
submerged in a freshwater stream (Hyde et al. 1999a);
South Africa, Kwa Zulu-Natal, Mt. Ubatuba, on submerged rachis of Raphia australia (Hyde et al. 1999a);
USA, Minnesota, west arm of Lake Itasca, Lake Itasca
State Park, on submerged Typha sp. (Campbell et al.
2003b).
Asexual morph: Undetermined
Notes: Specimens collected from freshwater: ILL
A4–10, ILL A4–11. ITS, LSU and SSU sequence data are
available.
Panorbis Campb. et al., Mycologia 95(3): 544 (2003)
Asexual morph Undetermined. Sexual morph Description
and illustrations see Campbell et al. (2003b).
Type species: Panorbis viscosus (I. Schmidt) Campb.
et al., Mycologia 95(3): 544 (2003)
Notes: Campbell et al. (2003b) established the genus
Panorbis to accommodate Halosarpheia viscosa. Only one
species was accepted in this genus and occurrence on
woody and herbaceous plant debris in freshwater, brackish
and marine habitats (Schmidt 1985; Campbell et al.
2003b).
Panorbis viscosus (I. Schmidt) Campb. et al.
: Halosphaeria viscosa I. Schmidt, Mycotaxon 24: 420
(1985)
Distribution: Philippines, Luzon, Laguna, Los Bados,
Maquiling Mountain, on wood submerged in a freshwater
stream (Hyde et al. 1999a); South Africa, Durban, Palmiet
River, on submerged wood (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: Holotype No. 148, JE. LSU and SSU sequence
data are available. This species was introduced by Schmidt
(1985) as Halosphaeria viscosa I. Schmidt, and was collected from rotten wood submerged in seawater, Germany.
Hyde et al. (1999b) collected Halosphaeria viscosa from
freshwater habitats in Philippines and South Africa.
Campbell et al. (2003b) established the genus Panorbis and
synonymised Halosphaeria viscosa as Panorbis viscosus
based on morphology and phylogeny.
Phaeonectriella R.A. Eaton & E.B.G. Jones, Nova Hedwigia 19(3–4): 779 (1971) [1970]
Asexual morph Undetermined. Sexual morph Ascomata
immersed or partially immersed, globose to subglobose,
hyaline to pale brown, membraneous, ostiolate, papillate,
periphysate. Peridium composed of textura angularis or
compressed cells. Catenophyses absent. Asci 8-spored,
clavate, pedicellate, thin-walled, apically truncate and
555
thickened, with apical pore and cytoplasm retracted below
the ascus apex, peesistent. Ascospores ellipsoidal to fusiform, bicelled, hyaline or pale brown, thin-walled, with
appendages at each end.
Type species: Phaeonectriella lignicola R.A. Eaton &
E.B.G. Jones, Nova Hedwigia 19(3–4): 779 (1971) [1970]
Notes: Phaeonectriella was described from test blocks
exposed in a number of water cooling towers by Eaton and
Jones (1970). Hyde et al. (1999b) provided descriptions
and illustrations for the type species Phaeonectriella lignicola and introduced the second species Phaeonectriella
appendiculata in this genus based on the collections from
freshwater habitats of Australia, Mauritius and Philippines.
Phaeonectriella appendiculata Hyde et al.
Distribution: Australia, north Queensland, Crystal
Cascades, on submerged wood (Hyde et al. 1999a);
Philippines, Luzon, Laguna, Los Bados, Mt. Maquiling, on
wood submerged in a freshwater stream (Hyde et al.
1999a).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2864 (now in IFRD).
sequence data is not available.
Phaeonectriella lignicola R.A. Eaton & E.B.G. Jones
Distribution: China, Yunnan Province, Kunming city,
on submerged wood in Dianchi lake (Luo et al. 2004);
Mauritius, near Tamarin, Black River, on submerged
wood (Hyde et al. 1999a).
Asexual morph: Undetermined
Notes: ITS, LSU and SSU sequence data are available.
Triadelphiaceae Y.Z. Lu, J.K. Liu, Z.L. Luo & K.D.
Hyde, fam. nov.
Index Fungorum number: IF 555668, Facesoffungi
number: FoF 05449
Etymology:‘Triadelphiaceae’ referring to the type
genus Triadelphia.
Saprobic on decaying wood, or pathogenic on human, or
found from the gut of red palm weevils. Sexual morph
Undetermined. Asexual morph Hyphomycetous. Conidiophores lacking. Conidiogenous cells holoblastic,
monoblastic, integrated, flask-shaped, fusiform, cylindrical
or clavate, arising from undifferentiated hyphae, hyaline to
pale brown, smooth-walled. Conidia acrogenous, solitary,
develop one to five forms in distinct species: form
(a) cylindrical, oblong, or clavate, straight or slightly
curved, brownish, 1–2 septate, septa covered with broad
dark bands; form (b) clavate, club-shaped, dark brown,
uniseptate, septum covered with a dark band; form (c) obclavate to acicular with a narrow long tip, hyaline or yellowish brown, multiseptate; form (d) fusiform, obclavate
with acicular tips, or rounded at the tip, multi-septate, end
cells pale brown, central cells brown to dark brown, dark
band covering over the central septa; form (e) allantoid or
123
556
Fungal Diversity (2019) 99:451–660
Fig. 40 Conioscypha aquatica (MFLU 18–1640, holotype) a Colony on natural substrate. b–g Conidia. h Germinating conidium. Culture on
PDA from above (i) and reverse (j). Scale bars: b–h 20 lm
reniform, hyaline or pale yellowish, 0–3-septate; form
(f) obovate to broadly ellipsoidal, pale brown, unicellular.
(Shearer and Crane 1971; Constantinescu and Samson
1982; Matsushima 1989, 1995; Tzean and Chen 1989;
Abdel-Sater and Soliman 2017; Li and Ye 2017; Lu et al.
2018b).
Type genus: Triadelphia Shearer & J.L. Crane,
Mycologia 63(2): 247 (1971)
Note: The genus Triadelphia was established by Shearer
and Crane (1971) based on the type species T. heterospora,
which is placed in the order Microascales as genera incertae sedis (Wijayawardene et al. 2017, 2018; Lu et al.
2018b). In this study, our multi-gene phylogenetic analyses
show that the Triadelphia clade shares a sister relationship
to the Graphiaceae clade with good bootstrap support (98%
ML) (Fig. 1, clade 41). However, morphologically, Triadelphia taxa are obviously distinct from the species of
Graphiaceae in conidial and conidiophore characters. Most
Triadelphia species are pleomorphic, produce 2–5 forms of
conidia, and lack or with a short conidiophores (Shearer
and Crane 1971; Constantinescu and Samson 1982; Matsushima 1989, 1995; Tzean and Chen 1989; Abdel-Sater
and Soliman 2017; Li and Ye 2017; Lu et al. 2018b), while
the species of Graphiaceae have synnematous
123
conidiophores which are erect and 103–300 lm long, with
a cylindrical stipe and terminated in a slimy head of
conidia (Gilgado et al. 2005; Jacobs et al. 2013). Therefore,
Triadelphiaceae fam. nov. is introduced here.
Triadelphia Shearer & J.L. Crane, Mycologia 63(2): 247
(1971)
Asexual morph Colonies on natural substrate sparse,
brown to dark brown. Mycelium immersed or superficial,
composing of colorless to pale brown, smooth, septate
hyphae. Conidiogenous cells gregarious to caespitose,
globose to subglobose to ampulliform, borne directly on
vegetative hyphae. Conidia aerogenous, solitary, dry, septate, pleomorphic. Sexual morph Undetermined.
Type species: Triadelphia heterospora Shearer & J.L.
Crane, Mycologia 63(2): 247 (1971)
Notes: The pleomorphic genus Triadelphia was introduced by Shearer and Crane (1971) with T. heterospora as
type species. The sporodochial hyphomycete was isolated
from wood blocks submerged in the Patuxent River, USA
and characterized by two different forms of conidia. Li and
Ye (2017) accepted 18 species in Triadelphia and provided
key to species of this genus. Three Triadelphia species
Fungal Diversity (2019) 99:451–660
557
Fig. 41 Conioscypha submersa (MFLU 18–1639, holotype) a–h Conidia. Culture on PDA from surface (i) and reverse (j). Scale bars: a,
f 20 lm, b–e, g, h 10 lm
have been reported from freshwater habitats (Shearer and
Crane 1971; Révay 1992; Lu et al. 2018b).
Triadelphia fusiformis Lu et al.
Distribution: Thailand, Krabi, on submerged wood in a
freshwater stream (Lu et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–1436. ITS, LSU and TEF1a
sequence data are available.
Triadelphia heterospora Shearer & J.L. Crane
Distribution: USA, Maryland, on wood blocks submerged in the Patuxent River (Shearer and Crane 1971).
Sexual morph: Undetermined
Notes: Holotype ILLS 34793, isotype DAOM 126798,
IMI 14403. ITS and LSU sequence data are available.
Triadelphia morgoensis Révay
Distribution: Hungary, on submerged wood (Révay
1992).
Sexual morph: Undetermined
Notes: Holotype BP 664Fn. Sequence data is
unavailable.
Torpedosporales Jones et al.
Juncigenaceae Jones et al.
Juncigena Kohlm. et al., Bot. Mar. 40(4): 291 (1997)
Asexual morph Mycelium composed of septate, branched,
hyaline hyphae. Conidiophores obsolete or micronematous,
cylindrical to clavate, acrogenous or laterally on the
hyphae, hyaline to lightbrown. Conidiogenous cells
monoblastic, integrated, terminal, determinate. Conidia
acrogenous, irregularly helicoid, solitary, septate, constricted at the septa, yellow-brown to brown. The count of
the conidial cells is less than ten. Sexual morph Ascomata
immersed, subglobose to pyriform, ostiolate, coriaceous,
fuscous, single. Neck cylindrical, hyaline to fuscous, periphysate. Peridium of ellipsoidal to subglobose cells,
forming a textura angularis, dark on the outside, hyaline on
the inside. Paraphyses composed of unbranched pseudoparaphyses attached at the top and bottom. Asci
8-spored, unitunicate, fusiform to cylindrical, short
pedunculate, with a non-amyloid apical ring. Ascospores
uni- to biseriate, fusiform to ellipsoidal, 3-septate, hyaline.
Type species: Juncigena adarca Kohlm. et al., Bot. Mar.
40(4): 291 (1997)
123
558
Notes: The monotypic genus Juncigena was introduced
by Kohlmeyer et al. (1997) and typified by J. adarca for an
asexual morph which was described as Cirrenalia adarca.
Later the asexual species was placed in the genus Moheitospora as M. adarca along with the type species, M.
fruticosae (Abdel-Wahab et al. 2010), thus Juncigena and
Moheitospora are synonyms. In case of Juncigena has
priority and has greater use in the literature, Réblová et al.
(2016b) recommended that Juncigena be used.
Juncigena fruticosae (Abdel-Wahab et al.) A.N. Mill. &
Shearer
: Moheitospora fruticosae Abdel-Wahab et al., Mycol.
Progr. 9(4): 551 (2010)
Distribution: Egypt, Alexandria, Lake Manzala, on
submerged decayed stems of Suaeda fruticosa (AbdelWahab et al. 2010).
Sexual morph: Undetermined
Notes: Holotype IMI 391650. LSU and SSU sequence
data are available. This species is only known from the
type locality.
Hypocreomycetidae genera incertae sedis
Papulaspora Preuss, Linnaea 24: 112 (1851)
Asexual morph Mycelium branched, septate, hyaline or
fuscous, more or less procumbent. Bulbils initially hyaline,
becoming light brown to black, and opaque at maturity as a
result of progressive darkening of cell walls, subglobose to
irregular, variable in size, intercalary or less frequently
terminal, developing on pedicels. Primordium a group of
terminal or intercalary, terminating the pedicel, consists of
one to three branches forming septate side branches by
dichotomous division, the final divisions widen to spherical
cells, which constitute the bulbil. Conidia on bottle-shaped
sterigmata. Sexual morph Undetermined.
Type species: Papulaspora sepedonioides Preuss, Linnaea 24: 112 (1851)
Notes: Papulaspora was introduced by Preuss (1851)
with P. sepedonioides Preuss as the type species which
produced bulbils. Species of Papulaspora are known to be
saprophytic (Hotson 1917, 1942; Fergus 1971). Papulaspora coprophila was reported as coprophilous fungus
(Hotson 1912, 1917). Several are endophytes (Gezgin and
Eltem 2009; Vitorino et al. 2012) and soil fungi (Watanabe
1991; He et al. 1997). Papulaspora equi can infect mammals (Shadomy and Dixon 1989). Among the Papulaspora
species, P. pulmonaria and P. sepedonioides were reported
from freshwater habitats (van Beverwijk 1954; Cai et al.
2006a).
Papulaspora pulmonaria Beverw
Distribution: UK, Netherlands, on decaying leaves in
water (van Beverwijk 1954)
Sexual morph: Undetermined.
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Fungal Diversity (2019) 99:451–660
Notes: Sequence data is not available. van Beverwijk
(1954) mentioned that the type of this species was found
from submerged leaves in an old tin filled with rain water
in a garden, but the author did not give herbarium number
for the type.
Papulaspora sepedonioides Preuss
Distribution: China, Yunnan Province, on submerged
bamboo culms (Cai et al. 2006a)
Sexual morph: Undetermined.
Notes: ITS, LSU, RPB1, RPB2, TEF1a and b-tubulin
sequence data are available.
Subclass Lulworthiomycetidae Dayar. et al.
Pisorisporiales Réblová & J. Fourn.
Pisorisporiaceae Réblová & J. Fourn.
Achroceratosphaeria Réblová et al., Fungal Divers 43: 79
(2010)
Asexual morph Undetermined. Sexual morph Ascomata
immersed to semi-immersed, nonstromatic, solitary or in
small groups, venter globose to subglobose, sometimes
depressed, subhyaline to pale brown. Neck cylindrical,
hyaline to subhyaline to grey, sometimes hairy, periphysate. Peridium leathery to fragile, two-layered. Paraphyses septate, hyaline, tapering towards the tip, longer
than the asci. Asci 8-spored, unitunicate, cylindrical to
clavate, stipitate. Ascospores cylindrical-fusiform to
cylindrical, septate, hyaline.
Type species: Achroceratosphaeria potamia Réblová
et al., Fungal Divers 43: 79 (2010)
Notes: The genus Achroceratosphaeria was described
for perithecial ascomycetes with similar morphology to
Ceratosphaeria and Pseudohalonectria of the Magnaporthaceae (Réblová et al. 2010). Presently, Achroceratosphaeria comprises two freshwater and one terrestrial
species characterised by immersed, subhyaline to pale
brown ascomata with a fragile, hyaline to pale brown
protruding neck, tapering paraphyses, unitunicate asci with
a nonamyloid apical annulus and ellipsoidal to fusiform,
septate ascospores (Réblová et al. 2010). For one of the
freshwater Achroceratosphaeria species, collected from
Hong Kong, China, was described as Achroceratosphaeria
sp. by Réblová et al. (2010). Although the sequence of
Achroceratosphaeria sp. was deposited in GenBank (under
the name Ascocollumdensa aquatica), this fungus has
never been validly published. The only known written
records are the photographs and descriptions made by
Ranghoo (1998) in her Ph. D thesis. New material needs to
be recollected.
Achroceratosphaeria potamia Réblová, J. Fourn. & K.D.
Hyde
Distribution: France, Ariège, Le Baup stream, on submerged wood of Platanus species (Réblová et al. 2010).
Asexual morph: Undetermined
Fungal Diversity (2019) 99:451–660
Notes: Holotype PRM 915719. LSU and SSU sequence
data are available. This species is known so far only from
the type locality.
Pisorisporium Réblová & J. Fourn., Persoonia 34: 45
(2015)
Asexual morph Undetermined. Sexual morph Description
and illustration see Réblová et al. (2015b).
Type species: Pisorisporium cymbiforme Réblová & J.
Fourn., Persoonia 34: 45 (2015)
Notes: The genus Pisorisporium was introduced by
Réblová et al. (2015b) with Pisorisporium cymbiforme
Réblová & J. Fourn as type species which was collected
from freshwater habitats. Presently, Pisorisporium comprises two species, one is from freshwater and another one
from terrestrial habitats.
Pisorisporium cymbiforme Réblová & J. Fourn
Distribution: France, Ariège, valley of La Maille brook,
on submerged decorticated wood of Alnus glutinosa and
Fraxinus excelsior (Réblová et al. 2015b).
Asexual morph: Undetermined
Notes: Holotype PRM 924377, other specimens collected from freshwater habitats: PRM 924378, PRM
924379. LSU, SSU and RPB2 sequence data are available.
Subclass Savoryellomycetidae Hongsanan et al.
Conioscyphales Réblová & Seifert
Conioscyphaceae Réblová & Seifert
Conioscypha Höhn., Annls mycol. 2(1): 58 (1904)
Asexual morph Conidiophores micronematous, mononematous, errect, hyaline, smooth walled. Conidiogenous
cells enteroblastic, terminal or intercalary, hyaline,
cyathiform to doliiform, with multilayered, cup-like collarettes. Conidia often with a basal pore, formed singly and
successively by percurrent regeneration of the apex of the
conidiogenous cell, liberating by apical rupture of the outer
wall of the conidiogenous cell, aseptate, brown,. Sexual
morph Ascomata perithecial, immersed to partially
immersed, or superficial, globose to subglobose, with
subcylindrical papilla or cylindrical elongated neck, ostiole
periphysate. Peridium leathery, waxy, comprising two
layers. Paraphyses filiform, unbranched, longer than the
asci. Asci 8-spored, unitunicate, persistent, with a pronounced non-amyloid apical annulus, cylindrical-clavate,
stipitate. Ascospores fusiform to fusiform-navicular,
transversely multiseptate, hyaline, without mucilaginous
sheath or appendages.
Type species: Conioscypha lignicola Höhn., Annls
mycol. 2(1): 58 (1904)
Notes: The dematiaceous hyphomycetous genus Conioscypha, with C. lignicola as type species, includes
species from freshwater and terrestrial habitats. The genus
Conioscyphascus, typified by C. varius, was originally
established for fungi with Conioscypha asexual morphs
559
(Réblová and Seifert 2004b). Based on priority and the
greater number of species, Réblová et al. (2016b) recommended to use the name Conioscypha.
Conioscypha aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555669, Facesoffungi
number: FoF 05450, Fig. 40
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–1640
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, dark brown to black. Conidiophores
reduced to conidiogenous cells. Conidia 19–23 lm long,
17–21 lm wide (
x = 21 9 19 lm, n = 20), dry, dark
brown to black, mostly globose to subglobose, smoothwalled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on submerged decaying wood, May 2015,
Z.L. Luo, S-447 (MFLU 18–1640, holotype; HKAS 92696,
isotype), ex-type living culture MFLUCC 18–1333,
KUMCC 15–0294.
Notes: Conioscypha aquatica resembles C. hoehnelii in
having globose to subglobose, dark brown conidia (Kirk
1984). However, Conioscypha aquatica differs from C.
hoehnelii in having larger conidia (19–23 9 17–21 vs.
12–17 9 11–15 lm). Phylogenetic analysis also shows
that Conioscypha aquatica is distinct from C. hoehnelii
(Fig. 1, clade 46).
Conioscypha gracilis (Munk) Zelski et al.
: Debaryella gracilis Munk, Bot. Tidsskr. 51: 226
(1954)
: Cryptoleptosphaeria gracilis (Munk) Rossman &
Samuels, Stud. Mycol. 42: 185 (1999)
: Conioscyphascus gracilis (Munk) Réblová & Seifert,
Stud. Mycol. 50:104 (2004)
Distribution: Peru, Cusco, Camanti, stream along
Quincemil Trail, on submerged woody debris (Zelski et al.
2015).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater habitats:
ILL 41215, ILL 41203. LSU sequence data is available.
Conioscypha lignicola Höhn.
Distribution: Australia, on submerged wood (Höhnel
1904).
Sexual morph: Undetermined
Notes: Holotype IMI 270438. LSU, SSU and RPB2
sequence data are available.
Conioscypha nakagirii Chuaseehar. et al.
Distribution: Thailand, Nakhon Ratchasima Province,
Pak Chong District, on unidentified submerged wood in a
stream (Chuaseeharonnachai et al. 2017).
Asexual morph: see Chuaseeharonnachai et al. (2017)
123
560
Notes: Holotype BBH 40587, paratype BBH 40588.
ITS, LSU, SSU and RPB2 sequence data are available.
Conioscypha peruviana Zelski et al.
Distribution: Peru, Cusco, Camanti, stream along
Quincemil Trail, on submerged woody debris (Zelski et al.
2015).
Asexual morph: see Zelski et al. (2015)
Notes: Holotype ILL 41202, other specimens collected
from freshwater habitats: ILL 41208, ILL 41209, ILL
41210, ILL 41211, ILL 41213, ILL 41214. LSU sequence
data is available.
Conioscypha submersa Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555670, Facesoffungi
number: FoF 05451, Fig. 41
Etymology: Referring to the submerse habitat of this
fungus.
Holotype: MFLU 18–1639
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, dark brown to black. Conidiophores
reduced to conidiogenous cells. Conidia 17–19 lm long,
15–17 lm wide (
x = 18 9 16 lm, n = 20), globose to
subglobose, or ovoid, dry, guttulate, pale brown when
young, dark brown to black when mature, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, S.M. Tang, S-904 (MFLU 18–1639, holotype), extype living culture DLUCC 0904.
Notes: Conioscypha submersa resembles C. lignicola in
having micronematous conidiophores, globose to subglobose, or ovoid, dark brown to black conidia with similar
size (Matsushima 1975). However, phylogenetic analysis
shows that Conioscypha submersa is distinct from C. lignicola (Fig. 1, clade 45).
Fuscosporellales Yang et al.
Fuscosporellaceae Yang et al.
Bactrodesmiastrum Hol.-Jech., Folia geobot. phytotax.
19(1): 103 (1984)
Asexual morph Emended description and illustration refer
to Hernández-Restrepo et al. (2015a). Sexual morph
Undetermined.
Type species: Bactrodesmiastrum obscurum Hol.-Jech.,
Folia geobot. phytotax. 19(1): 105 (1984)
Notes: The genus Bactrodesmiastrum is characterized
by solitary or aggregated conidiophores, mostly reduced to
brown, monoblastic, conidiogenous cells, arising from
pulvinate to subpustulate sporodochial conidiomata, and
moniliform or beaded hyphoid cells (Holubová-Jechová
1984; Hernández-Restrepo et al. 2015a). Five species are
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Fungal Diversity (2019) 99:451–660
accepted in this genus and one of them was collected from
freshwater habitats (Calduch et al. 2002).
Bactrodesmiastrum obovatum (Calduch et al.) Mena et al.
: Janetia obovata Calduch et al., Mycologia 94(2): 355
(2002)
Distribution: Spain, Balearic Islands, Malorca, Tramuntana Mountain, on unidentified submerged wood in
Reservoir (Calduch et al. 2002).
Sexual morph: Undetermined
Notes: Holotype IMI 380443, Isotype FMR 6482, other
specimen collected from freshwater habitats: FMR 7274.
LSU sequence data is available.
Fuscosporella Yang et al., Cryptog. Mycol. 37(4): 457
(2016)
Asexual morph Descriptions and illustrations see Yang
et al. (2016). Sexual morph Undetermined.
Type species: Fuscosporella pyriformis Yang et al.,
Cryptog. Mycol. 37(4): 458 (2016)
Notes: Fuscosporella is the type genus of the family
Fuscosporellaceae with F. pyriformis as the type species
(Yang et al. 2016a). Currently, two species are included in
the genus and both are reported from freshwater streams in
Thailand. Fuscosporella morphologically resembles Bactrodesmisatrum and Bactrodesmium which they share the
characters as having sporodochial conidiomata,
monoblastic conidiogenous cells and brown septate conidia
(Hernández-Restrepo et al. 2013, 2015a). However, species
in Fuscosporella have hyphoid, hyaline conidiogenous
cells, while Bactrodesmiastrum species are characterized
by solitary or aggregated conidiophores reduced to brown,
single or moniliform conidiogenous cells (Holubová-Jechová 1984; Hernández-Restrepo et al. 2015a). Bactrodesmium differs from Fuscosporella in having hyaline
or brown, simple or branched conidiophores supporting
mono- or polyblastic conidiogenous cells (Ellis 1971;
Holubová-Jechová 1972). Phylogenetic studies indicate
that Bactrodesmium lies within the Dothideomycetes
(Hernández-Restrepo et al. 2015a) while Fuscosporella
belongs to Fuscosporellales, Sordariomycetes (Yang et al.
2016a, 2017).
Fuscosporella aquatica J. Yang & K.D. Hyde
Distribution: Thailand, Phang Nga Province, Bann Tom
Thong Khang, on decaying wood submerged in a freshwater stream (Yang et al. 2017).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–1973. ITS, LSU and SSU
sequence data are available.
Fuscosporella pyriformis Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on submerged wood in freshwater stream (Yang et al.
2016a).
Sexual morph: Undetermined
Fungal Diversity (2019) 99:451–660
561
Fig. 42 Phaeoisaria filiformis (MFLU 18–1462, holotype). a, b
Ascomata necks on decaying wood. c Ascomata d Section of
ascomata. e Section of peridium. f Paraphyses. g–k Asci. l–
n Ascospore. o germinating ascospore. Cultures on PDA from above
(p) and reverse (q). Scale bars: d 100 lm, e 20 lm, f–o 30 lm
Notes: Holotype MFLU 16–1979, isotype HKAS 95050.
ITS, LSU, SSU and RPB2 sequence data are available.
Asexual morph Descriptions and illustrations see Yang
et al. (2016a). Sexual morph Undetermined.
Mucispora Yang et al., Cryptog. Mycol. 37(4): 466 (2016)
123
562
Type species: Mucispora obscuriseptata J. Yang, Bhat
& K.D. Hyde, Cryptog. Mycol. 37(4): 466 (2016)
Notes: In the family Fuscosporellaceae, Mucispora is
different from the asexual genera Bactrodesmisatrum,
Fuscosporella and Parafuscosporella by its distinctive
longer brown conidiophores. Mucispora is morphologically
similar to Acrogenospora and Monotosporella in producing
unbranched, percurrent conidiophores and brown conidia
(Cai et al. 2006b). However, conidia in Mucispora and
Monotosporella are septate while those in Acrogenospora
are unicellular. Presently, two species are included in
Mucispora and both were collected from freshwater habitats in Thailand (Yang et al. 2016a, 2017).
Mucispora obscuriseptata Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, stream flowing outside Kaeng Krachan National
Park, on decaying submerged wood (Yang et al. 2016a).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1153, isotype HKAS 95047.
ITS, LSU and SSU sequence data are available.
Mucispora phangngaensis J. Yang & K.D. Hyde
Distribution: Thailand, Phang Nga Province, Bann Tom
Thong Khang, on decaying wood submerged in a freshwater stream (Yang et al. 2017).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–1974. ITS, LSU and SSU
sequence data are available.
Parafuscosporella J. Yang & K.D. Hyde, Cryptog. Mycol.
37(4): 458 (2016)
Asexual morph Descriptions and illustrations see Yang
et al. (2016a). Sexual morph Undetermined.
Type species: Parafuscosporella moniliformis J. Yang,
Bhat & K.D. Hyde, Cryptog. Mycol. 37(4): 460 (2016)
Notes: The generic delimitation of Parafuscosporella
and Fuscosporella is narrow. However, the two genera are
phylogenetically distinct. Based on the morphology of
sporulating on media, Fuscosporella is distinct from
Parafuscosporella by multi-celled, filamentous to helicoid
conidia, while Parafuscosporella produces globose to
obpyriform, uniseptate conidia in culture. Three species are
included in Parafuscosporella and all were reported from
freshwater habitats in Thailand.
Parafuscosporella garethii Boonyuen et al.
Distribution: Thailand, Chiang Mai Province, Chiang
Dao District, a small stream at Ban Hua Thung community
forest, on unidentified decaying submerged wood (Boonyuen et al. 2016).
Sexual morph: Undetermined
Notes: Holotype BBH 40839, paratype BBH 40840.
LSU, SSU and RPB2 sequence data are available.
Parafuscosporella moniliformis Yang et al.
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Fungal Diversity (2019) 99:451–660
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Yang
et al. 2016a).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1161, isotype HKAS 95049.
ITS, LSU and SSU sequence data are available.
Parafuscosporella mucosa Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, stream flowing outside Kaeng Krachan National
Park, on decaying submerged wood (Yang et al. 2016a).
Sexual morph: Undetermined
Notes: Holotype MFLU 16–1980, isotype HKAS 95051.
ITS, LSU and SSU sequence data are available.
Plagiascoma Réblová & J. Fourn, Persoonia 37: 69 (2015)
[2016]
Asexual morph Undetermined. Sexual morph Ascomata
perithecial, immersed, gradually erumpent to semi-immersed, non-stromatic, conical, dark brown, lying obliquely to horizontally, papillate or with a neck. Ostiole
periphysate. Peridium fragile, 2-layered. Paraphyses
abundant, persistent, septate. Asci 8-spored, unitunicate,
cylindrical to cylindricalfusiform, stipitate, apex with a
non-amyloid apical annulus. Ascospores fusiform, transversely septate, hyaline.
Type species: Plagiascoma frondosum Réblová & J.
Fourn., Persoonia 37: 69 (2015) [2016]
Notes: Plagiascoma is a monotypic genus placed at the
basal branch in Fuscosporellales (Réblová et al. 2016a;
Yang et al. 2016a, 2017; Hernández-Restrepo et al. 2017).
Plagiascoma frondosum resembles species in Annulatascus, e.g. A. nilensis and A. tropicalis, in having cylindrical
asci with obvious non-amyloid apical annulus and fusiform, hyaline, multi-septate ascospores. Most Annulatascus
species have mucilaginous sheaths while no sheath or
appendages were observed in P. frondosum. Presently, only
one species is included in this genus which was collected
from freshwater habitats and only known from the type
locality.
Plagiascoma frondosum Réblová & J. Fourn.
Distribution: France, Midi-Pyrénées, Ariège, Rimont,
valley of La Maille brook, submerged decorticated wood of
Fraxinus excelsior (Réblová et al. 2016a)
Asexual morph: Undetermined
Notes: Holotype PRM 933854. LSU, SSU, RPB2 and
MCM7 sequence data are available.
Pseudoascotaiwania Yang et al., Cryptog Mycol 37(4):
471 (2016)
Asexual morph Undetermined. Sexual morph Ascomata
scattered, immersed, urniform, rostrate, membranous,
black, paraphysate, ostiolate. Ostiole periphysate. Peridium
several layers thick, brown, with textura angularis. Paraphyses simple or branched. Asci unitunicate, 8-spored,
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563
Fig. 43 a Pleurothecium recurvatum (DLU 0611) a Colonies on substrate. b, c Conidiophore with conidia. d–e conidiogenous cells with
conidia. g–h Conidium. Scale bars: b, c 100 lm, d–f 25 lm, g–h 15 lm
cylindrical, persistent, pedicellate, with a non-amyloid
apical ring. Ascospores uniseriate, cylindrical to fusiform,
5–6-septate, sometimes slightly constricted at the septa,
with central cells light brown, end cells hyaline.
Type species: Pseudoascotaiwania persoonii (Fallah
et al.) Yang et al., Cryptog. Mycol. 37(4): 471 (2016)
Notes: Ascotaiwania persoonii is phylogenetically distinct from Ascotaiwania species in Savoryellales. Yang
et al. (2016a) introduced a new genus Pseudoascotaiwania
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564
to accommodate Ascotaiwania persoonii and assigned this
genus in Fuscosporellales.
Pseudoascotaiwania persoonii (Fallah et al.) Yang et al.
: Ascotaiwania persoonii Fallah et al., Can. J. Bot.
77(1): 87 (1999)
Distribution: Canada, Manitoba, Grass River, on
decorticated submerged wood (Fallah et al. 1999).
Asexual morph: Undetermined
Notes: Holotype ILLS 52298. LSU sequence data is
available.
Pleurotheciales Réblová & Seifert
Pleurotheciaceae Réblová & Seifert
Monotosporella S. Hughes, Can. J. Bot. 36: 786 (1958)
Asexual morph Colonies on natural substrate effuse.
Mycelium immersed, consisting of brown, septate, thinwalled, branched hyphae. Conidiophores macronematous,
mononematous, solitary, errect, straight, unbranched,
cylindrical, septate, smooth, thick-walled. Conidiogenous
cells holoblastic, monoblastic, integrated, terminal, cylindrical. Conidia acrogenous, obovoid to pyriform, or clavate, septate, brown to black, paler towards the base,
smooth-walled. Conidial sessesion schizolytic. Sexual
morph Undetermined.
Type species: Monotosporella setosa (Berk. & M.A.
Curtis) S. Hughes, Can. J. Bot. 36: 787 (1958)
Notes: Monotosporella was introduced by Hughes
(1958) with M. setosa as the type species. Monotosporella
is characterized by unbranched, percurrently proliferating
conidiophores, septate, pyriform to obvoid, dark brown to
black conidia. There are 12 species in this genus and two of
them were reported from freshwater habitats (Ranghoo
et al. 1999; Tsui et al. 2001e; Cai et al. 2003b).
Monotosporella microaquatica (Tubaki) Sv. Nilsson
: Dactylella microaquatica Tubaki, Bull. natn. Sci.
Mus., Tokyo, N.S. 3: 256 (1957)
Distribution: Philippines, on submerged wood in the
Liput River (Cai et al. 2003b)
Sexual morph: Undetermined
Notes: Holotype is not indicated. Sequence data is not
available.
Monotosporella rhizoidea V. Rao & de Hoog
Distribution: China, Hong Kong, on submerged wood
(Ranghoo et al. 1999; Tsui et al. 2001e).
Sexual morph: Undetermined
Notes: Sequence data is not available.
Phaeoisaria Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 330 [56 repr.] (1909)
Asexual morph Conidiomata synnematal, indeterminate,
scattered, erect, rigid, dark brown to black, composed of
compactly and parallelly adpressed conidiophores. Conidiophores macronematous, synnematous, brown to dark
brown, smooth. Synnemata erect, rigid, dark brown to
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black, velvety, smooth, composed of compactly and parallelly adpressed conidiophores, with flared conidiogenous
cells in the above half. Conidiogenous cells polyblastic,
terminal and intercalary, mostly discrete, sometimes integrated, recurved, smooth, sympodial extended, denticulate,
subhyaline to pale brown, sympodial, each with one to
several denticulate conidiogenous loci. Conidia ellipsoidal
to obovoidal or cylindric-ovate, straight, guttulate, smoothwalled. Sexual morph Ascomata superficial to immersed,
globose to elongate globose, with a long neck. Paraphyses
numerous, filamentous, branched, septate, smooth. Asci
8-spored, unitunicate, cylindrical, thin-walled, with a small
refractive apical apparatus, smooth-walled. Ascospores
filiform, guttulate, tapered at both ends, septate, hyaline,
smooth-walled.
Type species: Phaeoisaria bambusae Höhn., Sber.
Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 329
(1909)
Notes: Phaeoisaria was introduced by Höhnel (1909)
for a collection on Gigantochloa sp. (Bambusae) and is
typified by P. bambusae Höhn. Three species in Phaeoisaria have been reported from freshwater habitats and all of
them were collected from southwestern China (Hyde et al.
2018; Luo et al. 2018b).
Phaeoisaria aquatica Luo et al.
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0918. ITS, LSU and RPB2
sequence data are available. This species is only known
from the type locality.
Phaeoisaria clematidis (Fuckel) S. Hughes
: Stysanus clematidis Fuckel, Jb. nassau. Ver. Naturk.
23-24: 365 (1870)
Distribution: China, Yunnan Province, Lancang River,
saprobic on submerged decaying wood (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater habitats:
MFLU 17–0919, MFLU 17–0920. ITS, LSU, SSU and
RPB2 sequence data are available.
Phaeoisaria guttulata J. Yang & K.D. Hyde
Distribution: China, Guizhou Province, Anshun city,
Gaodang village, on decaying wood submerged in Suoluo
River (Hyde et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0139; ITS, LSU, SSU
sequence data are available.
Phaeoisaria filiformis D.F. Bao, Z.L. Luo, K.D. Hyde &
H.Y. Su, sp. nov.
Index Fungorum number: IF 555671, Facesoffungi
number: FoF 05452, Fig. 42
Etymology: Referring to the filiform ascospores of this
fungus.
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565
Fig. 44 Sterigmatobotrys uniseptata (HKAS 83988) a Conidiophore with conidia. b Conidiogenous cells with coidia. c–j Conidia.
k Germinating conidium. Scale bars: a 50 lm, b 25 lm, c 15 lm, d–k 10 lm
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566
Holotype: MFLU 18–1462
Saprobic on decaying wood submerged in freshwater.
Asexual morph Undetermined. Sexual morph Ascomata
350–450 lm diam., 400–450 lm high, immersed, globose
to elongate globose, with a long, cylindrical, black neck.
Peridium 29–31 lm thick, composed 2 layers, inner layer
composed of hyaline, thin-walled cells of textura angularis, outer layer comprised of small angular cells with
dark brown amorphous material. Paraphyses 7–11 lm
wide, numerous, filamentous, branched, septate, smooth.
Asci 120–136 9 12–14 lm (
x = 128 9 13 lm, n = 30), 8spored, unitunicate, cylindrical, thin-walled, with a small
refractive apical apparatus, smooth. Ascospores 88–
116 9 2.5–3.5 lm (
x = 102 9 3 lm, n = 30), filiform,
guttulate, tapering at both ends, multi-septate, hyaline,
smooth-walled.
Material examined: THAILAND, Sai khu Waterfall, on
submerged decaying wood, August 2017, K. Vinit, B-33
(MFLU 18–1462, holotype), ex-type living culture,
MFLUCC 18–0214.
Notes: Phaeoisaria filiformis resembles species of Ophioceras and Ceratosphaeria in having immersed ascomata with
a long, cylindrical, black neck, unitunicate, 8-spored, cylindrical asci and hyaline, guttulate, filiform ascospores (Niessl
1876; Saccardo 1883; Hu et al. 2012c). However, phylogenetic analysis shows that Phaeoisaria filiformis belongs to the
Pleurotheciales while Ophioceras and Ceratosphaeria belong
to Magnaporthales (Fig. 1, clade 19, 20, 43).
Pleurotheciella Réblová et al., Mycologia 104(6): 1304
(2012)
Asexual morph: Descriptions and illustrations see Réblová
et al. (2012), Luo et al. (2018b). Sexual morph: Descriptions and illustrations see Réblová et al. (2012).
Type species: Pleurotheciella rivularia Réblová et al.,
Mycologia 104(6): 1304 (2012)
Notes: Pleurotheciella was introduced by Réblová et al.
(2012) with two new species, P. rivularia and P. centenaria and both were collected from freshwater habitats.
Presently, eleven species are accepted in this genus.
Pleurotheciella is a representative freshwater fungal genus
as all species in this genus were reported from freshwater
and most of them were collected from Asia (Hyde et al.
2018; Luo et al. 2018b).
Pleurotheciella aquatica Luo et al.
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0911. ITS, LSU, SSU and
RPB2 sequence data are available.
Pleurotheciella centenaria Réblová et al.
Distribution: Canada, Ontario, Centennial Lake, on
decaying submerged wood (Réblová et al. 2012).
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Sexual morph: Undetermined
Notes: Holotype DAOM 229631. ITS, LSU, SSU and
RPB2 sequence data are available.
Pleurotheciella fusiformis Luo et al.
Distribution: China, Yunnan Province, Erhai Lake, on
decaying submerged wood (Luo et al. 2018b).
Asexual morph: see Luo et al. (2018b).
Notes: Holotype MFLU 17–0909, isotype HKAS 92582,
other specimens collected from freshwater MFLU
17–0907, MFLU 17–0908, MFLU 17–0910. ITS, LSU,
SSU and RPB2 sequence data are available.
Pleurotheciella guttulata Luo et al.
Distribution: China, Yunnan Province, Dulong River,
on decaying submerged wood (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0914, isotype HKAS 92699.
ITS, LSU, SSU and RPB2 sequence data are available.
Pleurotheciella krabiensis J. Yang & K.D. Hyde
Distribution: Thailand, Krabi Province, on decaying
wood submerged in a freshwater stream near Morakot lake
(Hyde et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0140. ITS, LSU and SSU
sequence data are available.
Pleurotheciella lunata Luo et al.
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0913; ITS, LSU, SSU and
RPB2 sequence data are available. This species is only
known from the type locality.
Pleurotheciella rivularia Réblová et al.
Distribution: France, on decaying submerged wood of
Hedera helix (Réblová et al. 2012).
Asexual morph: see Réblová et al. (2012)
Notes: Holotype PRM 899852, other specimen collected
from freshwater habitats: PRM 899853. ITS, LSU, SSU
and RPB2 sequence data are available.
Pleurotheciella saprophytica Luo et al.
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0915. ITS, LSU, SSU and
RPB2 sequence data are available. This species is only
known from the type locality.
Pleurotheciella submersa Z.L. Luo & K.D. Hyde
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0916. ITS, LSU, SSU and
RPB2 sequence data are available. This species is only
known from the type locality.
Pleurotheciella tropica J. Yang & K.D. Hyde
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Distribution: Thailand, Phang Nga Province, Bann Tom
Thong Khang, on decaying wood submerged in a freshwater stream (Hyde et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 18–0141; ITS, LSU and SSU
sequence data are available. This species is only known
from the type locality.
Pleurotheciella uniseptata (Matsush.) Seifert.
: Dactylaria uniseptata Matsush.
Distribution: China, Yunnan Province, Jinsha River,
saprobic on submerged decaying wood (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater habitats:
MFLU 17–0912. ITS, LSU and RPB2 sequence data are
available.
Pleurothecium Höhn., Ber. dt. bot. Ges. 37: 154 (1919)
Holomorph Descriptions and illustrations see Réblová
et al. (2012).
Type species: Pleurothecium recurvatum (Morgan)
Höhn., Centbl. Bakt. ParasitKde, Abt. II 60: 26 (1923)
[1924]
Notes: Pleurothecium was introduced by Höhnel (1919)
with P. recurvatum (Morgan) Höhn as type species. Ten
species are accepted in the genus Pleurothecium (Monteiro
et al. 2016a; Hyde et al. 2017; Luo et al. 2018b) and four of
them have been reported from freshwater habitats.
Pleurothecium aquaticum Luo et al.
Distribution: China, Yunnan Province, Dali, on decaying wood submerged in a stream (Luo et al. 2018b).
Sexual morph: Undetermined
Notes: Holotype MFLU 17–0922. ITS and LSU
sequence data are available. This species is only known
from the type locality.
Pleurothecium floriforme J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying wood submerged in a freshwater stream (Hyde
et al. 2017).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1163, isotype GZAAS
17–0013. ITS, LSU and SSU sequence data are available.
This species is only known from the type locality.
Pleurothecium pulneyense Subram. & Bhat
Distribution: China, Yunnan Province, Jinsha River,
saprobic on decaying wood submerged (Luo et al. 2018b);
India, Western Ghats, on decomposing woody litter in a
freshwater stream (Sridhar and Sudheep 2011).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater MFLU
17–0921. LSU, SSU and RPB2 sequence data are
available.
Pleurothecium recurvatum (Morgan) Höhn.
567
: Acrotheciumr ecurvatum Morgan. J. Cincinnati Soc
Nat Hist 18: 44 (1895)
Facesoffungi number: FoF 05453, Fig. 43
Saprobic on decaying wood submerged in freshwater.
Asexual morph Colonies effuse, superficial, hairy, brown
to dark brown. Mycelium immersed or superficial. Conidiophores
299–371 lm
long,
6–10 lm
wide
(
x = 335 9 8 lm, n = 20), macronematous, mononematous, solitary, straight or slightly flexuous, septate, smooth,
brown to dark brown, paler towards the apex. Conidiogenous cells holoblastic, polyblastic, integrated, terminal,
denticulate, denticles cylindrical, hyaline or grayish.
Conidia 25–31 lm long, 7–9 lm wide (
x = 28 9 8 lm,
n = 20), acrogenous, cylindrical or clavate, straight or
slightly curved, rounded at both ends, 3-septate, guttulate,
hyaline, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on decaying wood submerged in
a freshwater stream, July 2015, X.J. Su, S-611 (DLU
0611), living culture DLUCC 0611.
Distribution: China, Yunnan Province, Baoshan city, on
submerged wood in a stream in Gaoligong Mountain (This
study).
Sexual morph: see Fernández et al. (1999)
Notes: Specimen collected from freshwater habitats:
DLU 0611. ITS, LSU, SSU and RPB2 sequence data are
available. Morgan (1895) introduced Acrothecium recurvatum and later this species was synonymised as Pleurothecium recurvatum by Höhnel (1919). During the
investigation on lignicolous freshwater fungi in China, a
Pleurothecium-like fungus was collected from Yunnan
Province. Morphologically, our new isolate fits well with
Pleurothecium recurvatum in having macronematous,
mononematous, solitary, straight or slightly flexuous, septate, smooth, brown to dark brown conidiophores paler
towards the apex, integrated, terminal, polyblastic, denticulate, hyaline or grayish conidiogenous cells and cylindrical or clavate, straight or slightly curved, rounded at
both ends, 3-septate, guttulate, smooth conidia with similar
size (Höhnel 1919; Goos 1969). Réblová et al. (2012)
provided sequence data for Pleurothecium recurvatum and
our phylogenetic analysis shows that the new isolate
clusters with P. recurvatum with high support value
(Fig. 1, clade 43). Based on the morphology and phylogeny, we identify our isolate as Pleurothecium recurvatum and provide descriptions and illustration.
Pleurothecium semifecundum Réblová et al.
Distribution: France, Haute Garonne, Mancioux, Oustaous, le Rieutort Brook, on decaying submerged wood of
Sambucus nigra (Réblová et al. 2012).
Asexual morph: see Réblová et al. (2012)
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Notes: Holotype PRM1 899854, other specimen collected from freshwater habitats: PRM 899855. ITS, LSU
and SSU sequence data are available.
Sterigmatobotrys Oudem.
Holomorph Descriptions and illustrations refer to Réblová
and Seifert (2011).
Type species: Sterigmatobotrys elata (Sacc.) Oudem.,
Ned. kruidk. Archf, 2 sér. 4: 548 (1886)
Notes: Sterigmatobotrys was introduced by Oudemans
(1886) with two species S. elata and S. papyrogena. It is a
conspicuous, cosmopolitan dematiaceous hyphomycete
genus with species occurring on decaying wood in both
terrestrial (Sutton 1973; Hughes 1978; Thomas and Polwart
2003; Réblová and Seifert 2011) and freshwater (Eaton and
Jones 1971; Eaton 1972; Chang 1991; Hyde and Goh 1999;
Kane et al. 2002; Réblová and Seifert 2011). Three species
are accepted in this genus (Réblová and Seifert 2011; Ertz
et al. 2016).
Sterigmatobotrys macrocarpa (Corda) S. Hughes
: Graphium macrocarpum Corda, Icon. Fung. 3: 13
(1839) (holotype PRM 155517; epitype PRM 915682)
: Harpographium macrocarpum (Corda) Sacc., Syll.
Fung. 4: 620 (1886)
= Acrothecium bulbosum Sacc., Michelia 1: 74 (1877)
= Stachybotrys elata Sacc., Michelia 2: 560 (1882)
Distribution: Canada, Ontario, on submerged wood in a
stream (Réblová and Seifert 2011); UK, England, on submerged wood in River Coln (Hyde and Goh 1999).
Holomorph: see Réblová and Seifert (2011)
Notes: Epitype PRM 915682. LSU sequence data is
available.
Sterigmatobotrys uniseptata H.S. Chang
Facesoffungi number: FoF 05454, Fig. 44
Saprobic on decaying wood submerged in freshwater.
Asexual morph Colonies effuse, hairy, brown to dark
brown. Mycelium immersed. Conidiophores 90–138 lm
long, 7–9 lm wide (
x = 114 9 8 lm, n = 20), macronematous, mononematous, dark brown, septate, arising from
stromatic cells, each composed of a well-defined stipe and
a complex penicillate head consisting of series of penicillate branches and terminating with conidiogenous cells and
a head of slimy conidia. Conidiogenous cells 32–48 lm
long, 3–5 lm wide (
x = 40 9 4 lm, n = 20), polyblastic,
hyaline, parallel, smooth-walled. Conidia 15–19 lm long,
5–9 lm wide (
x = 17 9 7 lm, n = 20), solitary, slimy,
straight, cylindrical to subclavate, uniseptate, broad and
rounded at the apex and gradually narrowing towards the
base, thin-walled, hyaline, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a
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freshwater stream, March 2014, X.Y. Liu, S-030 (HKAS
83988), living culture MFLUCC 15–0358.
Distribution: China, Taiwan, on submerged wood
(Chang 1991), Yunnan Province (This study).
Notes: Holotype WLG 0612, other specimen collected
from freshwater HKAS 83988. ITS, LSU and SSU
sequence data are provide in this study. Sterigmatobotrys
uniseptata was introduced by Chang (1991). This species is
characterized by its macronematous, mononematous, dark
brown, septate conidiophores arising from stromatic cells,
each composed of a well-defined stipe and a complex
penicillate head consisting of series of penicillate branches
and terminating with conidiogenous cells and a head of
slimy conidia, polyblastic, hyaline conidiogenous cells and
solitary, slimy, thin-walled, hyaline, smooth, uniseptate,
straight, cylindrical to subclavate conidia (Chang 1991).
During the investigation of lignicolous freshwater fungi in
China, a Sterigmatobotrys-like fungus was collected from
Yunnan Province. Morphologically, our isolate fits well
with Sterigmatobotrys uniseptata (Chang 1991). Phylogenetic analysis shows that our isolate clusters together with
Sterigmatobotrys macrocarpa (DAOM230059, MR2973)
and Sterigmatobotrys rudis (DAOM 229838), but the
topology supports they are distinct species (Fig. 1, clade
43). Based on morphology and phylogeny, we identify our
isolate as Sterigmatobotrys uniseptata and provide the
descriptions and illustrations for this species as well as
sequence data.
Savoryellales Boonyuen et al.
Savoryellaceae Jaklitsch & Réblová
Ascotaiwania Sivan. & H.S. Chang, Mycol. Res. 96(6):
481 (1992)
Asexual morph Colonies scattered, glistening, black.
Mycelium mostly immersed, consisting of hyaline hyphae.
Conidiophores micronematous or semi-micronematous,
mononematous, solitary, erect, smooth, pale brown, septate, sometimes reduced to conidiogenous cells. Conidiogenous cells monoblastic, integrated, terminal, cylindrical,
hyaline to pale brown. Conidia acrogenous, fusiform,
ellipsoidal or obovoid, dark brown, paler at the basal cell,
smooth, 2–5-euseptate, with darker bands obscuring the
septa. Sexual morph Ascomata immersed, semi-immersed,
or superficial, solitary or aggregated. perithecioid, globose
or subglobose, coriaceous, brown or black, papillate, ostiolate. Ostiole mostly central or if ascomata lying horizontal
to the host surface, then at one end and curving upwards,
long or short, usually brown or black, periphysate. Peridium comprising several layers of brown thick-walled
compressed angular cells, hyaline at the inner walls. Paraphyses hyaline, filiform, tapering distally, septate. Asci
8-spored, unitunicate, cylindrical, pedicellate, truncate at
the apex, with a distinct non-amyloid apical annulus.
Fungal Diversity (2019) 99:451–660
569
Fig. 45 Dematiosporium aquaticum (HKAS 92564, holotype) a Colony on natural substrate. b–k Conidia. Scale bars: b 25 lm, c–k 15 lm
Ascospores uniseriate or overlapping biseriate, fusiform,
straight or somewhat curved, 3–7-euseptate, sometimes
slightly constricted at the septa, with central cells brown,
end cells hyaline or pale colour, smooth, thick-walled, with
or without mucilaginous sheaths and appendages.
Type species: Ascotaiwania lignicola Sivan. & H.S.
Chang, Mycol. Res. 96(6): 481 (1992)
Notes: Ascotaiwania was introduced by Sivanesan and
Chang (1992) with A. lignicola as the type species. Savoryella is comparable to Ascotaiwania, but species of
Savoryella have 3-euseptate ascospores, while in
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Ascotaiwania, they can be 37-euseptate. Molecular studies
revealed the placement of Ascotaiwania within Savoryelales along with the genera Canalisporium (Ascothailandia) and Savoryella, except A. hughesii which was placed
in Pleurotheciales and A. persoonii (Pseudoascotaiwania
persoonii) to Fuscosporellales (Boonyuen et al. 2011;
Réblová et al. 2016a; Yang et al. 2016a). However, Hernández-Restrepo et al. (2017) investigated the phylogeny
of Savoryelales and reported Ascotaiwania was polyphyletic within the order. We follow Boonyuen et al.
(2011), Réblová et al. (2016c) and Yang et al. (2016) to
treat Ascotaiwania as a monophyletic clade in
Savoryelales.
Ascotaiwania fusiformis Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
on decaying submerged wood in a stream (Yang et al.
2016a).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1156, isotype HKAS 95048.
LSU, SSU and RPB2 sequence data are available. Yang
et al. (2016a) introduced this asexual morph species based
on collection from freshwater habitats in Thailand and it is
only known from the type locality.
Ascotaiwania hsilio H.S. Chang & S.Y. Hsieh
Distribution: China, Taiwan, on submerged wood
(Chang et al. 1998); USA, (Fallah et al. 1999; Raja et al.
2009b).
Asexual morph: see Chang (2001)
Notes: Holotype HAST, Chang WL0121-K-3. Sequence
data is not available.
Ascotaiwania hughesii Fallah et al.
Distribution: USA, Wisconsin, Sparkling lake and Trout
lake, on submerged wood (Fallah et al. 1999).
Asexual morph: Helicoon farinosum Linder (Fallah
et al. 1999)
Notes: Holotype ILLS 53605, other specimens collected
from freshwater ILLS 53607, ILLS 53608, ILLS 53609,
ILLS 53610. LSU sequence data is available.
Ascotaiwania limnetica (H.S. Chang & S.Y. Hsieh)
Réblová & J. Fourn.
: Savoryella limnetica H.S. Chang & S.Y. Hsieh,
Mycol. Res. 102: 715. (1998)
Distribution: China, Taiwan, on submerged wood
(Chang et al. 1998); France, on submerged wood (Van
Vooren 2010; Réblová et al. 2016a); India, north Maharashtra, on submerged wood (Patil and Borse 2011);
Asexual morph: see Réblová et al. (2016c)
Notes: Holotype HAST, Chang WL0713-T-2, other
specimens collected from freshwater PRM 933849, PRM
933851. ITS, LSU, SSU and MCM7 sequence data are
available.
Ascotaiwania mitriformis Ranghoo & K.D. Hyde
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Distribution: China, Hong Kong, Plover Cove Reservoir, on submerged wood (Ranghoo and Hyde 1998).
Asexual morph: see Ranghoo and Hyde (1998)
Notes: Holotype IFRD 8696. LSU sequence data is
available. This species is only known from the type
locality.
Ascotaiwania pallida K.D. Hyde & Goh
Distribution: Australia, on submerged wood (Vijaykrishna and Hyde 2006); Brunei, Tutong River, on submerged wood (Fryar et al. 2004); UK, Gloucestershire, nr
Cheltenham, Chedworth Woods, River Coln, on submerged wood (Hyde and Goh 1999).
Asexual morph: Undetermined
Notes: Holotype IFRD 8697. Sequence data is not
available.
Ascotaiwania palmicola K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood in a stream in Mt. Lewis (Hyde and Goh 1997)
Asexual morph: Undetermined
Notes: Holotype BRIP 22744. Sequence data is not
available. This species is only known from the type
locality.
Ascotaiwania sawadae H.S. Chang & S.Y. Hsieh
Distribution: China, on submerged wood (Chang et al.
1998); Thailand, Khao Yai National Park, on well-rotted,
decorticated hard wood, submerged in a freshwater stream
(Sivichai et al. 1998a).
Asexual morph: see state Sivichai et al. (1998a)
Notes: Holotype HAST, Chang HTP0329-A0-2. ITS,
LSU, SSU, RPB2 and TEF1a sequence data are available.
This species is only known from the type locality.
Ascotaiwania wulai H.S. Chang & S.Y. Hsieh
Distribution: China, Taiwan, on submerged wood in
freshwater stream (Chang et al. 1998).
Asexual morph: Undetermined
Notes: Holotype HAST, Chang HTP0421-W2.
Sequence data is not available. This species is only known
from the type locality.
Canalisporium Nawawi & Kuthub., Mycotaxon 34(2): 477
(1989)
Asexual morph Colonies sporodochial, scattered, punctiform, dark brown to black, glistening. Mycelium mostly
immersed, consisting of branched, septate, smooth, thinwalled, pale to brown hyphae. Conidiophores micronematous or semi-macronematous, septate, simple or sparsely
branched, loosely fasciculate, becoming vesiculate and
disintegrating as conidia mature, subhyaline or lightly
pigmented, smooth, thin-walled. Conidiogenous cells
holoblastic, monoblastic, integrated, terminal, determinate,
hyaline or pale brown, globose to ellipsoid. Conidia
acrogenous, solitary, dry, pale or pigmented, muriform,
with ordered arrangement of vertical and transverse septa,
Fungal Diversity (2019) 99:451–660
slightly constricted at the septa, complanate, usually with a
pale basal cell, rarely with several small cells at the base,
with cell lumen connected by septal canals, sometimes
with one-cell layer thickened, smooth-walled. Sexual
morph Ascomata immersed, semi-immersed or superficial,
perithecioid, globose or subglobose, dark brown to black,
solitary or scattered, ostiolate. Ostiole mostly central but
curving upwards at one end if ascomata lying horizontal to
the host surface, long or short, usually brown or black,
periphysate. Paraphyses hypha-like, tapering distally, not
embedded in a gelatinous matrix. Asci 8-spored, unitunicate, long cylindrical, pedunculate, truncate at the apex,
with a refractive non-amyloid apical ring. Ascospores
uniseriate, fusiform, straight or slightly curved, 3-euseptate
and versicolurus.
Type species: Canalisporium caribense (Hol.-Jech. &
Mercado) Nawawi & Kuthub., Mycotaxon 34(2): 479
(1989)
Notes: Canalisporium was established to accommodate
Berkleasmium caribense, B. pulchrum with a new species
C. elegans. Twelve species are included in the genus with
ten species encountered in freshwater habitats. Sri-indrasutdhi et al. (2010) introduced the only known sexual
morph Ascothailandia grenadoidea in the genus with
Canalisporium asexual morph sporulating from the culture.
Further molecular studies supported the connection
between Ascothailandia and Canalisporium and placed the
genus within Savoryelales (Sri-indrasutdhi et al. 2010;
Jones et al. 2016; Réblová et al. 2016a). With the one
fungus one name (Hawksworth 2011), the genus Canalisporium has priority and thus Ascothailandia was synonymized under Canalisporium (Réblová et al. 2016c).
Canalisporium caribense (Hol.-Jech. & Mercado) Nawawi
& Kuthub.
: Berkleasmium caribense Hol.-Jech. & Mercado,
Česká Mykol 38: 99 (1984)
Distribution: Brunei, Temburong, Batu Apoi Forest
Reserve, on submerged wood (Goh et al. 1998a); China,
Hong Kong, New Territories, Tai Po, Tai Po Kau Country
Park, on submerged wood (Goh et al. 1998a); India,
Karnataka, River Kali (Sridhar et al. 2011), North Maharashtra (Patil et al. 2014); Malaysia, submerged wood in
stream (Ho et al. 2001); Panama, Colon Province,
Soberania National Park/Juan Grande River/Limbo River/
Conrad stream, on submerged decoricated wood (Ferrer
and Shearer 2005); Thailand, Chiang Rai Province, Hui
Kang Pla waterfall National Park/Khun Korn Waterfall, on
submerged wood (Zhang et al. 2014); USA, Florida (Raja
et al. 2009b).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater MFLU
11–1071. ITS, LSU, SSU and RPB2 sequence data are
available.
571
Canalisporium elegans Nawawi & Kuthub
Distribution: Brunei, Temburong, Batu Apoi Forest
Reserve, on submerged wood (Goh et al. 1998a); Panama,
Colon Province, Soberania National Park/Limbo River/
Frijoles River/Gatum lake/Wheeler stream, on submerged
decoricated wood (Ferrer and Shearer 2005); Thailand,
Chiang Rai Province, Hui Kang Pla waterfall National
Park, on submerged wood (Zhang et al. 2014).
Sexual morph: Undetermined
Notes: Holotype IMI 326603, other specimen collected
from freshwater MFLU 10–0181. ITS, LSU, SSU, RPB2
and TEF1a sequence data are available.
Canalisporium exiguum Goh & K.D. Hyde
Distribution: Australia, north Queensland, Atherton
Tablelands, Lake Barrine, on submerged wood (Goh et al.
1998a);
Sexual morph: Undetermined
Notes: Holotype IFRD 8720. ITS, LSU, SSU and RPB2
sequence data are available.
Canalisporium grenadoideum Sri-indr. et al.
Distribution: Thailand, Narathiwat Province, Hala Bala
Wildlife Sanctuary, on submerged wood of Wrightia
tomentosa (Sri-indrasutdhi et al. 2010).
Sexual morph: see Sri-indrasutdhi et al. (2010)
Notes: Holotype BBH 26384. LSU, SSU, RPB2 and
TEF1a sequence data are available.
Canalisporium jinghongensis Cai et al.
Distribution: China, Yunnan Province (Cai et al.
2003c); Thailand, Chiang Rai Province, Hui Kang Pla
waterfall National Park, on submerged wood (Zhang et al.
2014).
Sexual morph: Undetermined
Notes: Holotype PDD 74130. ITS, LSU, SSU, RPB2
and TEF1a sequence data are available.
Canalisporium kenyense Goh et al.
Distribution: Panama, Colon Province, Conrad stream/
Wheeler stream on submerged decoricated wood (Ferrer
and Shearer 2005); USA, Florida (Raja et al. 2009b).
Sexual morph: Undetermined
Notes: Holotype IMI 285428a. Sequence data is not
available.
Canalisporium pallidum Goh et al.
Distribution: China, Hong Kong, New Territories, Tai
Po, Tai Po Kau Country Park, on submerged wood (Goh
et al. 1998a).
Sexual morph: Undetermined
Notes: Holotype HKU (M) 5903 (now in IFRD). ITS,
LSU, SSU, RPB2 and TEF1a sequence data are available.
This species is only known from the type locality.
Canalisporium panamense A. Ferrer & Shearer
Distribution: Panama, Colon Province, Soberania
National Park, on submerged decoricated wood (Ferrer and
Shearer 2005).
123
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Fungal Diversity (2019) 99:451–660
Fig. 46 Chaetosphaeria aquatica (MFLU 18–1618, holotype). a–d Conidiophores with conidia. e, f Conidiogenous cells. g–j conidia.
k Germinating conidium. l, m Culture on PDA from above and reverse. Scale bars: a–d 30 lm, e–f 15 lm, g–k 10 lm
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Sexual morph: Undetermined
Notes: Holotype ILL, Ferrer E074–1. Sequence data is
not available.
Canalisporium pulchrum (Hol.-Jech. & Mercado) Nawawi
& Kuthub
: Berkleasmium pulchrum Hol.-Jech. & Mercado,
Česká Mykol 38(2): 101 (1984)
Distribution: Australia, north Queensland, Atherton
Tablelands, Lake Barrine, on submerged wood; Brunei,
Temburong, Batu Apoi Forest Reserve, on submerged
wood, China, Hong Kong, New Territories, Plover Cove
Reservoir, on submerged wood (Goh et al. 1998a);
Panama, Colon Province, Frijoles stream, on submerged
decoricated wood (Ferrer and Shearer 2005); Thailand,
Chiang Rai Province, Hui Kang Pla waterfall National
Park, Chiang Mai Province, Doi Inthanon National Park,
on submerged wood (Zhang et al. 2014).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater MFLU
10–0168, MFLU 11–0973. ITS, LSU, SSU, RPB2 and
TEF1a sequence data are available.
Canalisporium variabile Goh & K.D. Hyde
Distribution: Australia, Queensland, Atherton Tablelands, Cowry Creek, on wood submerged in creek (Goh
and Hyde 2000a).
Sexual morph: Undetermined
Notes: Holotype IFRD 8721. Sequence data is not
available.
Dematiosporium Z.L. Luo, K.D. Hyde & H.Y. Su, gen.
nov.
Index Fungorum number: IF 555672, Facesoffungi
number: FoF 05455
Etymology: ‘‘Dematiosporium’’ referring to dematiaceous conidia of this genus.
Saprobic on decaying wood. Asexual morph Colonies
on natural substrate superficial, effuse, dark brown to
black. Conidiophores reduced to conidiogenous cells.
Conidia dry, dark brown to black, mostly globose to subglobose, smooth. Sexual morph Undetermined.
Type species: Dematiosporium aquaticum Z.L. Luo,
K.D. Hyde & H.Y. Su
Notes: During our study on lignicolous freshwater fungi
of Erhai lake, a hyphomycetous fungus was collected from
submerged wood. Morphologically, our collection resembles asexual morph of Conioscypha in having micronematous conidiophores, dry, dark brown to black, globose to
subglobose conidia. However, phylogenetic analysis shows
that our collection (MFLU 18–1641) belongs in
Savoryellaceae (Savoryellales), while Conioscypha
belongs in Conioscyphaceae (Conioscyphales) (Fig. 1,
clade 44, 45). Phylogenetic analysis also shows that the
species (MFLU 18–1641) is distinct from other genera in
573
Savoryellaceae (Fig. 1, clade 44). We therefore introduce a
new genus Dematiosporium to accommodate this asexual
fungus.
Dematiosporium aquaticum Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555673, Facesoffungi
number: FoF 05456, Fig. 45
Etymology: Referring to the aquatic habitat of this
fungus.
Holotype: MFLU 18–1641
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, dark brown to black. Conidiophores
reduced to conidiogenous cells. Conidia 25–31 lm long,
24–28 lm wide (
x = 28 9 26 lm, n = 20), dry, mostly
globose to subglobose, dark brown to black, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Erhai
lake, saprobic on submerged decaying wood, April 2015,
Q. Dai, S-623 (MFLU 18–1641, holotype; HKAS 92564,
isotype).
Notes: Dematiosporium aquaticum mostly resembles
Conioscypha aquatica in having micronematous conidiophores, dry, dark brown to black, globose to subglobose
conidia. However, Dematiosporium aquaticum differs from
Conioscypha aquatica in having larger conidia
(25–31 9 24–28 vs. 19–23 9 17–21 lm) and D. aquaticum belongs to Savoryellaceae while C. aquatica belongs
to Conioscyphaceae.
Savoryella E.B.G. Jones & R.A. Eaton, Trans. Br. mycol.
Soc. 52(1): 161 (1969)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Maharachchikumbura et al.
(2016).
Type species: Savoryella lignicola E.B.G. Jones & R.A.
Eaton, Trans. Br. Mycol. Soc. 52(1): 162 (1969)
Notes: The genus Savoryella was established by Jones
and Eaton (1969) with S. lignicola as the type species
which colonized on the test-blocks in water cooling towers.
Savoryella species are commonly reported from submerged
wood in aquatic habitats and among the ten known species,
of which six are from freshwater, four from marine or
brackish water (including water cooling towers), while S.
melanospora is known on driftwood from coastal sand
dunes in Australia (Abdel-Wahab and Jones 2000; Jones
et al. 2015, 2016). Savoryella is shown to be a monophyletic group as sister clade to Ascotaiwania and Canalisporium within Savoryellales (Boonyuen et al. 2011;
Réblová et al. 2016a). Jones et al. (2016) provided a review
on the genus, which discussed the history, significance of
the genus, illustration of its morphology and discussed its
role in the colonization and biodeterioration of
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Fungal Diversity (2019) 99:451–660
Fig. 47 Chaetosphaeria catenulata (MFLU 18–1620, holotype) a, b Colonies on wood. c, d Conidiophores with conidia. e, f Conidiogenous
cells. g–k Conidia. l Germinating conidium. m, n Culture on PDA from above and reverse. Scale bars: c, d 50 lm, e–g 15 lm, h–l 10 lm
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575
Fig. 48 Chaetosphaeria cubensis (MFLU 18–1621) a, b Colonies on wood. c d Conidiophores with conidia. e, f Conidiogenous cells with
conidia. g–n conidia. o Germinating conidium. p, q Culture on PDA from above and reverse. Scale bars: c, d 40 lm, e, f 20 lm, g–o 5 lm
123
576
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Fig. 49 Chaetosphaeria guttulata (MFLU 18–1617, holotype). a, b Conidiophores with conidia. c, d Conidiogenous cells. e–i conidia.
j Germinating conidia. k, l Culture on PDA from above and reverse. Scale bars: a, b 35 lm, c, d 20 lm, e–j 10 lm
123
Fungal Diversity (2019) 99:451–660
lignocellulosic materials. The role of Savoryella in soft
rotten decaying wood was also discussed (Jones et al.
2016).
Savoryella aquatica K.D. Hyde
Distribution: Australia, north Queensland, Clohesy
River, on submerged wood in freshwater (Hyde 1993a);
Brunei, Temburong, Sungai Sitam, on submerged wood;
Malaysia, Kuala Lumpur, State Negara, Lipur Lentang
Nature Reserve, on submerged wood; Philippines, Mindanao, Bukidnon, Natigbasam Creek, on submerged wood
(Ho et al. 1997); South Africa, Durban, Inharca River, on
submerged wood (Ho et al. 1997), Palmiet River, on submerged wood (Hyde et al. 1998b); Thailand, Chiang Mai
Province, on submerged wood (Ho et al. 1997).
Asexual morph: Undetermined
Notes: Holotype BRIP 19327, other specimen collected
from freshwater habitats: BRIP 20613. ITS, LSU, SSU,
RPB1, RPB2 and TEF1a sequence data are available.
Savoryella curvispora Ho et al.
Distribution: Malaysia, State Negara, Lipur Lentang
Nature Reserve, in ramulo submerse; Mauritius, Tamarin,
Black River, on submerged wood; Philippines, Luzon,
Laguna, Los Ban4 os, Mt Makiling, on wood submerged in
a small stream; South Africa, Durban, Palmiet River, on
submerged wood (Ho et al. 1997).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2881 (now in IFRD).
Sequence data is not available.
Savoryella fusiformis Ho et al.
Distribution: Brunei, Temburong, Sungai Sitam, on
submerged wood (Ho et al. 1997).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 2923 (now in IFRD). ITS,
MCM7, SSU, RPB2 and TEF1a sequence data are
available.
Savoryella grandispora K.D. Hyde
Distribution: Australia, Cape Tribulation, Cow Bay, on
wood submerged in a River, Malaysia, State Negara, Lipur
Lentang Nature Reserve, on submerged wood (Ho et al.
1997).
Asexual morph: Undetermined
Notes: Holotype BRIP 20918, other specimen collected
from freshwater habitats: BRIP 20919. Sequence data is
not available.
Savoryella lignicola E.B.G. Jones & R.A. Eaton
Distribution: Australia, north Queensland, Clohesy
River, on submerged wood in freshwater (Hyde 1993a),
Queensland, on submerged wood in a stream in Mt. Lewis
(Hyde and Goh 1997); Brunei, Temburong, Sungai Sitam,
on submerged wood, China, Hong Kong, New Territory,
Tai Po Kau Forest Stream, on submerged wood (Ho et al.
1997); Japan, Koito River, on submerged wood (Tsui et al.
2001a); Malaysia, State Negara, Lipur Lentang Nature
577
Reserve, on submerged wood, South Africa, Durban,
Palmiet River, on submerged wood (Ho et al. 1997).
Asexual morph: Undetermined
Notes: Holotype IMI 129784. ITS, LSU, SSU, RPB2
and TEF1a sequence data are available.
Savoryella verrucosa Minoura & T. Muroi
Distribution: Australia, north Queensland, Crystal
Cascade, near Cairns, on wood submerged in a River,
Brunei, Temburong, Sungai Sitam, on submerged wood,
Japan, Tochigi Pref. Nasugun, Nishinasuno, Shizunuma
Pond, on submerged balsa wood, Malaysia, State Negara,
Lipur Lentang Nature Reserve, on submerged wood (Ho
et al. 1997).
Asexual morph: Undetermined
Notes: Holotype HUT 40006. ITS, LSU, SSU, RPB2,
TEF1a and MCM7 sequence data are available.
Savoryellomycetidae genera incertae sedis
Flammispora Pinruan et al., Stud Mycol 50(2): 384 (2004)
Asexual morph Undetermined. Sexual morph Ascomata
immersed or partially immersed, coriaceous, ostiolate,
solitary, black. Asci unitunicate, 8-spored, clavate to
cylindrical clavate, or fusiform, deliquescent, thin-walled.
Ascospores biseriate, fusiform or cylindrical, hyaline,
septate, appendaged.
Type species: Flammispora bioteca Pinruan, Sakay.,
K.D. Hyde & E.B.G. Jones, Stud. Mycol. 50(2): 384 (2004)
Notes: The genus Flammispora was introduced by
Pinruan et al. (2004a) with F. bioteca as type species.
Flammispora resembles aquatic genera with polar appendaged ascospores, especially some species with cylindrical
to filiform ascospores and the taxa such as Ascosacculus
aquaticus, Ascosalsum cincinnatulum, A. viscidulum and A.
unicaudatum. However, these species differ from Flammispora in having hamate polar appendages, initially closely adpressed to the ascospore wall, then separating and
eventually unfurling to form long, narrow appendages
(Campbell et al. 2003b). Two species were accepted in the
genus Flammispora, and F. pulchra occurred in a lake,
while F. bioteca was reported from a peat swamp (Pinruan
et al. 2004a; Raja and Shearer 2008).
Flammispora bioteca Pinruan et al.
Distribution: Thailand, Narathiwat, Sirindhorn Peat
Swamp Forest, on submerged dead leaves of Licuala
longecalycata (Pinruan et al. 2004a).
Asexual morph: Undetermined
Notes: Holotype BBH WAH 134. SSU sequence data is
available.
Flammispora pulchra Raja & Shearer
Distribution: USA, Florida, Wildcat Lake, on submerged decorticated woody debris (Raja and Shearer 2008)
Asexual morph: Undetermined
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Fig. 50 Chaetosphaeria myriocarpa (HKAS 92985) a Colonies on wood. b–e Conidiophores with conidia. f Conidiogenous cells. g Conidia. h,
i Culture on PDA from surface and reverse. Scale bars: b–e 30 lm, f 10 lm, g 5 lm
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Fig. 51 Phylogram generated from maximum likelihood analysis
based on ITS and LSU sequence data for species of Chaetosphaeriaceae (with Sordaria fimicola and Gelasinospora tetrasperma as
outgroup). The best scoring RAxML tree with a final likelihood value
of - 20964.682820 is presented. RAxML bootstrap support values
equal to or greater than 75% are given before the forward slash
579
(black). Bayesian posterior probability equal to or higher than 0.95
are given after the forward slash (red). Hyphen (‘-’) indicates a value
lower than 75% for RAxML and bayesian posterior probability lower
than 0.95. Newly generated sequences are in red. Ex-type or exepitype strains are in bold
123
580
Fungal Diversity (2019) 99:451–660
Fig. 51 continued
Notes: Holotype ILL 40119. Sequence data is not
available. This species is only known from the type
locality.
123
Subclass Sordariomycetidae O.E. Erikss & Winka
Chaetosphaeriales Huhndorf et al.
Chaetosphaeriaceae Réblová et al.
Fungal Diversity (2019) 99:451–660
581
Fig. 51 continued
Brunneodinemasporium Crous & R.F. Castañeda, Persoonia 28: 128 (2012)
Asexual morph descriptions and illustrations refer to
Crous et al. (2012). Sexual morph Undetermined.
Type species: Brunneodinemasporium brasiliense Crous
& R.F. Castañeda, Persoonia 28: 129 (2012)
Notes: Brunneodinemasporium was introduced by Crous
et al. (2012) with B. brasiliense as the type species, to
accommodate a Dinemasporium-like species which has
tightly aggregated brown conidiogenous cells and pale
brown conidia. Lu et al. (2016) introduced the second
species for this genus and named as Brunneodinemasporium jonesii from freshwater habitats in China.
Brunneodinemasporium jonesii Lu et al.
Distribution: China, Guangxi Province, on decaying
wood submerged in a freshwater stream (Lu et al. 2016).
Sexual morph: Undetermined
123
582
Notes: Holotype GZAAS 16–0062. ITS and LSU
sequence data are available. This species is known so far
only from the type locality.
Chaetosphaeria Tul. & C. Tul., Select. fung. carpol. (Paris)
2: 252 (1863)
Holomorph
Descriptions
and
illustration
see
Maharachchikumbura et al. (2016).
Type species: Chaetosphaeria innumera Berk. & Broome
ex Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 252 (1863)
Notes: The genus Chaetosphaeria was introduced by
Tulanse and Tulanse (1863) and is typified by C. innumera.
The genus Chaetosphaeria is a lignicolous perithecial
ascomycete and the genus has world-wide distribution. It is
characterized by non-stromatic, dark, papillate ascomata,
persistent paraphyses, unitunicate asci with a shallow,
refractive Japical annulus and hyaline, ellipsoidal, fusiform
to filiform, one to several-septate hyaline ascospores,
although several species with versicolorous ascospores are
also accommodated in the genus. Four species in this genus
have been reported from freshwater habitats. Conidial
phases in the genus Chaetosphaeria largely resemble
asexual morph genus Catenularia Grove.
Chaetosphaeria anglica P.J. Fisher & Petrini
Distribution: Brunei, Batu Apoi Forest Reserve, on
submerged wood (Ho et al. 2001); Sudan, Blue Nile, on
submerged wood (Fisher and Petrini 1983); UK, River Exe,
on submerged wood (Fisher and Petrini 1983).
Asexual morph: Undetermined
Notes: Holotype IMI 273041, other specimen collected
from freshwater habitats: IMI 273042; Sequence data is not
available.
Chaetosphaeria aquatica Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555674, Facesoffungi
number: FoF 05457, Fig. 46
Etymology: Referring to the aquatic habitat of this fungus
Holotype: MFLU 18–1618
Saprobic on submerged decaying wood. Asexual
morph Colonies amphigenous, effuse, brown or black,
thinly hairy. Mycelium partly superficial, partly immersed,
composed of septate, branched, dark brown, smooth
hyphae. Conidiophores (111–)181–271 lm long, 5–7 lm
wide
(
x = 226 9 6 lm,
n = 20),
monomenatous,
macronematous, solitary or in small groups, erect, straight
or slightly flexuous, dark brown at base, pale brown at
apex, septate, unbranched, percurrently growing, smooth.
Conidiogenous cells monophialidic, with conspicuous
collarette, integrated, percurrently proliferating, terminal,
later becoming subterminal, pale brown. Conidia 20–
24 lm long, 6–8 lm wide (
x = 22 9 7 lm, n = 20), solitary, acropleurogenous, cylindro-oblong to reniform,
rounded at apex, pointed ant base, straight or slightly
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curved, 1–3-septate, greyish brown when yong, subhyaline
at the end cells, brown at the 3 cells, remaining attached to
the sides of conidiogenous cells on release, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, May 2015,
Z.L. Luo, S-752 (MFLU 18–1618, holotype), ex-type living culture MFLUCC 18–1341.
Notes: Chaetosphaeria aquatica resembles asexual
morph of C. conirostris in having multiseptate, cylindrical,
unbranched conidiophores and oblong to reniform conidia
with similar size (Fernández and Huhndorf 2005). However, Chaetosphaeria aquatica differs from C. conirostris
in having terminal to subterminal conidiogenous cells, 1–3septate, grayish brown to brown conidia. Phylogenetic
analysis based on ITS and LSU sequence data also shows
that Chaetosphaeria aquatica is distinct from C. conirostris (Fig. 51).
Chaetosphaeria catenulata Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555675, Facesoffungi
number: FoF 05458, Fig. 47
Etymology: Referring to the catenulate conidia of this
fungus.
Holotype: MFLU 18–1620
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on the substratum
superficial, scattered, brown. Mycelium immersed, brown
to dark brown, composed of septate, brown hyphae.
Conidiophores 187–283 lm long, 6–10 lm wide
(
x = 235 9 8 lm, n = 20), macronematous, mononematous, cylindrical, single, erect, straight, unbranched, thickwalled, dark brown, becoming paler towards the apex,
smooth. Conidiogenous cells monophialidic, integrated,
terminal, cylindric-clavate, flared collarette. Conidia 13–
15 lm long, 12–14 lm wide (
x = 14 9 13 lm, n = 20),
acrogenous, formed in chains, aseptate, turbinate-triangular, with 3 blunt protruding edges at the broader distal end,
viewed from above 3-lobed or cruciform with blunt protruding corners, hyaline to subhyaline when yong, greyish
brown at mature, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-891 (MFLU 18–1620, holotype), ex-type
living culture DLUCC 0891; saprobic on decaying wood
submerged in Nujiang River, May 2015, Z.L. Luo, S-298.
Notes: Chaetosphaeria catenulata resembles C. cubensis in having mononematous, macronematous, septate,
unbranched, cylindrical conidiophores, integrated, terminal, monophialidic, percurrent conidiogenous cells and
cuneiform, aseptate conidia (Holubova-Jechova 1982; this
study). However, Chaetosphaeria catenulata differs from
Fungal Diversity (2019) 99:451–660
C. cubensis in having larger conidia (13–15 9 12–14 vs.
5.5–9 9 3.5–5.5 lm). Phylogenetic analysis also shows
that Chaetosphaeria catenulata is distinct from C. cubensis
(Fig. 51)
Chaetosphaeria cubensis Hol.-Jech.
Facesoffungi number: FoF 05459, Fig. 48
Saprobic on submerged decaying wood. Asexual
morph Colonies on substratum, effuse, brown to black,
superficial, groups, hairy. Mycelium partly superficial,
composed of septate, branched, brown hyphae. Conidiophores 175–219 lm long, 5–7 lm wide (
x = 197 9 6 lm,
n = 20), mononematous, macronematous, cylindrical,
solitary or in small groups, straight or slightly flexuous,
septate, greyish brown, gradually becoming paler towards
the apex, smooth, septate, percurrently proliferating.
Conidiogenous cells monophialidic, integrated, terminal,
percurrent, calyciform. Conidia 6–8 lm long, 4–6 lm
wide (
x = 7 9 5 lm, n = 20), cuneiform, with 3 blunt
protruding edges at the broader distal end, aseptate, greyish
brown to brown, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on submerged decaying wood, May 2015,
Z.L. Luo, S-375 (MFLU 18–1621), living culture
MFLUCC 18–1331.
Notes: Morphologically, our collection fits well to the
Chaetosphaeria species with Catenularia asexual morph,
such as Ch. cubensis, Ch. cupulifera and Ch. novaezelandiae (Réblová 2004). The new collection resembles Ch.
cubensis in having mononematous, macronematous, septate, unbranched, cylindrical conidiophores with similar
size, integrated, terminal, monophialidic, percurrent conidiogenous cells and cuneiform, aseptate conidia with
similar size (Holubova-Jechova 1982). Phylogenetic analysis based on ITS and LSU sequence data shows that our
isolate (MFLUCC 18–1331) clusters with Ch. cubensis
(SMH 3258) with high support values (100 ML/1.00 PP)
(Fig. 51). Based on the morphology and phylogeny, we
identify our isolate as Ch. cubensis and it is the first record
for China and first collection from freshwater habitats as
well.
Chaetosphaeria guttulata Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555676, Facesoffungi
number: FoF 05460, Fig. 49
Etymology: Referring to the conidia with large guttules.
Holotype: MFLU 18–1617
Saprobic on submerged decaying wood. Asexual
morph Colonies on substratum superficial, effuse, scattered, hairy, dark brown. Mycelium partly immersed,
composed of septate brown, smooth, hyphae. Conidiophores 115–157 lm long, 5–7 lm wide (
x = 136 9 6 lm,
n = 20), macronematous, mononematous, erect, straight or
583
slightly flexuous, septate, solitary, unbranched, smooth,
brown to dark brown and gradually becoming paler
towards the apex. Conidiogenous cells holoblastic, polyblastic, with many tiny protuberant conidiogenous loci,
integrated, terminal, light brown or subhyaline, Conidia
17–21 lm long, 5–7 lm wide (
x = 19 9 6 lm, n = 20),
acropleurogenous,, ovoid or fusiform, dry, 3-septate, guttulate, rounded at apex, pointed at base, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan
Mountain, saprobic on submerged decaying wood in a freshwater stream, June 2016, Q.X. R, S-771 (MFLU 18–1617,
holotype), ex-type living culture MFLUCC 17–1703.
Notes: Chaetosphaeria guttulata resembles asexual
morph of Chaetosphaeria lignomollis in having conidiophores septate, unbranched, brown to dark brown and
gradually becoming paler towards the apex, conidiogenous
cells polyblastic, integrated, terminal and conidia septate,
hyaline, rounded at apex (Fernández and Huhndorf 2005).
However, Chaetosphaeria guttulata differs from C. lignomollis by its ovoid or fusiform, guttulate, smaller conidia
(17–21 9 5–7 vs. 26–31 9 7.3–8.6 lm). Phylogenetic
analysis also shows that Chaetosphaeria guttulata and C.
lignomollis are distinct species (Fig. 51). Chaetosphaeria
guttulata clusters with C. myriocarpa (MUCL34784) and
C. pygmaea (UPSC2523) with strong support (Fig. 51),
however, their morphology are obviously different (Booth
1957; Gams and Holubová-Jechová 1976).
Chaetosphaeria lentomita W. Gams & Hol.-Jech
Distribution: China, Hong Kong, on submerged wood
(Ho et al. 2001).
Asexual morph: Chloridium pachytrachelum
Notes: Holotype CBS 645.75. ITS and LSU sequence
data are available. Ho et al. (2001) found this species from
freshwater in China, but did not provide description and
illustration in their study.
Chaetosphaeria lignomollis F.A. Fernández & Huhndorf
Distribution: Brazil, Bahia, on submerged twig (Barbosa et al. 2013).
Asexual morph: see Fernández and Huhndorf (2005).
Notes: Holotype F, Huhndorf 3015 (SMH 3015), specimens collected from freshwater habitats: HUEFS 158056,
HUEFS 158081. ITS, LSU and b-tubulin sequence data are
available. Barbosa et al. (2013) collected this species from
freshwater habitats in Brazil and provided descriptions for
this fungus.
Chaetosphaeria myriocarpa (Fr.) C. Booth
: Sphaeria myriocarpa Fr., Syst. mycol. 2(2): 459
(1823)
Facesoffungi number: FoF 05461, Fig. 50
Saprobic on submerged decaying wood. Sexual morph:
Undetermined. Asexual morph Colonies on the substratum
superficial, scattered, hairy, brown to dark brown.
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Fig. 52 Chaetosphaeria submersa (MFLU 18–1616, holotype). a, b Colonies on wood. c–e Conidiophores with conidia. f, g Conidiogenous
cells. h–k Conidia. Scale bars: c, e 50 lm, d 40 lm, f–g 30 lm, h–k 15 lm
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Mycelium mostly immersed, consisting of branched, septate, brown, smooth hyphae. Conidiophores 63–117 lm
long, 2.5–3.5 lm wide (
x = 90 9 3 lm, n = 20),
macronematous, mononematous, cylindrical, straight or
flexuous, septate, dark brown at the base and gradually
paler towards the apex, unbranched, percurrently growing,
smooth-walled. Conidiogenous cells monophialidic, percurrent, determinate, integrated, with flared pale brown
collarette. Conidia 1–2 lm (
x = 1.5 lm, SD = 0.5, n = 20)
diam., sphaerical or subsphaerical, aseptate, hyaline,
smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Jinsha
River, saprobic on submerged decaying wood, April 2015,
X.J. Su, S-408 (HKAS 92985), living culture MFLUCC
16–0977, KUMCC 15–0416.
Notes: Chaetosphaeria myriocarpa was described by
Fries (1823) under Sphaeria as S. myriocarpa. Booth
(1957) synonymized Sphaeria myriocarpa as Chaetosphaeria myriocarpa based on morphology. Crous et al.
(2018) collected an asexual morph of Chaetosphaeria
myriocarpa from Ukraine and provided ITS, LSU and btubulin sequence data for this species. During the study of
lignicolous freshwater fungi in China, a hyphomycetous
fungus was collected from submerged wood in Yunnan
Province and the morphological characters of our fungus
fits well with Chaetosphaeria myriocarpa (Booth 1957;
Crous et al. 2018). Phylogenetic analysis also shows that
our isolate (MFLUCC 16–0977) clusters together with
Chaetosphaeria myriocarpa (Fig. 51). Based on the morphology and phylogeny, we identify our collection as
Chaetosphaeria myriocarpa and it is the first report of this
species from freshwater habitats.
Chaetosphaeria rivularia Réblová & J. Fourn
Distribution: France, Midi-Pyrénées, Ariège, Rimont,
Grand Bois forest, Maury brook, on submerged wood of
Fagus sylvatica associated with Minutisphaera japonica
(Ariyawansa et al. 2015).
Asexual morph: see Ariyawansa et al. (2015).
Notes: Holotype PRM 933847. ITS and LSU sequence
data are available. Ariyawansa et al. (2015) introduced this
species based on collections from freshwater in France and
it is only known from the type locality.
Chaetosphaeria submersa Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555677, Facesoffungi
number: FoF 05462, Fig. 52
Etymology: Referring to the submerged habitat of this
fungus.
Holotype: MFLU 18–1616
Saprobic on submerged decaying wood. Asexual
morph Colonies on the substratum effuse, dark brown,
with glistening mass of conidia at the tip of conidiophores.
Mycelium partly immersed, partly superficial, composed of
585
branched septate, dark brown, smooth hyphae. Conidophores 380–596(–691) lm long, 15–21 lm wide
(
x = 493 9 18 lm, n = 20), macronematous, mononematous, erect, solitary, cylindrical, straight or slightly flexuous, unbranched, percurrently growing, rarely branched at
apex, thick-walled, 9–13-septate, brown to dark brown
below, pale brown to subhyaline toward the apex, smooth.
Conidiogenous cells monophialidic, percurrently proliferating, integrated, terminal, pale brown, cylindric-clavate.
Conidia
21–27 lm
long,
12–14 lm
wide
(
x = 24 9 13 lm, n = 20), acrogenous, solitary, cuneiform, aseptate, guttulate, rounded at apex, truncate at base,
hyaline when young, light brown at mature, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on submerged decaying wood in
freshwater stream, October 2016, Z.L. Luo, S-824 (MFLU
18–1616, holotype), ex-type living culture MFLUCC
18–1342.
Notes: Chaetosphaeria submersa resembles asexual
morph of Ch. novae-zelandiae in having unbranched,
septate conidiophores which brown to dark brown below,
paler toward the apex, cuneiform, aseptate, brown conidia
rounded at apex, truncate at base (Hughes 1965). However,
Ch. submersa differs from Ch. novae-zelandiae by its larger conidiophores (380–596(–691) vs. 50–330 lm) and
conidia (21–27 9 12–14 vs. 12.6–17.5 9 11.4–18.2 lm).
Ch. submersa also resembles Catenularia longispora in
having cylindrical, unbranched, septate conidiophores
which brown to dark brown below, pale brown to subhyaline toward the apex, monophialidic, integrated, terminal conidiogenous cells and aseptate, cuneiform conidia
(Hughes 1965). However, Chaetosphaeria submersa differs from Catenularia longispora in having 9–13-septate,
longer conidiophores (380–596(–691) vs. 200–330 lm)
and
smaller
conidia
(21–27 9 12–14
vs.
27–45 9 16.8–24 lm).
Chloridium Link, Mag. Gesell. naturf. Freunde, Berlin 3(12): 13 (1809)
Asexual morph Colonies effuse, velvety, reverse dark
brown to black. Mycelium partly immersed, partly superficial, composed of branched, hyaline, septate, smooth
hypha. Conidiophores erect, unbranched, pigmented, apical, integrated, arising from submerged hyphae. Conidiogenous cells phialidic to poly-enteroblastic, with a wide or
narrow, deep or short collarette, in some species with
multiple conidiogenous loci. Conidia arranged in slimy
heads or gradually extending cirrhi, rarely in simple chains,
but then mostly imbricate, globose to ellipsoidal or cylindrical, mostly containing one or two guttules, hyaline or
almost hyaline, smooth-walled. Chlamydospores present in
some species, pigmented, globose or elongated, terminal,
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Fig. 53 Chloridium gonytrichii (HKAS 93031) a–c Conidiophore. d, e Conidiogenous cells. f–k Conidia. l Germinating conidium. Scale bars:
a–c 50 lm, d, e 30 lm, f 10 lm, g–l 5 lm
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587
Fig. 54 Codinaea yunnanensis (MFLU 18–1611, holotype). a Colonies on wood. b, d, e Conidiophores with conidia. e Conidiogenous cells
with conidium. f–i Conidia. j Germinating conidium. k, l Culture on PDA from above and reverse. Scale bars: b, e 30 lm, c, d 15 lm, f–j 10 lm
123
588
lateral or more rarely intercalary. Sexual morph Ascomata
broadly ovoid to globose, dark brown, separate, papillate,
with a few, scattered setae, brown, multiseptate, slender,
tapering to an attenuated apex. Ascomatal wall of textura
angularis in surface view, composed of pseudoparenchymatic cells. Ascomatal apex papillate, opaque. Paraphyses
unbranched, septate, hyaline. Asci 8-spored, cylindricalclavate, short-stalked, thin-walled. Ascospores uniseriate,
ellipsoid, septate, easily disarticulating into part-spores,
hyaline (obtained from Fernández and Huhndorf (2005)).
Type species: Chloridium viride Link, Mag. Gesell.
naturf. Freunde, Berlin 3(1–2): 13 (1809)
Notes: Index Fungorum (December 2018) lists 73 epithets of the genus Chloridium which was typified by C.
viride. Three species in this genus have been reported from
freshwater habitats in China.
Chloridium gonytrichii (F.A. Fernández & Huhndorf)
Réblová & Seifert
: Melanopsammella gonytrichii F.A. Fernández &
Huhndorf, Fungal Divers 18: 42 (2005)
: Chloridium aseptatum M.J. Wei & H. Zhang, Phytotaxa 362: 191 (2018)
Facesoffungi number: FoF 05463, Fig. 53
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, hairy, brown to
dark brown. Mycelium superficial. Conidiophores 196–
240 lm long, 6–8 lm wide (
x = 218 9 7 lm, n = 20),
macronematous, mononematous, single, unbranched, dark
brown becoming light brown to subhyaline towards the
apex, multiseptate, with 2–4 whorls of phialides in midsection, with a single phialide at the apex. Conidiogenous
cells phialides, cylindrical to lageniform, producing conidia from multiple entero-blastic conidiogenous loci, phialides borne on collar hyphae around the conidiophore,
percurrent proliferation observed on the substrate. Conidia
3–5 lm long, 2–4 lm wide (
x = 4 9 3 lm, n = 20), globose to subglobose, aseptate, with a guttulate, light green.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April
2015, Z.L. Luo, S-574 (HKAS 93031), living culture
DLUCC 0574; Jinsha River, saprobic on submerged
decaying wood, April 2015, H.Y. Su, S-360, living culture
MFLUCC 16–1111, KUMCC 15–0444; Ibid., saprobic on
submerged decaying wood, April 2015, Z.L. Luo, S-380
(HKAS 93053), living culture MFLUCC 16–1095,
KUMCC 15–0424.
Notes: Holotype SMH 3785, specimens collected from
freshwater habitats: HKAS 93031, MFLU 11–1051. ITS,
LSU, SSU, RPB2 and TEF1a sequence data are available.
Chloridium gonytrichii is originally described by Fernández and Huhndorf (2005) as Melanopsammella gonytrichii.
Réblová et al. (2016b) synonymised Melanopsammella
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gonytrichii under Chloridium gonytrichii. During our study
on lignicolous freshwater fungi in China, three
hyphomycetous fungi were collected, morphological
characters of these fungi fit well with asexual of Chloridium gonytrichii in having macronematous, mononematous,
single, multiseptate, unbranched conidiophores with whorls
of phialides in midsection, globose to subglobose, light
green, aseptate conidia. Based on the morphology, we
identify our isolates as Chloridium gonytrichii and provided the sequence data for Chloridium gonytrichii.
Chloridium lignicola (F. Mangenot) W. Gams & Hol.-Jech
: Bisporomyces lignicola F. Mangenot, Revue Mycol.,
Paris 18: 136 (1953)
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
Notes: ITS and LSU sequence data are available.
Chloridium matsushimae W. Gams & Hol.-Jech
Distribution: China, Hong Kong, on submerged wood
(Tsui et al. 2000).
Sexual morph: Undetermined
Notes: Holotype MFC-1640. Sequence data is not
available.
Chloridium pachytrachelum W. Gams & Hol.-Jech
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
Notes: Holotype CBS 645.75. Sequence data is not
available.
Codinaea Maire, Publ. Inst. Bot. 3(4): 15 (1937)
Asexual morph Colonies on the natural substrate, scattered, hairy, black. Mycelium superficial or immersed,
composed of septate, branched, brown, smoothwalled
hyphae. Conidiophores macronematous, mononematous or
synnematous, brown to pale brown, septate. Conidiogenous
cells mono- to polyphialidic, with flared collarette, terminal, determinate or indeterminate with a few sympodial
extensions and sometimes with enteroblastic percurrent
regenerations. Conidia unicellular or septate, falcate to
lunate, hyaline, with a filiform appendage at each end.
Sexual morph Undetermined.
Type species: Codinaea aristata Maire, Publ. Inst. Bot.
3(4): 15 (1937)
Notes: Codinaea was introduced by Maire (1937) with
C. aristata Maire as the type species. The genus Dictyochaeta (Spegazzini 1923) was considered as an earlier
name for Codinaea (Maire 1937), but following the
molecular analyses of Réblová and Winka (2000), both
genera are accepted. Previous studies recommended that
species with filiform appendages (setulate conidia) can be
retained in Codinaea and taxa lacking appendages (asetulate conidia) can be placed in Dictyochaeta (Réblová and
Fungal Diversity (2019) 99:451–660
589
Fig. 55 Dictyochaeta cangshanensis (MFLU 18–1614, holotype). a Colonies on wood. b, c Conidiophores with conidia. d–f Conidiogenous
cells. g–i Conidia. k Germinating conidium. l, m Culture on PDA from above and reverse. Scale bars: b–c 30 lm, d–f 25 lm, g–k 15 lm
123
590
Winka 2000; Li et al. 2012; Oliveira et al. 2015a). However, some other studies also introduced some Dictyochaeta species with filiform appendages based on
molecular sequence data and morphology (Liu et al. 2016;
Wei et al. 2018).
Codinaea aquatica Castaneda et al.
Distribution: Brazil, Pernambuco, on submerged
decaying branches of unidentified plant in a River (Oliveira
et al. 2015a).
Sexual morph: Undetermined
Notes: Holotype URM 87707. Sequence data is not
available. This species is only known from the type
locality.
Codinaea yunnanensis Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555678, Facesoffungi
number: FoF 05464, Fig. 54
Etymology: Referring to the location where this fungus
was collected, Yunnan Province, China.
Holotype: MFLU 18–1611
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, hairy, greyish brown, shining, in
groups. Mycelium immersed, consisting of greyish brown,
branched, smooth hyphae. Conidiophores (62–)83–127 lm
long, 4.5–5.5 lm wide (
x = 105 9 5 lm, n = 20),
macronematous, mononematous, erect, unbranched, percurrently growing, straight or slightly flexuolls, smooth,
septate, brown, paler towards the apex, smooth. Conidiogenous cells initially monophialidic, with a conspicuous
collarette, later becoming poly-enteroblastic with sympodially conspicuous conidial primordia, integrated, terminal,
cylindrical, hyaline to subhyaline. Conidia 15–17 lm long,
2.5–3.5 lm wide (
x = 16 9 3 lm, n = 20), acrogenous,
aggregating in a globose mucoid mass, naviculate to long
fusiform, narrow at both of ends, with polar appendage,
aseptate, hyaline, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April
2015, Z.L. Luo, S-439 (MFLU 18–1611, holotype, HKAS
92839, isotype), ex-type living culture MFLUCC 17–0468.
Notes: Codinaea yunnanensis resembles C. pini in
having mononematous, unbranched, septate conidiophores,
terminal conidiogenous cells, aseptate, hyaline, setulate
conidia aggregating in a globose mucoid mass (Crous et al.
2014b). However, C. yunnanensis differs from C. pini by
its poly-enteroblastic conidiogenous cells which C. pini has
mono- to rarely polyphialidic conidiogenous cells, and C.
yunnanensis has larger conidia (15–17 vs. 13–15 lm).
Phylogenetic analysis also shows that Codinaea yunnanensis is distinct from C. pini (Fig. 51). In addition, Codinaea yunnanensis is phylogenetically close to
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Dictyochaeta fertilis (CBS 624.77), but their morphologies
are different (Menzies 1973).
Dictyochaeta Speg., Physis, Rev. Soc. Arg. Cienc. Nat. 7:
18 (1923)
Asexual morph Colonies effuse, brown to dark brown.
Mycelium mostly immersed, comprising pale to medium
brown, smooth, septate, branched hyphae. Setae fertile,
erect, septate, straight, thick-walled, dark brown, gradually
becoming paler towards the apex, smooth. Conidiophores
macronematous, mononematous, simple, erect, smooth,
straight to slightly flexuous, septate, pale brown to brown.
Conidiogenous cells monophialidic, terminal, integrated,
with a conspicuous collarete, smooth, pale brown to subhyaline. Conidia acicular with rounded bases, aseptate or
septate, aggregated in colourless slimy masses, hyaline,
smooth-walled. Sexual morph Undetermined.
Type species: Dictyochaeta fuegiana Speg., Physis,
Rev. Soc. Arg. Cienc. Nat. 7: 18 (1923)
Notes: Reblová and Winka (2000) revealed that setulate
and asetulate conidia of Dictyochaeta species grouped into
distinct sub groups in their phylogenetic analysis. Therefore, they suggested to maintain the genus name Dictyochaeta for species without setulae and Codinaea for
species with setulae. However, some researchers do not
agree with Reblová (2000), whereas some follow. More
fresh collections are needed to clarify the relationship and
distinction of these groups.
Dictyochaeta aciculata S.S. Silva & Gusmão
Distribution: Brazil, Caracol, on submerged petiole
(Silva and Gusmão 2013).
Sexual morph: Undetermined
Notes: Holotype HUEFS 192225. Sequence data is not
available. This species is only known from the type
locality.
Dictyochaeta aquatica W. Dong & H. Zhang
Distribution: Thailand, Prachuap Khiri Khan Province,
n submerged wood in a small River (Wei et al. 2018).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–2691. ITS and LSU
sequence data available. This species is only known from
the type locality.
Dictyochaeta assamica (Agnihothr.) Aramb. et al.
: Menisporella assamica Agnihothr.
Distribution: Malaysia, Pahang, Lepar Forest Reserve,
on decaying leaves of unidentified dicotyledon plant submerged in fast-flowing freshwater stream (Kuthubutheen
and Nawawi 1991b).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater: IMI
335369. Sequence data is not available. Kuthubutheen and
Nawawi (1991b) collected four samples for this species and
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Fig. 56 Dictyochaeta ellipsoidea (MFLU 18–1612, holotype). a, b Colonies on wood. c, d Conidiophores with conidia. e, f Conidiogenous cells.
g–i conidia. j Germinating conidium. Culture on PDA from above (k) and reverse (l). Scale bars: c–d 40 lm, e–f 25 lm, g–j 10 lm
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Fig. 57 Dictyochaeta lignicola (MFLU 18–1613, holotype). a, b Colonies on wood. c Conidiophores with conidia. d Conidiophores. e,
f Conidiogenous cells. g–j Conidia. k, l Culture on PDA from above and reverse. Scale bars: c, d 30 lm, e 20 lm, f–j 10 lm
one of them was collected from freshwater habitats (IMI
335369).
Dictyochaeta cangshanensis Z.L. Luo, K.D. Hyde & H.Y.
Su, sp. nov.
Index Fungorum number: IF 555679, Facesoffungi
number: FoF 05465, Fig. 55
123
Etymology: Referring to the location where the fungus
was collected, Cangshan Mountain, China.
Holotype: MFLU 18–1614
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on substrate effuse,
gregarious, brown, shining. Mycelium mostly immersed,
composed of branched, septate, smooth, thin-walled, brown
Fungal Diversity (2019) 99:451–660
hyphae. Setae 125–175 lm long, 4.5–6.5 lm wide
(
x = 150 9 5.5 lm, n = 20), erect, dark brown at the base,
paler towards the apex, 7–10-septate, acerose to subacerose, unbranched, smooth. Conidiophores 39–53 lm
long, 3.5–4.5 lm wide (
x = 46 9 4 lm, n = 20),
macronematous, mononematous, in groups from the
mycelial knots from the bases of setae, short, brown,
straight or slightly flexuous, 3–6-septate, unbranched,
cylindrical, smooth. Conidiogenous cells monophialidic,
with flared collarette, determinate, terminal, subhyaline.
Conidia
15–18 lm
long,
2.5–3.5 lm
wide
(
x = 16.5 9 3 lm, n = 20), acrogenous, solitary, aggregating in a globose mass at apex of conidiophore, aseptate,
cylindrical or long fusiform, curved, with hair-like appendages at both ends, hyaline, smooth-walled. Sexual morph
Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on submerged decaying wood in a
freshwater stream, July 2016, H.Y. Su, S-829 (MFLU
18–1614, holotype), ex-type living culture MFLUCC
17–2214.
Notes: Dictyochaeta cangshanensis resembles D.
aquatica in having septate, unbranched setae dark brown at
the base, paler towards the apex, mononematous, septate,
cylindrical, unbranched conidiophores with similar size,
aseptate, cylindrical or long fusiform, hyaline conidia with
hair-like appendages (Wei et al. 2018). However, phylogenetic analysis shows that D. cangshanensis is distinct
from D. aquatica. Dictyochaeta cangshanensis also
resembles D. siamensis in having septate, unbranched setae
dark brown at the base, paler towards the apex, mononematous, septate, cylindrical, unbranched conidiophores,
aseptate, cylindrical or long fusiform, hyaline conidia with
hair-like appendages and aggregating in a globose mass at
apex of conidiophore (Liu et al. 2016). However, D.
cangshanensis differs from D. siamensis in having shorter
setae (125–175 vs. 165–365 lm), shorter conidiophores
(39–53 vs. 60–100 lm) and monophialidic conidiogenous
cells while D. siamensis has monophialidic or rarely
polyphialidic conidiogenous cells.
Dictyochaeta coffeae (Maggi & Persiani) Aramb. &
Cabello
: Codinaea coffeae Maggi & Persiani
Distribution: China, Hong Kong, Lam Tsuen River, on
submerged wood (Tsui et al. 2001b).
Sexual morph: Undetermined
Notes: Holotype H.B.R 107 S. Sequence data is not
available.
Dictyochaeta curvispora Cai et al.
Distribution: Philippines, saprobic on submerged bamboo (Cai et al. 2004b).
Sexual morph: Undetermined
593
Notes: Holotype PDD 75040. Sequence data is not
available. This species is only known from the type
locality.
Dictyochaeta ellipsoidea Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555680, Facesoffungi
number: FoF 05466, Fig. 56
Etymology: Referring to ellipsoid conidia of this fungus.
Holotype: MFLU 18–1612
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, greyish brown, shining. Mycelium
partly immersed, composed of septate, hyaline hyphae.
Conidiophores 254–288 lm long, 8–10 lm wide
(
x = 271 9 9 lm, n = 20), macronematous, mononematous, septate, unbranched, erect, flexuous, dark brown
below, pale towards apex, smooth, percurrently prolieferating. Conidiogenous cells monophialidic, confined to
apical region on conidiophore, with conspicuous, flared,
funnel-shaped collarette. Conidia 13–15 lm long, 5.5–
6.5 lm wide (
x = 14 9 6 lm, n = 20), acrogenous, ellipsoid, solitary, aggregating in a globose mucoid mass,
curved, slightly pointed at both ends, with polar appendage,
aseptate, guttulate, often accumulating at tip of phialide,
hyaline, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-898 (MFLU 18–1612, holotype), ex-type
living culture MFLUCC 18–1574.
Notes: Dictyochaeta ellipsoidea mostly resembles Dictyochaeta tropicalis in having mononematous, septate,
unbranched conidiophores dark brown below, pale towards
apex, monophialidic, terminal conidiogenous cells with
funnel-shaped collarette and hyaline, aseptate, ellipsoid
conidia with one setulae at each end of conidium (Bhat and
Kendrick 1993). However, Dictyochaeta ellipsoidea differs
from D. tropicalis in having longer conidiophores
(254–288 vs. 90–130 lm) and guttulate, larger conidia
(13–15 9 5.5–6.5 vs. 7.5–9.5 9 3–5 lm). Dictyochaeta
ellipsoidea also shares similar morphology with D. multisetula in having mononematous, septate, unbranched
conidiophores dark brown at base, pale towards apex,
monophialidic, terminal conidiogenous cells with funnelshaped collarette and hyaline, aseptate, ellipsoid conidia
(Whitton et al. 2000). However, Dictyochaeta ellipsoidea
differs from D. multisetula in different sizes of conidiophores and conidia, the conidia of D. ellipsoidea with only
one setulae at each end, but the conidia of D. multisetula
are multi-setulate at each ends (Whitton et al. 2000).
Dictyochaeta fertilis (S. Hughes & W.B. Kendr.) Hol.-Jech
: Codinaea fertilis S. Hughes & W.B. Kendr.
Distribution: Malaysia, Kuala Lumpur, on decaying
leaves of unidentified dicotyledon plant submerged in fast-
123
594
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Fungal Diversity (2019) 99:451–660
b Fig. 58 Dictyochaeta
submersa (MFLU 18–2321, holotype)
a Colonies on wood. b Conidiophores with conidia. c Conidiophores
with setae. d Conidiogenous cells. e–h Conidia. i Germinating
conidium. j, k Culture on PDA from above and reverse. Scale bars: b,
c 50 lm, d, e 15 lm, f–i 5 lm
flowing freshwater stream (Kuthubutheen and Nawawi
1991b).
Sexual morph: Undetermined
Notes: Holotype PDD 20965, specimen collected from
freshwater habitats: IMI 335368. ITS sequence data is
available.
Dictyochaeta guadalcanalensis (Matsushima) Kuthubutheen & Nawawi
: Phialogeniculata guadalcanalensis Matsush.
Distribution: Malaysia, Bako National Park, on decaying twig of unidentified angiosperm submerged in fastflowing freshwater stream (Kuthubutheen and Nawawi
1991b).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater habitats:
IMI 343460; Sequence data is not available.
Dictyochaeta lignicola Z.L. Luo, H.Y. Su & K.D. Hyde,
sp. nov.
Index Fungorum number: IF 555681, Facesoffungi
number: FoF 05467, Fig. 57
Etymology: Referring to this fungus dwelling on wood.
Holotype: MFLU 18–1613
Saprobic on submerged decaying wood. Asexual
morph Colonies on natural substrate, effuse, superficial,
hairy, greyish brown, in groups. Mycelium immersed,
composed of branched septate, smooth haphae. Conidiophores
204–276 lm
long,
7.5–8.5 lm
wide
(
x = 240 9 8 lm, n = 20), macronematous, mononematous, solitary, cylindrical, erect, straight or slightly flexuous, unbranched, septate, brown, becoming pale brown
towards the apex, smooth. Conidiogenous cells mono- to
polyphialidic, determinate, terminal, subhyaline. Conidia
13–15 lm long, 4.5–5.5 lm wide (
x = 14 9 5 lm,
n = 20), solitary, aggregating in a globose mass at apex of
conidiophore, aseptate, navicular to fusiform, curved, with
appendages at both ends, hyaline, smooth-walled. Sexual
morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-899 (MFLU 18–1613, holotype), ex-type
living culture DLUCC 0899.
Notes: Dictyochaeta lignicola resembles D. renispora in
having mononematous, cylindrical, unbranched, septate
conidiophores, hyaline, aseptate, setulate conidia aggregating in a globose mucoid mass (Whitton et al. 2000).
However, D. lignicola differs from D. renispora in having
595
longer conidiophores (204–276 vs. 95–220 lm) and
polyphialidic conidiogenous cells, while D. renispora with
monophialidic conidiogenous cells and D. lignicola has
larger conidia (13–15 9 4.5–5.5 vs. 6–8.5 9 3–4.5 lm).
Dictyochaeta longispora (S. Hughes & W.B. Kendr.)
Whitton et al.
: Codinaea longispora S. Hughes & W.B. Kendr.
Distribution: Malaysia, Pahang, Lepar Forest Reserve,
on decaying leaves of unidentified dicotyledon plant submerged in fast-flowing freshwater stream (Kuthubutheen
and Nawawi 1991b).
Sexual morph: Undetermined
Notes: Holotype PDD 25755, specimen collected from
freshwater habitats: IMI 335366; Sequence data is not
available.
Dictyochaeta lunulospora (Hewings & J.L. Crane) Hol.Jech.
: Codinaea lunulospora Hewings & J.L. Crane
Distribution: Malaysia, Selangor, Mimaland, on
decaying twig of unidentified dicotyledon plant submerged
in fast-flowing freshwater stream (Kuthubutheen and
Nawawi 1991b).
Sexual morph: Undetermined
Notes: Holotype Dumont-VE 4619 (NY), specimen
collected from freshwater habitats: IMI 335363; Sequence
data is not available.
Dictyochaeta plovercovensis Goh & K.D. Hyde
Distribution: China, Hong Kong, Plover Cove Reservoir, on submerged wood (Goh and Hyde 1999), Yunnan
Province, Dianchi lake, on submerged wood (Luo et al.
2004); Philippines, Liput River, on submerged bamboo
(Cai et al. 2003b).
Sexual morph: Undetermined
Notes: Holotype IFRD 8744; Sequence data is not
available. Dictyochaeta plovercovensis was originally
collected from freshwater by Goh and Hyde (1999) in
Hong Kong, China. Cai et al. (2003a) reported this species
from Philippines. Luo et al. (2004) reported this species
from a freshwater lake in Yunnan Province, China. However, both the later studies did not provide descriptions and
illustrations for this species.
Dictyochaeta subfuscospora Kuthub. & Nawawi
Distribution: Australia, Queensland, on submerged
wood in a stream in Mountain Lewis (Hyde and Goh 1997).
Sexual morph: Undetermined
Notes: Holotype IMI 343458; Sequence data is not
available.
Dictyochaeta siamensis J. Yang, K.D. Hyde & J.K. Liu
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, Kaeng Krachan, on submerged wood (Liu et al.
2016).
Sexual morph: Undetermined
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Fungal Diversity (2019) 99:451–660
Fig. 59 Sporidesmium brachypus (MFLU 18–1615). a Colonies on wood. b–d Conidiophores with conidia. e Conidiogenous cells. f–h conidia
i Germinating conidium. j, k Culture on PDA from above and reverse. Scale bars: b–d 40 lm, e 10 lm, f–i 20 lm
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Fungal Diversity (2019) 99:451–660
Notes: Holotype MFLU 15–1149; ITS, LSU and TEF1a
sequence data are available. This species is only known
from the type locality.
Dictyochaeta submersa Z.L. Luo, K.D. Hyde & H.Y. Su,
sp. nov.
Index Fungorum number: IF 555682, Facesoffungi
number: FoF 05468, Fig. 58
Etymology: Referring to the submerged habitat of this
fungus.
Holotype: MFLU 18–2321
Saprobic on submerged decaying wood. Asexual
morph Colonies on the substratum superficial, effuse,
greyish brown, shining. Mycelium partly immersed, partly
superficial, consisting of branched, septate, brown, smooth
hyphae. Setae erect, cylindrical, straight, septate, unbranched, pale brown to brown. Conidiophores 62–122(–152)
lm long, 3–5 lm wide (
x = 92 9 4 lm, n = 20),
macronematous, mononematous, erect, solitary or in small
groups, unbranched, cylindrical, 4–6-septate, straight or
slightly flexuous, pale brown, smooth. Conidiogenous cells
monophialidic, terminal, determinate, subhyaline, narrowing below the collarette. Conidia 13.5–16.5 lm long, 2.5–
3.5 lm wide (
x = 15 9 3 lm, n = 20), acrogenous, solitary, aggregating in a globose mass at apex of conidiophore, fusiform, curved, rounded and narrow at both ends,
with polar appendages, aseptate, hyaline, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Jizu
Mountain, saprobic on decaying wood submerged in a
freshwater stream, March 2017, N. Zhao, S-1014 (MFLU
18–2321, holotype), ex-type living culture DLUCC 1014.
Notes: Dictyochaeta submersa resembles D. aquatica in
having mononematous, unbranched, cylindrical, septate
conidiophore, monophialidic, terminal conidiogenous cells
with conspicuous funnel-shaped collarettes and aseptate,
hyaline conidia with polar appendages (Wei et al. 2018).
However, D. submersa differs from D. aquatica in having
longer conidiophores (62–122(–152) vs. 47–68 lm) and
aseptate, fusiform conidia, while D. aquatica has 0–1septate, oblong to allantoid conidia. Phylogenetic analysis
also shows that D. submersa and D. aquatica are distinct
species (Fig. 51).
Sporidesmium brachypus (Ellis & Everh.) S. Hughes
: Helminthosporium brachypus Ellis & Everh.,Publications of the Field Columbia Museum, Bot. series 1: 92
(1896)
: Ellisembia brachypus (Ellis & Everh.) Subram.,
Proc. Indian natn Sci. Acad., Part B. Biol. Sci. 58(4): 183
(1992)
Facesoffungi number: FoF 05469, Fig. 59
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, scattered, dark
brown. Mycelium partly immersed, composed of septate,
597
branched, smooth, hyphae. Conidiophores 86–114 lm
long, 5–7 lm wide (
x = 100 9 6 lm, n = 20), macronematous, mononematous, solitary, erect, unbranched, 7–9septate, straight or flexuous, percurrently growing, dark
brown, smooth. Conidiogenous cells holoblastic,
monoblastic, integrated, terminal, dark brown. Conidia 45–
63 lm long, 13–17 lm wide (
x = 54 9 15 lm, n = 20),
acrogenous, solitary, ovoid to fusiform, 5–6-pseudoseptate,
truncate at base, with a short and hyaline rostrate tip at
apex,
brown,
smooth-walled.
Sexual
morph
Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-868 (MFLU 18–1615), living culture
MFLUCC 18–1573.
Notes: Specimens collected from freshwater habitats:
MFLU 18–1615. ITS, LSU, SSU, RPB2 and TEF1a
sequence data are available. Ellis (1971) described the
species Sporidesmium brachypus from twigs of various
trees. Shirouzu and Harada (2004) collected and described
this species as Ellisembia brachypus and provided
descriptions and illustrations. Shenoy et al. (2006) provided
LSU sequence data for this species (Ellisembia brachypus
HKUCC 10555). Su et al. (2016) treated Ellisembia as a
synonym of Sporidesmium based on the molecular data, we
follow this and use Sporidesmium brachypus in this study.
During an investigation on lignicolous freshwater fungi in
China, a Sporidesmium-like fungus was collected from
submerged wood in Yunnan Province, and morphological
characters of this fungus fit well with Sporidesmium
brachypus (Shirouzu and Harada 2004). Phylogenetic
analysis shows that our strain clusters with Ellisembia
brachypus (HKUCC 10555) with good support value. We
therefore identify our fungus as Sporidesmium brachypus
based on the morphology and molecular evidence and
provide descriptions and illustrations.
Menisporopsis S. Hughes, Mycol. Pap. 48: 59 (1952)
Asexual morph Colonies superficial, scattered, effuse,
white to pale brown. Mycelium partly immersed, composed
of brown hyphae. Setae central, solitary, erect, straight,
brown, septate, thick-walled, septate. Conidiophores synnematous, macronematous, septate, unbranched, cylindrical, lower part narrow, upper part wider, erect, straight or
slightly flexuous, brown, smooth, thin- to thick-walled.
Conidiogenous cells monophialidic, terminal, integrated,
pale brown, with collarette. Conidia acrogenous, fusiform,
gently curved or straight, with appendages at each end,
aseptate, hyaline, thin-walled, smooth. Sexual morph
Undetermined.
Type species: Menisporopsis theobromae S. Hughes,
Mycol. Pap. 48: 59 (1952)
123
598
Notes: The genus Menisporopsis was established by
Hughes (1952) with M. theobromae S. Hughes as the type
species. There are ten species accepted in the genus
Menisporopsis. Liu et al. (2016) provided the molecular
data for the type species Menisporopsis theobromae and it
is the first time provide sequence data for this genus. Two
species of Menisporopsis have been reported from freshwater habitats.
Menisporopsis multisetulata Tsui et al.
Distribution: China, Hong Kong, New Territories, Tai
Po, Lam Tsuen River, on submerged wood (Tsui et al.
1999).
Sexual morph: Undetermined
Notes: Holotype IFRD 8806. Sequence data is not
available. This species is known so far only from the type
locality.
Menisporopsis theobromae S. Hughes
Distribution: Thailand, Chiang Rai Province, stream
flowing near Tham Luang Nang Non Cave, on submerged
wood (Liu et al. 2016).
Sexual morph: Undetermined
Notes: Holotype IMI 39099a, specimen collected from
freshwater habitats: MFLU 15–1168. ITS and LSU
sequence data are available. Liu et al. (2016) collected this
species from freshwater in northern Thailand and provided
descriptions and illustrations.
Nawawia Marvanová, Trans. Br. mycol. Soc. 75(2): 227
(1980)
Asexual morph Colonies on the natural substratum effuse,
hairy, dark brown, with glistening mass of conidia at the tip
of conidiophores. Mycelium partly immersed and partly
superficial, composed of pale brown, septate hyphae.
Conidiophores cylindrical to slightly clavate, single or in
small groups, erect, straight, unbranched, smooth. Conidiogenous cells integrated, terminal, phialidic. Conidia with
a filiform appendage at each blunt corner, hyaline. Sexual
morph Undetermined.
Type species: Nawawia filiformis (Nawawi) Marvanová,
Trans. Br. mycol. Soc. 75(2): 227 (1980)
Notes: The asexual genus Nawawia was introduced by
Marvanová (1980) based on Clavatospora filiformis
Nawawi (Nawawi 1973), a hyphomycete found on submerged decaying twigs and petioles. Six species of
Nawawia have thus far been described, three from
Malaysia (Nawawi 1973; Kuthubutheen et al. 1992; Crous
et al. 2009; Goh et al. 2014), one from South Africa (Hyde
et al. 1996), one from Russia (Mel’nik and Hyde 2006) and
one from China (Peng et al. 2016). Presently, five Nawawia
species have been reported from freshwater habitats.
Nawawia dendroidea K.D. Hyde, Goh & Steinke
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Distribution: South Africa, Durban, Palmiet River, on
submerged pieces of stems of Phragmites (Hyde et al.
1996).
Sexual morph: Undetermined
Notes: Holotype BRIP 22870, Paratype BRIP 22871.
Sequence data is unavailable.
Nawawia filiformis (Nawawi) Marvanová
: Clavatospora filiformis Nawawi, Trans. Br. mycol.
Soc. 61(2): 390 (1973)
Distribution: Malaysia, on submerged wood (Nawawi
1973; Marvanová 1980; Kuthubutheen et al. 1992); Thailand, Trat Province, on decaying wood submerged in a
freshwater stream (Yang et al. 2018b)
Sexual morph: Undetermined
Notes: Holotype IMI 177451, other specimens collected
from freshwater habitats: MFLU 18–1500, MFLU
18–1501, HKAS 102155. ITS, LSU, SSU, RPB2 and
TEF1a sequence data are available.
Nawawia nitida Kuthub. et al.
Distribution: Malaysia, Pahang, Leper Forest Reserve,
on decaying wood submerged in a fast-flowing freshwater
stream (Kuthubutheen et al. 1992).
Sexual morph: Undetermined
Notes: Holotype IMI 344361. Sequence data is
unavailable.
Nawawia quadrisetulata Goh et al.
Distribution: Malaysia, Perak, Menglembu, Bukit Kledang, on submerged wood (Goh et al. 2014).
Sexual morph: Undetermined
Notes: Holotype UTAR(M)-0004. Sequence data is
unavailable.
Nawawia oviformis J. Peng & Z.F. Yu
Distribution: China, Sichuan Province, Gaosun County,
on submerged leaves of an unidentified dicotyledonous
plant in a stream (Peng et al. 2016).
Sexual morph: Undetermined
Notes: Holotype YMF 1.04361. Sequence data is
unavailable.
Phialogeniculata Matsush., Sci. Mus., Tokyo, N.S. 14: 471
(1971)
Asexual morph Colonies on natural substrate effuse,
brown. Conidiophores macronematous, mononematous,
solitary, erect, simple, straight or repeatedly geniculate,
septate, sometimes slightly constricted at the septa,
smooth-walled. Conidiogenous cells monoblastic or
polyphialidic, integrated, terminal. Conidia formed in
colourless slimy masses, obclavate, hyaline, smooth-walled. Sexual morph Undetermined.
Type species: Phialogeniculata guadalcanalensis Matsush., Bull. natn. Sci. Mus., Tokyo 14(3): 472 (1971)
Notes: The genus Phialogeniculata was introduced
Matsushima (1971) with P. guadalcanalensis as the type
Fungal Diversity (2019) 99:451–660
599
Fig. 60 Sporoschisma juvenile (MFLU 18–1608). a, b Colonies on wood. c, d Conidiophores with conidia. e, f Conidiogenous cells. g–
j Conidia. k Germinating conidium. Scale bars: c, d 50 lm, e–h 20 lm, i–k 15 lm
123
600
species. Two more species were described by Matsushima
(1993), namely Phialogeniculata dimorpha and P.
mulliseplata. Hyde et al. (1998a) introduced a freshwater
species in this genus named as Phialogeniculata africana
which was collected from freshwater in South Africa.
Phialogeniculata africana Goh et al.
Distribution: South Africa, Duban, Palmiet River, on
submerged wood (Hyde et al. 1998b).
Sexual morph: Undetermined
Notes: Holotype IFRD 8830. Sequence data is not
available. This species is only known from the type
locality.
Sporoschisma Berk. & Broome, Gard. Chron., London:
540 (1847)
Asexual morph Colonies on the substratum superficial,
effuse, gregarious, hairy. Mycelium immersed, composed
of pale to dark brown hyphae. Setae scattered, capitate,
with hyaline, mucilaginous substances at the swollen apex,
septate, smooth-walled. Conidiophores macronematous,
mononematous, smooth, black, erect, straight or slightly
flexuous, solitary, each composed of a bulbous base, a
cylindrical stipe and a swollen venter. Conidiogenous cells
monophialidic, integrated, terminal, determinate, brown.
Conidia solitary, in pseudo-chains, cylindrical, septate,
verruculose or smooth, developing endogenously in basipetal succession. Sexual morph Ascomata superficial,
solitary, scattered, anchored to substrate by an indistict
basal stroma, wall smooth and shiny. Setae stiff, erect,
unbranched, dark brown to blact, capitate, arising from all
over surface of ascomatal wall. Paraphyses filiform, septate, longer than asci, hyaline. Asci unitunicate, cylindrical
to narrowly clavate, apex with a refractive, non-amyloid
ring. Ascospores partially biseriate to biseriate, fusiform,
slightly curved, pale brown.
Type species: Sporoschisma mirabile Berk. & Broome,
London: 540 (footnote) (1847)
Notes: The genus Sporoschisma was introduced by
Berkeley (1847) with S. mirabile Berk. & Broome as the
type species. Sporoschisma is cosmopolitan distribution,
largely reported from moist terrestrial habitats in Asia,
Europe and Africa (Berkeley 1847; Rao and Rao 1964;
Bhat and Kendrick 1993; Goh et al. 1997; Sivichai et al.
2000b; Zelski et al. 2014; Yang et al. 2016a, b). Several
species are known from aquatic habitats (Goh et al. 1997;
Sivichai et al. 2000b; Zelski et al. 2014; Luo et al. 2016;
Yang et al. 2016a, b). There are 14 species accepted in
Sporoschisma.
Sporoschisma australiense (Goh & K.D. Hyde) Réblová
: Sporoschismopsis australiensis Goh & K.D. Hyde,
Mycol. Res. 101: 1302. 1997.
Distribution: Australia, On submerged wood in a
freshwater stream (Goh et al. 1997).
Sexual morph: Undetermined
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Notes: Holotype HKU (M) 2313 (now in IFRD).
Sequence data is not available. Goh et al. (1997) introduced
this species as Sporoschismopsis australiensis, Réblová
et al. (2014) transferred Sporoschismopsis australiensis to
the genus Sporoschisma as Sporoschisma australiense.
Goh et al. (1997) mentioned the holotype of this species as
HKU (M) 2313, the isotype of this species was deposited in
IMI herbarium.
Sporoschisma hemipsila (Berk. & Broome) Zelski et al.
: Sphaeria hemipsila Berk. & Broome, Bot. J. Linn.
Soc. 14: 126 (1873).
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in Nujiang River (Luo et al.
2016); Thailand, Nakorn Ratchassima Province/Prachuap
Khiri Khan Province: on submerged wood in freshwater
(Sivichai et al. 2000b; Yang et al. 2016b); Perú, Cusco, on
submerged, decomposing woody and herbaceous debris in
freshwater habitats (Zelski et al. 2014).
Sexual morph: see Goh et al. (1997)
Notes: Specimens collected from freshwater habitats:
HKAS 92767, HKAS 92843, MFLU 15–1150, MFLU
16–1324, PE0177–21a, PE0177–21b, PE0177–21c. ITS,
LSU and RPB2 sequence data are available.
Sporoschisma juvenile Boud.
: Sporoschisma aquaticum Luo et al.
Facesoffungi number: FoF 03440, Fig. 60
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, gregarious, hairy, shining, pale
brown to black. Mycelium mostly immersed, composed of
of septate, smooth, pale to dark brown hyphae. Setae in
groups mixed with conidiophores, erect, straight or flexuous, 0–3-septate, smooth, brown, paler towards the apex.
Conidiophores 211–233 lm long, 12–14 lm wide
(
x = 222 9 13 lm, n = 20), macronematous, mononematous, erect, solitary or in groups of 2–3, associated with 2–6
capitate setae, straight or slightly flexuous, smooth, dark
brown, each composed of a bulbous base, a cylindrical
stipe and a swollen venter with a long cylindrical neck.
Conidiogenous cells monophialidic, terminal, integrated,
pale brown. Conidia 25–31 lm long, 9–10 lm wide
(
x = 28 9 9.5 lm, n = 20), solitary, forming linear falsechains, cylindrical, 2–3-septate, guttulate at maturity,
hyaline to pale brown when young, brown at maturity,
smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-906 (MFLU 18–1608), living culture
MFLUCC 18–1348.
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in Nujiang River (Luo et al.
2016); Perú, Cusco, on submerged, decomposing woody
Fungal Diversity (2019) 99:451–660
601
Fig. 61 Tainosphaeria lunata (MFLU 18–1610, holotype). a, b Colonies on wood. c–e Conidiophores with conidia. f, g Conidiogenous cells. h–
k conidia l Germinating conidium. m, n Culture on PDA from above and reverse. Scale bars: c 50 lm, d–e 30 lm, f–g 10 lm, h–l 5 lm
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Fig. 62 Tainosphaeria obclavata (MFLU 18–1455, holotype). a, b Colonies on wood. c–d Conidiophores with conidia. f–i Conidiogenous cells
and conidia. j–l Conidia. n Germinating conidium. o, p Culture on PDA from above and reverse. Scale bars: c 30 lm, d–i 20 lm, j–n 10 lm
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and herbaceous debris in freshwater habitats (Zelski et al.
2014).
Sexual morph: Undetermined
Notes: Epitype E00817146, other specimens collected
from freshwater habitats: DLU 628, MFLU 18–1608. ITS,
LSU, SSU and TEF1a sequence data are available.
Sporoschisma longicatenatum J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province:
Hua Hin, stream outside Kaeng Krachan National Park, on
submerged wood (Yang et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 16–1325, isotype HKAS 95044.
ITS and LSU sequence data are available.
Sporoschisma mirabile Berk. & Broome
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in Dulong River (Luo et al.
2016).
Sexual morph: Undetermined
Notes: Specimen collected from freshwater: HKAS
92727. ITS and LSU sequence data are available.
Sporoschisma nigroseptatum D. Rao & P. Rag. Rao
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in a stream in Cangshan
Mountain (Luo et al. 2016); India, Western Ghats, Kali
River, submerged woody litter (Sudheep and Sridhar
2011).
Sexual morph: Undetermined
Notes: Holotype V.V.C.B.L. No. 355, other specimen
collected from freshwater habitats: DLU 629. Sequence
data is not available.
Sporoschisma palauense J. Yang & K.D. Hyde
Distribution: Thailand, Prachuap Khiri Khan Province:
Hua Hin, stream outside Kaeng Krachan National Park, on
submerged wood (Yang et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1151, isotype HKAS 95042.
ITS and LSU sequence data are available.
Sporoschisma phaeocentri Ho et al.
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in a stream in Cangshan
Mountain (Luo et al. 2016).
Sexual morph: Undetermined
Notes: Holotype HKU (M) 2897 (now in IFRD), other
specimen collected from freshwater habitats: HKAS
84039. Sequence data is not available.
Sporoschisma saccardoi E. W. Mason & S. Hughes
Distribution: Perú, Cusco, on submerged, decomposing
woody and herbaceous debris in freshwater habitats (Zelski
et al. 2014).
Sexual morph: see (Zelski et al. 2014).
Notes: Holotype IMI 25206, other specimen collected
from freshwater habitats: PE0349–1. Sequence data is not
available. Zelski et al. (2014) reported Sporoschisma
603
saccardoi from freshwater habitats and provided the
descriptions and illustrations for asexual and sexual morph
of this species.
Sporoschisma taitense (Mugambi & Huhndorf) A.N. Mill.
: Melanochaeta taitensis Mugambi & Huhndorf,
Sydowia 60: 263 (2008).
Distribution: China, Yunnan Province, saprobic on
decaying wood submerged in a stream in Cangshan
Mountain (Luo et al. 2016).
Sexual morph: see Mugambi and Huhndorf (2008)
Notes: Holotype EA, Mugambi 156 N, other specimen
collected from freshwater habitats: DLU 272. ITS and LSU
sequence data are available.
Sporoschisma uniseptatum Bhat & W.B. Kendr
Distribution: India, Western Ghats, Kali River, submerged woody litter (Sudheep and Sridhar 2011); Perú,
Cusco, on submerged, decomposing woody and herbaceous
debris in freshwater habitats (Zelski et al. 2014).
Sexual morph: see Sivichai et al. (2000b)
Notes: Holotype DAOM 214614, other specimen collected from freshwater habitats: PE0172-8. Sequence data
is not available.
Tainosphaeria F.A. Fernández & Huhndorf, Fungal Divers
18: 44 (2005)
Asexual morph Mycelium composed of partly immersed
and partly superficial, hyaline to pale brown, septate, erect
hyphae with glistening conidial masses at their apices.
Conidiophores superficial, macronematous, mononematous, erect, unbranched, bown to dark brown below half,
pale brown towards the apex, septate, smooth-walled.
Conidiogenous cells monophialidic, integrated, terminal,
determinate. Conidia fusiform or cylindrical, aseptate,
gently curved, rarely straight, hair-like appendages at both
ends, hyaline, smooth, thin-walled. Sexual morph Ascomata superficial on the substratum, subglobose to broadly
ovoid, dark brown, roughened surface, separate to gregarious, distinctly papillate. Ascomatal wall in surface view,
opaque in water, of textura angularis in lactophenol. Ascomatal apex broad, papillate, short. Paraphyses unbranched, septate, hyaline, tapering. Asci unitunicate,
cylindrical to clavate, short-stalked, thin-walled, apical
ring very small. Ascospores narrow-fusiform,, septate, with
rounded ends, often inequilateral, hyaline.
Type species: Tainosphaeria crassiparies F.A. Fernández & Huhndorf, Fungal Divers 18: 44 (2005)
Notes: The genus Tainosphaeria was introduced by
Fernández and Huhndorf (2005) based on the type species
T. crassiparies and both the asexual and sexual morphs
were found from the substrate and described. Liu et al.
(2016) described the second species in the genus Tainosphaeria based on a collection from freshwater habitats in
123
604
Thailand. Lu et al. (2016) introduced another species collected from freshwater in China.
Tainosphaeria jonesii Lu et al.
Distribution: China, Guangxi Province, on decaying
wood submerged in a freshwater stream (Lu et al. 2016).
Sexual morph: Undetermined
Notes: Holotype GZAAS 16–0065. ITS and LSU
sequence data are available.
Tainosphaeria siamensis Yang et al.
Distribution: Thailand, Prachuap Khiri Khan Province,
Hua Hin, Kaeng Krachan, on submerged wood (Liu et al.
2016).
Sexual morph: Undetermined
Notes: Holotype MFLU 15–1142. ITS and LSU
sequence data are available. This species is only known
from the type locality.
Tainosphaeria lunata Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555683, Facesoffungi
number: FoF 05470, Fig. 61
Etymology: Referring to the lunate conidia.
Holotype: MFLU 18–1610
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, greyish brown, in groups. Mycelium partly superficial, composed of septate, branched,
brown, smooth hyphae. Conidiophores 71–103 lm long,
6–8 lm wide (
x = 87 9 7 lm, n = 20), macronematous,
mononematous, aseptate or 1–2-septate, unbranched, erect,
flexuous, greyish black, pale towards apex, in groups,
arising from a common stroma. Conidiogenous cells
monophialidic, with conspicuous, flared collarette, terminal, determinate. Conidia 16–19 lm long, 4.5–5.5 lm
wide (
x = 17.5 9 5 lm, n = 20), acrogenous, fusiform,
solitary, guttulate, aseptate, curved, narrow and rounded at
both ends, without polar appendages, hyaline, smoothwalled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, October
2016, Z.L. Luo, S-871 (MFLU 18–1610, holotype), ex-type
living culture MFLUCC 18–0642.
Notes: Tainosphaeria lunata resembles T. jonesii and
T. siamensis in having macronematous, unbranched conidiophores, monophialidic, integrated, terminal conidiogenous cells and smooth, hyaline, aseptate conidia (Liu
et al. 2016; Lu et al. 2016). However, T. lunata differs from
T. jonesii in having wider conidiophores (6–8 vs.
2.5–3.5 lm) and conidia without polar appendages. T.
lunata differs from T. siamensis by its wider conidiophores
(6–8 vs. 2.5–4 lm) and fusiform, guttulate conidia without
polar appendages. Phylogenetic analysis shows that Tainosphaeria lunata is distinct from other Tainosphaeria
species.
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Tainosphaeria obclavata D.F. Bao, Z.L. Luo, K.D. Hyde
& H.Y. Su, sp. nov.
Index Fungorum number: IF 555684, Facesoffungi
number: FoF 05471, Fig. 62
Etymology: Referring to the obclavate conidia of this
fungus
Holotype: MFLU 18–1455
Saprobic on submerged decaying wood. Asexual
morph Colonies effuse, greyish brown, shining. Mycelium
mostly immersed, composed of septate, branched, brown,
smooth hyphae. Conidiophores 60–106 lm long, 4.5–
5.5 lm wide (
x = 83 9 5 lm, n = 20), macronematous,
mononematous, single or in groups, 3–6-septate, unbranched, erect, flexuous, greyish brown, paler towards the
apex, smooth. Conidiogenous cells monophialidic, with
conspicuous, flared, funnel-shaped collarette, light brown,
terminal, sometimes becoming intercalary, determinate,
sometimes sympodial, narrowing below the collarette.
Conidia 17–21 lm long, 5–6 lm wide (
x = 19 9 5.5 lm,
n = 20), solitary, straight, obclavate, gradually narrow
towards the apex, tapering at the apex, rounded at the base,
1-septate, with 2–3 guttules in each cell, hyaline, smoothwalled. Sexual morph Undetermined.
Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon, on
submerged decaying wood, October 2017, Z.L. Luo, B-49
(MFLU 18–1455, holotype), ex-type culture, MFLUCC
18–0260.
Notes: Tainosphaeria obclavata resembles T. jonesii, T.
siamensis and T. lunata in having macronematous,
unbranched conidiophores, monophialidic, integrated, terminal conidiogenous cells with conspicuous, flared, funnelshaped collarette and smooth, hyaline conidia (Liu et al.
2016; Lu et al. 2016). However, T. obclavata can easily be
distinguished from other Tainosphaeria species by having
obclavate, uniseptate, guttulate conidia which gradually
narrow towards the apex. Phylogenetic analysis also shows
that Tainosphaeria obclavata is distinct from other Tainosphaeria species (Fig. 51).
Thozetella Kuntze, Revis. gen. pl. (Leipzig) 2: 873 (1891)
Asexual morph Colonies cream and light brown, flat,
woolly to subfelty, margin incised and indistinct. Conidiomata sporodochial or effuse, superficial, sessile, forming
a convex or flat hymenium, topped by a moist spore mass.
Conidiophores macronematous, brown, irregularly cylindrical, branched, arising from a basal plate. Conidiogenous
cells monophialidic, integrated, determinate, terminal, light
brown, irregularly cylindrical, with no or minute collarette,
with periclinal wall thickening. Microawns produced from
conidiophores,sigmoid or L-shaped, aseptate, smooth,
apical part acerose, straight or slightly undulating, thickwalled, hyaline, aseptate. Conidia lunate, aseptate, finely
Fungal Diversity (2019) 99:451–660
guttulate, hyaline, smooth-walled. Sexual morph
Undetermined.
Type species: Thozetella nivea (Berk.) Kuntze, Revis.
gen. pl. (Leipzig) 2: 873 (1891)
Notes: Thozetella O. Kuntze (1891) is a nomen novum
for Thozetia Berk. et F. Mueller, which was antedated by
Thozetia F. Mueller ex Bentham (Asclepiadaceae).
Pirozynski and Hodges (1973) revised the genus and
accepted four species. Since then, a further 18 species were
introduced in this genus.
Thozetella coronata Monteiro et al.
Distribution: Brazil, Pará State, submerged decaying
plant materials in freshwater stream in Brazilian Amazon
forest (Monteiro et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype HUEFS 196471. Sequence data is not
available.
Thozetella nivea (Berk.) Kuntze
: Thozetia nivea Berk
Distribution: Thailand, Khao Yai National Park, Tad Ta
Phu, on submerged wood in a freshwater stream (Sivichai
et al. 2002).
Sexual morph: Undetermined
Notes: ITS and LSU sequence data are available. Sivichai et al. (2002) reported this species from freshwater in
Thailand, but did not provide descriptions and illustrations.
Thozetella ypsiloidea Monteiro et al.
Distribution: Brazil, Pará State, submerged decaying
plant materials in freshwater stream in Brazilian Amazon
forest (Monteiro et al. 2016b).
Sexual morph: Undetermined
Notes: Holotype HUEFS 196464. Sequence data is not
available.
Helminthosphaeriaceae Samuels, Cand. & Magni
Diplococcium Grove, J. Bot., Lond. 23: 167 (1885)
=Luxuriomyces R.F. Castañeda, Fungi Cubenses III: 23
(1988)
=Tretendophragmia Subram., Kavaka 20/21: 58 (1995)
Asexual morph Colonies effuse, dark, cottony or velvety.
Mycelium partly superficial, partly immersed. Conidiophores macronematous, mononematous, straight or flexuous, brown, smooth or verruculose, simple or branched.
Conidiogenous cells polytretic, integrated, terminal and
intercalary, determinate, cylindrical, brown. Conidia catenate, acropleurogenous, simple, short, clavate, ellipsoidal
or oblong rounded at the ends, 0–12-septate but mostly
1-septate, constricted at the septa, pale or dark brown,
smooth-walled. Sexual morph Undetermined.
Type species: Diplococcium spicatum Grove, J. Bot.,
Lond. 23: 167 (1885)
Notes: The genus Diplococcium was introduced by
Grove (1885) to accommodate D. spicatum Grove.
605
Presently, 28 species are accepted in the genus (Goh and
Hyde 1998; Wang and Sutton 1998; Hernández-Restrepo
et al. 2012; Ma et al. 2012; Silva et al. 2014; Almeida et al.
2015). The genus Diplococcium is morphologically similar
to Spadicoides by having mononematous, simple or branched conidiophores with polytretic, terminal or intercalary
conidiogenous cells producing acropleurogenous, euseptate
conidia (Ellis 1971; Seifert et al. 2011; Ma et al. 2012). The
sole distinguishing diagnostic character between these two
genera is that conidia are solitary in Spadicoides whereas
those of Diplococcium are catenate (Sinclair et al. 1985;
Ma et al. 2012).
Diplococcium aquaticum Goh et al.
Distribution: Australia, Queensland, Atherton Tablelands, Lake Barrine, on submerged wood (Goh et al.
1998b).
Sexual morph: Undetermined
Notes: Holotype HKU (M) 3201 (now in IFRD).
Sequence data is not available.
Endophragmiella B. Sutton, Mycol. Pap. 132: 58 (1973)
Asexual morph Colonies effuse to tufted, hairy to velutinous. Mycelium partly superficial, partly immersed. Conidiophores macronematous, mononematous, straight or
flexuous, branched, septate, pale brown to dark brown,
smooth. Conidiogenous cells monoblastic, integrated, terminal, sometimes determinate mostly percurrent, cylindrical. Conidia acrogenous, solitary, dry, simple, straight or
slightly curved, broadly fusiform, ellipsoidal, oblong
rounded at the ends or clavate, 1–3-septate, with a small,
protuberant thin-walled peg at the base, pale brown,
smooth or minutely verruculose. Sexual morph Phaeotrichosphaeria like and Lasiosphaeria like.
Type species: Endophragmiella pallescens B. Sutton,
Mycol. Pap. 132: 62 (1973)
Notes: The genus Endophragmiella was proposed by
Sutton (1973) for two species, E. pallescens B. Sutton and
E. canadensis (Ellis & Everh.) B. Sutton (= E. subolivacea
(Ellis & Everh.) S. Hughes). Hughes (1979) revised the
genera Endophragmia Duvernoy et Maire and Endophragmiella, some of the Endophragmia species were
transferred to Endophragmiella and an emended description of Endophragmiella was provided. The reported sexual morphs that linked to the genus Endophragmiella were
Phaeotrichosphaeria and Lasiosphaeria (Sivanesan 1983;
Hilber et al. 1987a). Selenosporella, a hyphomycetous
genus, was reported as the synanamorph of the Endophragmiella (Hughes 1979; Matsushima 1989).
Endophragmiella mexicana Mena et al.
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
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Fungal Diversity (2019) 99:451–660
Fig. 63 Cordana abramovii (MFLU 18–1454). a–e Conidiophores with conidia. f–g Conidiogenous cells and conidia. h–l Conidia.
m Germinating conidium. Scale bars: a–e 80 lm, f, g 30 lm, h 20 lm, i–m 10 lm
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607
Fig. 64 Cordana aquatica (MFLU 18–1625, holotype). a, b Colony on natural substrate. c, d Conidiophore with conidia. e Conidiogenous cells
with conidia. f–i Conidium. j Germinating conidium. Culture on PDA from above (k) and reverse (l). Scale bars: c–e 50 lm, f–i 5 lm
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b Fig. 65 Cordana lignicola (MFLU 18–1624, holotype). a, b Colony
on natural substrate. c Conidiophore with conidia. d, e Conidiogenous
cells with conidia. f–o Conidium. p Germinating conidium. Culture
on PDA from above (q) and reverse (r). Scale bars: c–e 50 lm,
f 25 lm, g–p 5 lm
Notes: Holotype XAL CB618. Sequence data is not
available. The species Endophragmiella mexicana was
introduced by Mercado et al. (1995). Ho et al. (2002b)
reported this species from freshwater habitats in China, but
did not provide descriptions and illustrations.
Endophragmiella occidentalis Castañeda et al.
Distribution: China, Hong Kong, Tai Po, Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
Notes: Holotype INIFAT C94/18. Sequence data is not
available.
Endophragmiella ramificata Hol.-Jech
Distribution: China, Hong Kong, Tai Po Kau Forest
Stream, on submerged wood (Ho et al. 2002a).
Sexual morph: Undetermined
Notes: Sequence data is not available.
Endophragmiella triseptata Tsui et al.
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Tsui et al. 2001).
Sexual morph: Undetermined
Notes: Holotype IFRD 8755. Sequence data is not
available.
Hilberina Huhndorf & A. N. Mill., Mycol. Res. 108(1): 31
(2004)
Asexual morph Undetermined. Sexual morph Descriptions refer to Miller and Huhndorf (2004).
Type species: Hilberina caudata (Fuckel) Huhndorf &
A.N. Mill., Mycol Res 108:31 (2004)
Notes: The genus Hilberina was introduced by Miller
and Huhndorf (2004) with H. caudata as type species.
There are 17 species accepted in this genus (Index Fungorum 2018) and only one species has been reported from
freshwater habitats (Ho et al. 2001; Luo et al. 2004).
Hilberina breviseta (P. Karst.) Huhndorf & A.N. Mill.
Distribution: Brunei, on submerged wood in freshwater
stream (Ho et al. 2001); China, Yunnan Province, on
submerged wood in Dianchi Lake (Luo et al. 2004).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Linocarpaceae Konta & K.D. Hyde
Linocarpon Syd. & P. Syd., Annls mycol. 15(3/4): 210
(1917)
Holomorph: Updated descriptions and illustrations refer to
Konta et al. (2017).
Type species: Linocarpon pandani Syd. & P. Syd.,
Annls mycol. 15(3/4): 210 (1917)
609
Notes: The genus Linocarpon was introduced by Sydow
and Sydow (1917), monographed by Hyde (1992d, 1997) with
23 accepted species. Konta et al. (2017) introduced a new
family Linocarpaceae with Linocarpon as the type genus
based on morphology and phylogenetic analyses. Four species
of Linocarpon have been reported from freshwater habitats.
Linocarpon apiculatum K.D. Hyde
Distribution: Indonesia, Irian Jaya, Manokwari, in
freshwater swamp on decaying palm petiole (Hyde 1997).
Asexual morph: Undetermined
Notes: Holotype IFRD 8785. Sequence data is not available. Hyde (1997) introduced this species from freshwater
habitats and is only known from the type locality.
Linocarpon appendisporum K.D. Hyde
Distribution: Indonesia, Irian Jaya, Manokwari, on dead
Pandanus leaves in freshwater swamp (Hyde 1997).
Asexual morph: Undetermined
Notes: Holotype IFRD 8786. Sequence data is not available. Hyde (1997) introduced this species from freshwater
habitats and is only known from the type locality.
Linocarpon bambusicola L. Cai & K.D. Hyde
Distribution: Philippines, saprobic on submerged bamboo (Cai et al. 2004b).
Asexual morph: Undetermined
Notes: Holotype PDD 75040. Sequence data is not available. Cai et al. (2004b) introduced this species from freshwater
habitats and is only known from the type locality.
Linocarpon pandanicola K.D. Hyde
Distribution: Indonesia, Irian Jaya, on decaying Pandanus leaves submerged in freshwater swamp (Hyde 1997).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 1120 (now in IFRD).
Sequence data is not available.
Coniochaetales Huhndorf et al.
Coniochaetaceae Malloch & Cain
Coniochaeta (Sacc.) Cooke, Grevillea 16(no. 77): 16
(1887)
Holomorph Descriptions and illustrations refer to
Maharachchikumbura et al. (2016).
Type species: Coniochaeta ligniaria (Grev.) Cooke,
Grevillea 16(no. 77): 16 (1887)
Notes: The genus Coniochaeta was introduced by Greville (1823–1824) with C. ligniaria as type species. Coniochaeta is an important ascomycete because its members
live in diversified habitats and nutritional modes. Most of
the species are reported from Europe and Eastern Asia
(Asgari et al. 2007; GBIF-accessed in 2018).
Coniochaeta gigantospora Fourn. et al.
Distribution: France, Vendée, Sauvaget, the River
Vendée, on submerged wood of Fraxinus excelsior (Raja
et al. 2012).
Asexual morph: Undetermined
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b Fig. 66 Cordana terrestris (MFLU 18–1623). a Colony on natural
substrate. b, c Conidiophore with conidia. d Conidiogenous cells with
conidia. e Conidiogenous cells. f–k Conidium. l Germinating conidium. Culture on PDA from above (m) and reverse (n). Scale bars: b,
c 50 lm, d, e 30 lm, f–k 5 lm
Notes: Holotype ILLS 60816. ITS and LSU sequence
data are available. This species is known only from the
type locality.
Coniochaeta leucoplaca (Sacc.) Cain
: Hypocopra leucoplaca Sacc., Syll. fung. (Abellini) 1:
244 (1882)
Distribution: USA, Illinois, Goose pond, on submerged
wood (Shearer and Crane 1986).
Asexual morph: Undetermined
Notes: Specimen collected from freshwater habitats:
ILLS 43771; LSU sequence data is available.
Coniochaeta ligniaria (Grev.) Cooke
: Sphaeria ligniaria Grev., Scott. crypt. fl. (Edinburgh)
2: 82 (1824)
Distribution: UK, England, on submerged wood in
cooling tower (Eaton 1972).
Asexual morph: Undetermined
Notes: Holotype Carmichael s.n.. ITS, LSU and RPB2
sequence data are available.
Coniochaeta renispora J.L. Crane & Shearer
Distribution: USA, Arkansas, on submerged wood in
freshwater (Crane and Shearer 1995).
Asexual morph: Undetermined
Notes: Holotype ILLS 51242. Sequence data is not
available.
Coniochaeta velutina (Fuckel) Cooke
: Rosellinia velutina Fuckel, Jb. nassau. Ver. Naturk.
23–24: 49 (1870)
Distribution: USA, Illinois, Goose pond, on submerged
wood (Shearer and Crane 1986).
Asexual morph: Undetermined
Notes: Specimen collected from freshwater habitats:
ILLS 37049. ITS, LSU, SSU and RPB2 sequence data are
available.
Coniochaeta kellermanii (Ellis & Everh.) Munk
: Rosellinia kellermanii Ellis & Everh., Proc. Acad.
nat. Sci. Philad. 42: 227 (1890)
Distribution: USA, Maryland and Illinois, Patuxent
River, on submerged wood (Shearer 1972; Shearer and
Crane 1986).
Asexual morph: Undetermined
Notes: Specimen collected from freshwater habitat:
ILLS 37699. Sequence data is not available.
Cordanales M. Hern.-Rest. & Crous
Cordanaceae Nann., Repert. mic. uomo: 498 (1934)
Cordana Preuss, Linnaea 24: 129 (1851)
611
Holomorph Descriptions and illustrations refer to Hernández-Restrepo et al. (2014) and Maharachchikumbura
et al. (2016).
Type species: Cordana pauciseptata Preuss, Linnaea 24:
129 (1851)
Notes: The genus Cordana was introduced by Preuss
(1851) for three species with C. pauciseptata as type species. Hernández-Restrepo et al. (2014) accepted 19 species
in this genus and provided a key for their identification.
Cordana species have been recorded from various temperate and tropical regions in the world, including Africa,
South America, South East Asia and New Zealand (Maharachchikumbura et al. 2016).
Cordana abramovii Seman & Davydk.
Facesoffungi number: FoF 05472, Fig. 63
Saprobic on submerged decaying wood. Asexual
morph colonies effuse, scattered, dark brown to black.
Mycelium partly immersed, composed of septate, branched,
dark brown, smooth hyphae. Conidiophores 348–490 lm
long, 7–11 lm wide (
x = 434 9 9 lm, n = 20), macronematous, mononematous, erect, straight or slightly flexuous,
unbranched, septate, singly or in small groups, brown, paler
towards the apex, smooth. Conidiogenous cells polyblastic,
terminal and intercalary, one swelling per cell, dark brown.
Conidia
20–24 lm
long,
11.5–13.5 lm
wide
(
x = 22 9 12.5 lm, n = 20), acropleurogenous, solitary,
tan to reddish brown, pyriform to obovate, truncated at the
base, rounded at the apex, 1-septate, mostly with one
guttlute in each cell, thick walled. Sexual morph
Undetermined.
Material examined: THAILAND, on submerged
decaying wood, October 2017, Z.L. Luo, B-98 (MFLU
18–1454), living culture MFLUCC 18–1605.
Distribution: Perú, Cusco, on submerged, decomposing
woody and herbaceous debris in freshwater habitats (Zelski
et al. 2014); Thailand, on submerged decaying wood (This
study).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater habitats:
ILL 41204, MFLU 18–1454. ITS, LSU, SSU, RPB2 and
TEF1a sequence data are available. Zelski et al. (2014)
redescribed Cordana abramovii based on the collections
from freshwater habitats in Perú and provided illustrations
and LSU sequence data. During an investigation of lignicolous freshwater fungi in Thailand, a Cordana-like fungus
was collected from southern Thailand. Morphologically,
our collection fits well with Cordana abramovii in having
macronematous, mononematous, erect, straight or slightly
flexuous, unbranched, septate conidiophores, polyblastic,
terminal and intercalary, conidiogenous cells with one
swelling per cell and tan to reddish brown, pyriform to
obovate, thick walled, 1-septate conidia with one guttlute
in each cell (Zelski et al. 2014). Phylogenetically, the
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Fungal Diversity (2019) 99:451–660
newly isolate clusters with other Cordana species in a wellsupported monophyletic clade and groups with Cordana
abramovii (PE0053–24a) with strong support (99% ML,
77% MP) (Fig. 67). Based on morphology and phylogeny,
we identify our isolate as Cordana abramovii and provide
the descriptions and illustration here. This is the first record
of this species in Thailand.
Cordana aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Fig. 67 Phylogram generated from maximum likelihood analysis
based on ITS sequence data for species of Cordana (with Coniochaeta velutina as outgroup). The best scoring RAxML tree with a
final likelihood value of - 3697.331198 is presented. RAxML
bootstrap support values equal to or greater than 75% are given
before the forward slash (black). Maximum parsimony bootstrap
support values equal to or greater than 75% are given after the
forward slash (red). Hyphen (‘-’) indicates a value lower than 75% for
RAxML and Maximum parsimony. Newly generated sequences are in
red. Ex-type strains are in bold
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Index Fungorum number: IF 555685, Facesoffungi
number: FoF 05473, Fig. 64
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1625
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, hairy, brown to dark brown. Mycelium
immersed, composed of septate, branched, smooth, subhyaline hyphae. Conidiophores 184–234 lm long, 5–7 lm
wide
(
x = 209 9 6 lm,
n = 20),
macronematous,
mononematous, solitary, erect, unbranched, septate,
straight or flexuous, with intercalary nodes, cylindrical,
dark brown at the base and lighter towards the apex.
Conidiogenous cells polyblastic, integrated, terminal,
becoming intercalary, percurrent, sympodial, denticulate,
usually inflated at the conidiogenous loci. Conidia 8–
10 lm long, 3–5 lm wide (
x = 9 9 4 lm, n = 20),
acropleurogenous, dry, mostly oblong, sometimes ellipsoid, 1-septate, slightly constricted at septate, rounded at
the apex, darker at septum, brown, smooth-walled. Sexual
morph Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on submerged decaying wood, May 2015,
Z.L. Luo, S-520 (MFLU 18–1625, holotype, HKAS 92702,
isotype), ex-type living culture MFLUCC 16–0954,
KUMCC 15–0293.
Notes: Cordana aquatica resembles C. verruculosa in
having macronematous, unbranched, smooth, straight or
flexuous conidiophores with intercalary nodes, polyblastic,
integrated, denticulate, terminal and intercalary conidiogenous cells usually inflated at the conidiogenous loci and
dry, solitary conidia (Hernández-Restrepo et al. 2014).
However, C. aquatica differs from C. verruculosa by its
septate, larger conidiophores (184–234 9 5–7 lm vs. up to
187 9 2.5–3 lm) and smooth, uniseptate, oblong, larger
conidia (8–10 9 3–5 lm vs. 3–5.5 9 2–3.5 lm), while C.
verruculosa has verruculose, aseptate, ellipsoid to obovoid
conidia. The phylogenetic analysis shows that Cordana
aquatica is close to C. lignicola, but their morphology is
different.
Cordana lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555686, Facesoffungi
number: FoF 05474, Fig. 65
Etymology: Referring to this fungus dwelling on wood
Holotype: MFLU 18–1624
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, hairy, brown to dark brown. Mycelium
immersed, composed of septate, branched, smooth,
613
subhyaline hyphae. Conidiophores 164–198 lm long, 4–
6 lm wide (
x = 181 9 5 lm, n = 20), macronematous,
mononematous, solitary, erect, unbranched, septate, straight
or flexuous, with intercalary nodes, cylindrical, dark brown
at the base and lighter towards the apex. Conidiogenous cells
polyblastic, integrated, terminal, becoming intercalary,
percurrent, sympodial, denticulate, usually inflated at the
conidiogenous loci. Conidia 9–11 lm long, 3.5–4.5 lm
wide (
x = 10 9 4 lm, n = 20), acropleurogenous, dry,
mostly oblong, sometimes ellipsoid, aseptate, 1–2-septate,
slightly constricted at septate, rounded at the apex, darker at
septum, hyaline when young, brown when mature, smoothwalled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in
a freshwater stream, April 2016, S.M. Tang, S-825
(MFLU 18–1624, holotype), ex-type living culture
MFLUCC 17–1332; Ibid., saprobic on decaying wood
submerged in a freshwater stream, April 2016, X.J. Su,
S-707 (MFLU 18–1626, Paratype), living culture
MFLUCC 17–1323.
Notes: Cordana lignicola resembles C. mercadiana in
having macronematous, unbranched, straight or flexuous
conidiophores with intercalary nodes, polyblastic, integrated, terminal, becoming intercalary, denticulate conidiogenous cells and dry, smooth, brown conidia slightly
constricted at septate (Hernández-Restrepo et al. 2014).
However, Cordana lignicola differs from C. mercadiana
by its septate, larger conidiophores (164–198 9 4–6 lm
vs. 57–150 9 2.5–3 lm) and 1–2-septate, oblong conidia,
while C. mercadiana have 0–1-septate, oblong, obovoid or
cylindrical conidia. Phylogenetic analysis shows that
Cordana lignicola is distinct from other Cordana species.
Cordana terrestris (Timonin) Hern.-Restr. et al.
: Spicularia terrestris Timonin, Can. J. Res. 18: 315
(1940)
: Pseudobotrytis terrestris (Timonin) Subram., Proc.
Indian Natn. Sci. Acad. Biol. Sci. 43:277, 1956
: Umbellula terrestris (Timonin) E.F. Morris,
Mycologia 47:603 (1955)
= Pseudobotrytis fusca Krzemien. & Badura, Acta Soc.
Bot. Pol. 23: 762 (1954)
= Porosphaerella borinquensis F.A. Fernández &
Huhndorf, Fungal Divers 17:12 (2004)
Facesoffungi number: FoF 05475, Fig. 66
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, hairy, pale brown. Mycelium immersed,
composed of septate, branched, smooth, subhyaline
hyphae. Conidiophores (90–)141–223 lm long, 5–7 lm
wide
(
x = 182 9 6 lm,
n = 20),
macronematous,
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Fungal Diversity (2019) 99:451–660
Fig. 68 Chaetomium globosum (S-164) a Colony on natural substrate. b–f Conidia. g Germinating conidia. Culture on PDA from above (h) and
reverse (i). Scale bars: b–g 15 lm
mononematous, solitary, erect, unbranched, septate,
straight or flexuous, cylindrical, dark brown at the base and
lighter towards the apex. Conidiogenous cells holoblastic,
polyblastic, terminal, discrete, hyaline, in groups of 8–14
arranged in umbels, with multiple conidiogenous loci
appearing as spinules, subhyaline. Conidia 6–8 lm long,
2.5–3.5 lm wide (
x = 7 9 3 lm, n = 20), acropleurogenous, mostly oblong, sometimes ellipsoid, guttulate,
uniseptate, slightly constricted at sptate, darker at septum,
light brown, smooth-walled. Sexual morph Undetermined.
123
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on submerged decaying wood, May 2015,
H.Y. Su, S-515 (MFLU 18–1623, ibid HKAS 92728),
living culture MFLUCC 16–0938, KUMCC 15–0308.
Distribution: China, Yunnan Province, Dulong River,
saprobic on decaying wood submerged in freshwater River
(This study).
Sexual morph: Undetermined
Notes: Specimens collected from freshwater habitat:
MFLU 18–1623, HKAS 92728. ITS, LSU, SSU, RPB2 and
Fungal Diversity (2019) 99:451–660
TEF1a sequence data are available. Hernández-Restrepo
et al. (2014) synonymised Spicularia terrestris, Pseudobotrytis fusca and Porosphaerella borinquensis as C.
terrestris. Our collection morphologically fits well with
asexual morph of Cordana terrestris in having macronematous, erect, unbranched, septate, straight or flexuous,
cylindrical conidiophores, terminal, discrete, hyaline
conidiogenous cells in groups of 8–14 arranged in umbels,
with multiple conidiogenous loci appearing as spinules and
light brown, uniseptate, guttulate conidia (HernándezRestrepo and Huhndorf 2004). The phylogenetic analysis
shows that our isolate (MFLUCC 16–0938) clusters with
Cordana terrestris with strongly support value (Fig. 67).
Based on the morphology and phylogeny, we identify our
fungus as Cordana terrestris. This is the first record of this
species in China and it is the first report collected from
freshwater habitats.
Cordana uniseptata Cai et al.
Distribution: China, Yunnan Province, Yiliang, on
Phyllostachys bambusoides submerged in a small forest
stream (Cai et al. 2004a).
Sexual morph: Undetermined
Notes: Holotype IFRD 8734. Sequence data is not
available.
615
large number of species is a result of naming species on the
basis of host association (Cannon 1988). Only one Phyllachora species has been reported from freshwater, namely
Phyllachora therophila growing on Juncus filiformis, collected in Austria (Magnes and Hafellner 1991).
Phyllachora therophila (Desm.) Arx & E. Müll.
: Sphaeria therophila Desm.
Distribution: Austria, Eastern Alps, on Juncus filiformis
in mountain lake (Magnes and Hafellner 1991).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Tamsiniella Wong et al., Can. J. Bot. 76(2): 334 (1998)
Asexual morph Undetermined. Sexual morph Ascomata
semi-immersed to immersed, dark brown, subglobose,
ostiolate, papillate, periphysate, solitary or gregarious.
Peridium comprising 1–2 layers of elongated angular,
brown cells, lined with a layer of very pale brown, disintegrated tissue. Paraphyses cellular, septate, sparse. Asci
8-spored, unitunicate, cylindrical, short pedicellate, apically truncate, with a nonamyloid, refractive, bifurcate
apical ring and an external thickening. Ascospores uniseriate to biseriate, ellipsoidal-fusiform, unicellular, surrounded by a narrow, roughened mucilaginous sheath,
hyaline.
Type species: Tamsiniella labiosa Wong et al., Can.
J. Bot. 76(2): 334 (1998)
Notes: The genus Tamsiniella was introduced by Wong
et al. (1998) with single species which was collected from
freshwater stream. There is only one species described in
this genus.
Tamsiniella labiosa Wong et al.
Distribution: Australia, north Queensland, Mount
Lewis, on wood submerged in a small stream (Wong et al.
1998b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8862. Sequence data is not
available. This species is only known from the type
locality.
Phyllachorales M.E. Barr
Phyllachoraceae Theiss. & H. Syd.
Ascovaginospora Fallah et al., Mycologia 89(5): 813
(1997)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Fallah et al. (1997).
Type species: Ascovaginospora stellipala Fallah et al.,
Mycologia 89(5): 813 (1997)
Notes: The genus Ascovaginospora was introduced by
Fallah et al. (1997) and typified by Ascovaginospora stellipala. There is only one species was accepted in this genus
and is only known from the type locality.
Ascovaginospora stellipala Fallah et al.
Distribution: USA, Northern Wisconsin, on submerged
dead stems and leaves of Carex limosa (Fallah et al. 1997). Sordariales Chad. ex D. Hawksw. & O.E. Erikss.
Asexual morph: Undetermined
Chaetomiaceae G. Winter
Notes: Holotype ILLS 52319, other specimens collected
Chaetomium Kunze, in Kunze & Schmidt, Mykologische
from freshwater habitats: ILLS 53005, ILLS 53006, ILLS
Hefte (Leipzig) 1: 15 (1817)
53007, ILLS 53009, ILLS 53010. LSU and SSU sequence
Holomorph
Descriptions
and
illustrations
see
data are available.
Maharachchikumbura et al. (2016).
Type species: Chaetomium globosum Kunze, in Kunze
Phyllachora Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
& Schmidt, Mykologische Hefte (Leipzig) 1: 16 (1817)
23–24: 216 (Fuckel 1870) [1869–70]
Notes: Chaetomium is a cosmopolitan genus with more
Holomorph Descriptions and illustrations refer to
than
150
species
(Asgari
and
Zare
2011;
Maharachchikumbura et al. (2016).
Maharachchikumbura et al. 2016). Chaetomium species
Type species: Phyllachora graminis (Pers.) Fuckel, Jb.
can be potential biological control agents (Soytong et al.
nassau. Ver. Naturk. 23–24: 216 (1870) [1869–70]
2001), they can produce bioactive metabolites (Wang et al.
Notes: Phyllachora is a large genus with more than 1500
species currently recognized (Index Fungorum 2018). This
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Fungal Diversity (2019) 99:451–660
2012; Li et al. 2014), or can produce mycotoxins and infect
skin and nails in humans (Pieckova 2003).
Chaetomium globosum Kunze
Facesoffungi number: FoF 02196, Fig. 68
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
effuse, dark brown to black. Conidiophores not observed.
Conidiogenous cell not observed. Conidia 12–14 lm long,
11–13 lm (
x = 13 9 12 lm, n = 20) wide, dry, globose to
subglobose, guttulate, dark brown to black, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on submerged decaying wood in a
freshwater stream, June 2014, Z.L. Luo, S-164 living culture KUMCC 15–0342; Ibid., saprobic on decaying wood
submerged in a freshwater stream, June 2014, Z. Li, S-166,
living culture MFLUCC 16–1102, KUMCC 15–0344.
Notes: Chaetomium globosum can produce bioactive
metabolites and is known as an endophytic fungus (Wang
et al. 2012; Li et al. 2014). During the investigation on
lignicolous freshwater fungi in China, a hyphomycetous
fungus was collected from submerged decaying wood in
freshwater streams. Phylogenetic analysis shows that our
isolate clusters with Chaetomium globosum (CBS 160.62)
with good support value (Fig. 1, clade 29). We therefore
identify our newly collections as asexual morph of Chaetomium globosum. It is the first report of this species collected from freshwater habitats.
Asexual morph Colonies effuse, velvety or cottony, pale
grey to browm or black. Mycelium superficial or immersed,
composed of hyaline to subhyaline, branched, septate,
smooth hyphae. Conidiophores micronematous, unranched,
hyaline, smooth, or differentiation inconspicuous. Conidiogenous cells monoblastic, terminal or lateral, hyaline to
pale brown, smooth. Conidia solitary, unicellular, mostly
spherical or subspherical, or sometimes pyriform, pale
brown to brown, smooth-walled. Sexual morph: Hyphae
hyaline, septate, branched, smooth-walled. Ascomata oval
to subglobose, ostiolate, brown to black-brown. Peridium
brown, textura irregularis. Rhizoids well developed, septate, pale brown. Ascomatal hair unbranched, tapering,
septate, yellow to brown, verrucose. Asci fasciculate, clavate, long-stalked. Ascospores limoniform to subglobose,
dark brown, thick-walled.
Type species: Humicola fuscoatra Traaen, Nytt Mag.
Natur. 52: 33 (1914)
Notes: Humicola was introduced by Traaen (1914) for
two species of fungi with hyaline hyphae, namely Humicola fuscoatra (type) Traaen and H. grisea Traaen. Only
one species has been reported from freshwater habitats
(Raja et al. 2009a, b).
Humicola asteroidea Udagawa & Y. Horie
Distribution: USA, Florida, on submerged wood in
freshwater (Raja et al. 2009b).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Corynascella Arx & Hodges, Stud. Mycol. 8: 23 (1975)
Asexual morph Undetermined. Sexual morph Ascomata
mostly superficial, scattered or clustered in small groups,
globose to subglobose, nonostiolate, dark brown to black.
Mycelium compose of hyaline to pale brown, branched,
septate hyphae. Peridium coriaceous, cephalothecoid,
brown to dark brown. Paraphyses hyaline, filiform, septate.
Asci 8-spored, clavate, rounded at the top, short-stipitate.
Ascospores broadly ellipsoidal, brown to dark brown,
smooth, thick-walled.
Type species: Corynascella humicola Arx & Hodges,
Stud. Mycol. 8: 23 (1975)
Notes: The genus Corynascella was introduced by von
Arx (1975) for a single species C. humicola. Four species
were accepted in this genus and only one species has been
reported from freshwater habitats.
Corynascella inquinata Udagawa & S. Ueda
Distribution: Japan, isolated from sewage sludge
(Udagawa and Ueda 1979).
Asexual morph: Undetermined
Notes: Holotype NHL 2841. ITS, RPB2 and TEF1a
sequence data are available.
Humicola Traaen, Nytt Mag. Natur. 52: 31 (1914)
Trichocladium Harz, Bull. Soc. Imp. nat. Moscou 44(1):
125 (1871)
Asexual morph Colonies effuse, at first hyaline and
becoming black. Mycelium composed of branched, septate,
light brown hyohae. Conidiophores micronematous,
mononematous, smooth, thin- or thick-walled, hyaline or
light brown. Conidiogenous cells holoblastic, integrated,
terminal or intercalary, smooth, cylindrical. Conidia solitary, subglobose to obpyriform, septate. Sexual morph
Undetermined.
Type species: Trichocladium asperum Harz, Bull. Soc.
Imp. nat. Moscou 44(1): 125 (1871)
Notes: Species of Trichocladium are frequently
encountered in aquatic environment (Crane and Shearer
1978; Kohlmeyer and Volkmann-Kohlmeyer 1995), and
five species have been reported from freshwater habitats.
Trichocladium englandense K.D. Hyde & Goh
Distribution: UK, Gloucestershire, River Coln, on submerged wood (Hyde and Goh 1999).
Sexual morph: Undetermined
Notes: Holotype IFRD 8867. Sequence data is not
available. This species is only known from the type
locality.
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617
Fig. 69 Cercophora caudata (HKAS 92795) a, b Conidiophores with conidia. c Conidiogenous cells with conidia. d–g Conidium.
h Germinating conidium. Culture on PDA from above (j) and reverse (k). Scale bars: a, b 150 lm, c 50 lm, d–h 20 lm
123
618
Trichocladium lignicola I. Schmidt
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Cai et al. 2002a); UK, England, River
Severn, on submerged wood (Kane et al. 2002).
Sexual morph: Undetermined
Notes: Holotype JE No. 238, paratype JE No. 227;
Sequence data is not available.
Trichocladium linderi J.L. Crane & Shearer
Distribution: USA, Maryland, on balsa wood submerged
in the Patuxent River (Crane and Shearer 1978).
Sexual morph: Undetermined
Notes: Holotype ILLS 37019, isotype NY-01042730.
Sequence data is not available.
Trichocladium lobatum Sutton
Distribution: USA, Illinois, from a Cypress Swamp, on
submerged wood (Crane and Shearer 1978).
Sexual morph: Undetermined
Notes: Holotype IMI 188290, specimen collected from
freshwater habitats: ILLS 37060. Sequence data is not
available.
Trichocladium moenitum J.L. Crane & Shearer
Distribution: USA, Illinois, from Elvira Cypress Swamp
and Wolf lake, on submerged wood (Crane and Shearer
1978).
Sexual morph: Undetermined
Notes: Holotype ILLS 36986, isotype NY-01042731,
other specimen collected from freshwater habitats: ILLS
37028. Sequence data is not available.
Lasiosphaeriaceae Nannf.
Apiosordaria Arx & W. Gams, Nova Hedwigia 13: 201
(1967)
Asexual morph Chrysosporium-like. Sexual morph Ascomata immersed to superficial, globose, subglobose to
pyriform, ostiolate or non-ostiolate, with or lacking
agglutinated hairs. Peridium membranaceous, brownishorange to brown; outer layer with textura intricata, inner
layer composed by thin-walled polygonal cells. Asci 4 or
8-spored, cylindrical to clavate, rounded at apex. Ascospores globose, subglobose or broadly fusiform, hyaline
when young, black when mature, two celled.
Type species: Apiosordaria verruculosa (C.N. Jensen)
Arx & W. Gams, Nova Hedwigia 13: 201 (1967)
Notes: Arx & Gams (1967) introduced the genus
Apiosordaria Arx & W. Gams (Lasiosphaeriaceae, Sordariales) based on the type species Pleurage verruculosa
C.N. Jensen [: A. verruculosa]. Krug et al. (1983) synonymised Echinopodospora and Lacunospora under
Apiosordaria based on morphological similarities.
Apiosordaria currently comprises 23 species with a
worldwide distribution in soil, dung or decaying wood
(Warcup 1951a, b; Stchigel et al. 2000). Some species have
also been found associated with plant material (Rostrup
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Fungal Diversity (2019) 99:451–660
et al. 1916) and the insect Plecia nearctica (Kish et al.
1974). Wu et al. (2016) introduced a new species
Apiosordaria hamata which was collected from sediment
in a freshwater lake.
Apiosordaria hamata Wu et al.
Distribution: China, Hubei Province, Wuhan City, isolated from sediment in Donghu Lake (Wu et al. 2016).
Asexual morph: Undetermined
Notes: Holotype HMAS 246231. ITS and LSU sequence
data are available.
Cercophora Fuckel, Jb. nassau. Ver. Naturk. 23–24: 244
(1870) [1869-70]
Asexual morph Obtained from culture. Hyphae thin-walled, hyaline to pale brown. Conidiogenous cells
monophialidic or polyphialidic, polytretic commonly produced from pale brown hyphae as single terminal or several
lateral phialides, delimited by a basal septum, monophialidic or polyphialidic, cylindrical to obpyriform, mostly
pale brown, constricted below the collarette, collarette
short, slightly flaring, inconspicuous, same colour as
phialide. Conidia subglobose to pyriform, truncate at base,
hyaline. Sexual morph Ascomata superficial, perithecial
ovoid or subglobose to broadly obpyriform, with a conical
to hemispherical neck, clustered in small groups, at times
in contact, dark brown to black. Ascomatal wall of textura
angularis in surface view, cells thickwalled, pale to dark
brown. Paraphyses filiform, hyaline, thin-walled, abundant, septate, unbranched. Asci 8-spored, unitunicate,
cylindrical, bi- to triseriate, spore-bearing part fusoidventricose, apex rounded, long-stipitate. Ascospores
cylindrical, ends rounded, straight to slightly sigmoid,
geniculate in lower quarter, aseptate, densely guttulate.
ascospore becoming differentiated into an apical swollen
head and a basal tail while inside the ascus; head ellipsoid,
hyaline, rarely pigmented, pale brown; tail septate, hyaline,
rarely pigmented, pale brown.
Type species: Cercophora mirabilis Fuckel, Jb. nassau.
Ver. Naturk. 23–24: 245 (1870)
Notes: The genus Cercophora Fuckel was established by
Fuckel (1870) for C. conica, C. fimiseda and C. mirabilis.
Fuckel (1873) synonymized Cercophora under Sordaria.
Lundqvist (1972) revised the Sordariaceae s. lat. and resurrected the genus Cercophora. The genus is characterized
by membraneous to coriaceous, dark colored, large ascomata and hyaline, cylindrical ascospores which develop a
swollen, pigmented head (Miller and Huhndorf 2001).
Currently, more than 50 species were accepted and most of
the species are lignicolous and coprophilous. Phylogenetic
studies inferred from LSU or multi-gene sequence data
indicated that Cercophora species are polyphyletic (Miller
and Huhndorf 2001; Chang et al. 2010). Asexual morph of
Cercophora species are cladorrhinum- or phialophora-like
Fungal Diversity (2019) 99:451–660
(Miller and Huhndorf 2001; Cai et al. 2006c; Doveri 2016).
Key to Cercophora species of the complex-sordarioides
and key to Cercophora species with a carbonaceous
peridium were provided by Doveri (2016), key to species
of Cercophora from China (Taiwan) was provided by
Chang and Wang (2005).
Cercophora appalachianensis O. Hilber & R. Hilber
Distribution: China, Hong Kong, Tai Po Kau Forest
stream, on submerged wood (Ho et al. 2001).
Asexual morph: Undetermined
Notes: Hilber et al. (1987b) mentioned the holotype of
Cercophora appalachianensis was deposited in BPI, isotype was deposited in NY, but did not give the herbarium
number. ITS and LSU sequence data are available. Ho
et al. (2001) reported this species from freshwater in Hong
Kong, China, but did not provide descriptions and
illustrations.
Cercophora aquatica Chaudhary et al.
Distribution: France, on submerged wood of Quercus
(Chaudhary et al. 2007).
Sexual morph: Phialophora-like, see Chaudhary et al.
(2007)
Notes: Holotype ILLS 58443. ITS, LSU and MCM7
sequence data are available.
Cercophora caudata (Sacc.) N. Lundq.
: Sphaeria caudata Curr., Trans. Linn. Soc. London
22: 320 (1859)
: Sordaria caudata Sacc., Syll. fung. (Abellini) 1: 236
(1882)
Facesoffungi number: FoF 05476, Fig. 69
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies effuse, hairy, dark
brown to black. Mycelium immersed, composed of septate,
branched, smooth, brown hyphae. Conidiophores 893–
1123 lm long, 12–22 lm wide (
x = 1008 9 17 lm,
n = 20), macronematous, mononematous, solitary, erect,
unbranched, septate, straight or flexuous, cylindrical, dark
brown to black, smooth. Conidiogenous cells polytretic,
integrated, terminal, brown. Conidia 18.5–21.5 lm long,
11–13 lm wide (
x = 20 9 12 lm, n = 20), acrogenous,
ellipsoidal or obovoid, papilate at the apex, truncate at the
base, guttulate, brown to dark brown, smooth-walled.
Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Nujiang
River, saprobic on submerged decaying wood, May 2015,
Z.L. Luo, S-779, living culture MFLUCC 17–0475; Ibid.,
saprobic on submerged decaying wood, May 2015, S.M.
Tang, S-346 (HKAS 92795), living culture MFLUCC
16–1271.
Distribution: China, Yunnan Province, on submerged
wood (This study).
Sexual morph: see Lundqvist (1972)
619
Notes: Holotype K(M) 68753, specimen collected from
freshwater habitats: HKAS 92795. ITS, LSU, SSU, RPB2,
TEF1a and b-tubulin sequence data are available.
Cercophora costariensis (Carroll & Munk) O. Hilber & R.
Hilber
: Bombardia costaricensis G.C. Carroll & Munk,
Mycologia 56(1): 80 (1964)
Distribution: China, Hong Kong, on submerged wood in
a freshwater stream (Tsui et al. 2000).
Asexual morph: Undetermined
Notes: Sequence data is not available.
Cercophora squamulosa Miller et al.
Distribution: France, Ariège, Le Pujol stream and
Peyrau brook, on submerged wood (Crous et al. 2016a).
Asexual morph: Phialophora-like, see Crous et al. (2016a)
Notes: Holotype ILLS 79803, other specimen collected
from freshwater habitats: ILLS 79954. ITS and LSU
sequence data are available.
Cercophora vinosa A.N. Mill. & J. Fourn
Distribution: France, Ariège, Artillac stream, on partly
submerged wood (Crous et al. 2016b).
Asexual morph: from culture, see Crous et al. (2016b)
Notes: Holotype ILLS 79802. ITS, LSU and b-tubulin
sequence data are available.
Cladorrhinum Sacc. & Marchal, in Marchal, Bull. Soc.
R. Bot. Belg. 24(1): 64 (1885)
Holomorph Descriptions refer to Mouchacca and Gams
(1993).
Type species: Cladorrhinum foecundissimum Sacc. &
Marchal, Bull. Soc. R. Bot. Belg. 24(1): 64 (1885)
Notes: The genus Cladorrhinum, typified by C. foecundissimum, was established by Marchal (1885) based on
a fungus isolated from dung. Cladorrhinum species are
found on dung and plant material and in soil.
Cladorrhinum samala (Subram. & Lodha) W. Gams &
Mouch
Distribution: Brunei, on submerged wood (Ho et al.
2001).
Sexual morph: Undetermined
Notes: Holotype IMI 113129. ITS and LSU sequence
data are available. Ho et al. (2001) reported this species
from freshwater habitats in Brunei, but did not provide
description and illustration.
Echria (N. Lundq.) Kruys et al., Fungal Divers 70: 106
(2015)
Asexual morph Undetermined. Sexual morph Ascomata
superficial to partially immersed, subiculate or not,
tomentose, scattered, obpyriform, ostiolate, membranous.
Neck cylindrical to conical, opaque, black, inside lined
with hyaline, filiform periphyses, covered with pointed,
unbranched, tapering, long tufts of agglutinated, rigid,
septate, cylindrical, brown hairs measuring. Peridium
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620
pseudoparenchymatous in surface view, semitransparent;
in longitudinal section, inner layer composed of elongated,
flattened, thin-walled, hyaline to brown cells; outer layer
composed of brown, thin-walled, angular cells. Paraphyses
hyaline, filiform-ventricose, septate, abundant. Asci
8-spored, unitunicate, thin-walled, broadly clavate, with a
long stipe, swelling in water. Ascospores irregularly
arranged, one-celled, hyaline when young, ranging from
golden to dark brown at maturity.
Type species: Kruys et al. (2015) raised the Arnium
section Echria to genus level but did not designated the
type species for this genus.
Notes: Kruys et al. (2015) raised the Arnium section
Echria to genus level and accepted two species, E.
gigantospora (Raja and Shearer) Kruys, Huhndorf and
A.N. Mill. and E. macrotheca (P. Crouan and H. Crouan)
Kruys, Huhndorf and A.N. Mill., based on morphology and
molecular data. The genus Echria is characterized by
having ascomata covered with long pointed tufts of
agglutinated rigid hairs and striated gelatinous sheaths that
swell in water and surround their ascospores (Kruys et al.
2015).
Echria gigantospora (Raja & Shearer) Kruys et al.
Distribution: USA, Ocala National Forest, Lake Kerr, on
submerged, decorticated wood (Kruys et al. 2015).
Asexual morph: Undetermined
Notes: Holotype ILL, slide made from a specimen on
submerged, decorticated wood, F77-1. ITS, LSU and btubulin sequence data are available.
Immersiella A. N. Mill. & Huhndorf, Mycol. Res. 108(1):
31 (2004)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations refers to Miller and Huhndorf (2004).
Type species: Immersiella immersa (P. Karst.) A.N.
Mill. & Huhndorf, Mycol. Res. 108(1): 31 (2004)
Notes: The genus Immersiella was introduced by Miller
and Huhndorf (2004) with two species and Immersiella
immersa as type. Immersiella is distinguished from Lasiosphaeria and Lasiosphaeris in having immersed to
erumpent ascomata covered with long, flexuous, brown
hairs (Hilber & Hilber 1983). Lasiosphaeria and Lasiosphaeris have superficial ascomata covered with either
tomentum or setae respectively. Immersiella can also be
distinguished from Lasiosphaeria in having hyaline centrum contents.
Immersiella immersa (P. Karst.) A.N. Mill. & Huhndorf
Distribution: China, Hong Kong, on submerged wood in
freshwater stream (Tsui et al. 2000).
Asexual morph: Undetermined
Notes: LSU sequence data is available.
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Zopfiella G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig)
1.2: 56 (1884)
Asexual morph Undetermined. Sexual morph Ascomata
solitary to gregarious, superficial or immersed in substrate,
covered with septate, flexuous, subhyaline hairs, cleistothecial, globose to subglobose, light brown. Peridium
pseudoparenchymatous, membranous, semi-transparent,
light brown. Asci 8-spored, cylindrical to clavate, apex
rounded, stipitate, evanescent. Ascospores triangular, sides
often concave, comers rounded, one truncate and bearing a
refractile scar at the former point of attachment of basal
cell, one bearing a germ pore, and one broadly rounded,
dark brown, thick-walled.
Type species: Zopfiella tabulata (Zopf) G. Winter,
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 56 (1884)
Notes: The genus Zopfiella was established by Winter
(1884) for Z. curvata and Z. tabulata. Lundqvist (1972)
revised the Sordariaceae s. lat. and the genus was accepted
within the family Lasiosphaeriaceae. Based on the phylogenetic analysis by using LSU sequence data, Huhndorf
et al. (2004) revised the family Lasiosphaeriaceae and
other families in the Sordariales, the genus Zopfiella was
transferred from Lasiosphaeriaceae to Chaetomiaceae since
the only one used Zopfiella species (Z. ebriosa) in their
study. To investigate the phylogenetic relationships of
Zopfiella and allied genera in the Sordariales, phylogenetic
analyses based on partial ITS, LSU and partial b-tubulin
sequences were carried out by Cai et al. (2006) and their
results showed that Zopfiella is polyphyletic, familial
replacement of Zopfiella in the family Chaetomiaceae was
not accepted. The most recent classification placed the
genus Zopfiella within the family Lasiosphaeriaceae
(Maharachchikumbura et al. 2016; Hongsanan et al. 2017;
Wijayawardene et al. 2017, 2018).
Zopfiella cephalothecoidea Guarro et al.
Distribution: Iraq, Euphrates, On submerged dead twig
(Guarro et al. 1996).
Asexual morph: Undetermined
Notes: Holotype FMR 5307, Isotype in IMI and BSRA
(10412). Sequence data are not available.
Zopfiella latipes (N. Lundq.) Malloch & Cain
: Tripterospora latipes N. Lundq., Bot. Notiser 122:
592 (1969)
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Cai et al. 2002a); Egypt, Nile River, on
submerged wood (Abdel-Aziz 2016).
Asexual morph: Undetermined
Notes: ITS, LSU and b-tubulin sequence data are
available. Zopfiella latipes is a fairly common species
found on various herbaceous and woody substrates in both
marine and terrestrial environments (Guarro et al.1991).
Fungal Diversity (2019) 99:451–660
621
Fig. 70 Cuspidatispora xiphiago (MFLU 18–1622). a Colony on substrate. b Conidiophores with conidia. c Conidiogenous cells with conidia.
d–m Conidia. n Germinating conidium. Culture on PDA from above (o) and reverse (p). Scale bars: b 140 lm, c 95 lm, d 15 lm, e–n 10 lm
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622
Cai et al. (2002a) reported this species from freshwater
lake in China, and provided photos for asci and ascospores
of this species but did not mention the information of
specimen.
Zopfiella leucotricha (Speg.) Malloch & Cain
: Sordaria leucotricha Speg., Michelia 1(no. 2): 225
(1878)
Distribution: Egypt, Nile River, on submerged wood
(Abdel-Aziz 2016).
Asexual morph: Undetermined
Notes: Sequence data is not available. Abdel-Aziz
(2016) reported this species from Nile River, but did not
provide descriptions and illustration for it.
Zopfiella lundqvistii Shearer & J.L. Crane
Distribution: USA, Illinois, Elvira Cypress Swamp, on
submerged wood (Shearer & Crane 1978).
Asexual morph: Undetermined
Notes: Holotype ILLS 36931, other specimens collected
from freshwater habitats: ILLS 36933, ILLS 36934; ITS
and LSU sequence data are available.
Zopfiella submersa Guarro, Al-Saadoon, Gené & Abdullah
Distribution: Iraq, Euphrates River, on submerged dead
culms of Phragmites sp. (Guarro et al. 1997).
Asexual morph: Undetermined
Notes: Holotype IMI 370952, isotype FMR 5646, other
specimen collected from freshwater habitats: FMR 5647.
Sequence data is not available.
Sordariales genera incertae sedis
Ascolacicola Ranghoo & K.D. Hyde, Mycologia 90(6):
1055 (1998)
Asexual morph Vegetative hyphae partly superficial and
partly immersed, branched and septate, not constricted at
the septa, hyaline. Conidiogenous cells thin-walled, hyaline, sometimes thickened and pigmented near the base,
subglobose, ampulliform, or irregularly cylindrical, bearing
a single terminal conidium. Conidia broadly obovoid,
l-septate near the base, slightly constricted at the septa
when mature, dark-brown, thick-walled. Sexual morph
Ascomata superficial, subglobose, black, coriaceous, solitary to gregarious and beaked. Peridium comprisirlg several layers of brown flattened angular cells. Paraphyses
filamentous, hyaline, numerous, flexuous, septate, rounded
at the ends. Asci 8-spored, unitunicate, cylindrical, short
pedicellate, apically rounded, with a J-, refractive, discoid,
apical ring. Ascospores overlapping uniseriate, ellipsoidal,
celled, not constricted at the septa, brown with hyaline to
pale brown end cells, smooth-walled.
Type species: Ascolacicola aquatica Ranghoo & K.D.
Hyde, Mycologia 90(6): 1056 (1998)
Notes: Ranghoo and Hyde (1998) introduced the genus
Ascolacicola Ranghoo & K.D. Hyde with A. aquatica
Ranghoo & K.D. Hyde as type species which was collected
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Fungal Diversity (2019) 99:451–660
from freshwater habitats. Up to now, there is one species
accepted in this genus and is only known from the type
locality.
Ascolacicola aquatica Ranghoo & K.D. Hyde
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
River, on submerged wood (Ranghoo and Hyde 1998).
Asexual morph: Trichocladium uniseptatum (Ranghoo
and Hyde 1998).
Notes: Holotype IFRD 8693. LSU sequence data is
available.
Cuspidatispora Shearer & Bartolata, Mycoscience 47(4):
220 (2006)
Asexual morph Mycelium immersed, composed of septate,
branched, smooth hyphae. Conidiophores macronematous,
mononematous, solitary, erect, unbranched, septate,
straight or flexuous, cylindrical, subhyaline to pale brown,
smooth. Conidiogenous cells holoblastic, polyblastic,
integrated, terminal, subhyaline. Conidia acrogenous,
obpyriform, papillate, guttulate, brown to dark brown,
smooth-walled. Sexual morph Descriptions and illustrations see Miller et al. (2006).
Type species: Cuspidatispora xiphiago Shearer & Bartolata, Mycoscience 47(4): 220 (2006)
Notes: The genus Cuspidatispora was introduced by
Miller et al. (2006) and typified by C. xiphiago which was
collected from submerged in a creek.
Cuspidatispora xiphiago Shearer & Bartolata
Facesoffungi number: FoF 05477, Fig. 70
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on natural substrate
superficial, effuse, brown. Mycelium immersed, composed
of septate, branched, smooth hyphae. Conidiophores 250–
300 lm long, 2–4 lm wide (
x = 275 9 3 lm, n = 20),
macronematous, mononematous, solitary, erect, unbranched, septate, straight or flexuous, cylindrical, subhyaline
to pale brown, smooth. Conidiogenous cells holoblastic,
polyblastic, integrated, terminal, subhyaline. Conidia 10–
12 lm long, 4–6 lm wide (
x = 11 9 5 lm, n = 20),
acrogenous, obpyriform, papillate, guttulate, brown to dark
brown, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Dulong
River, saprobic on submerged decaying wood, May 2015,
H.Y. Su, S-505 (MFLU 18–1622, HKAS 90735), living
culture MFLUCC 16–1277, KUMCC 15–0312.
Distribution: China, Yunnan Province, Dulong River, on
submerged wood (This study); USA, Illinois, Vermilion
County, Jordan Creek, on submerged wood (Miller et al. 2006).
Sexual morph: see Miller et al. (2006)
Notes: Holotype ILL, Shearer A184-1, isotype ILLS
57515, other specimens collected from freshwater habitats:
MFLU 18–1622, HKAS 90735. ITS, LSU, SSU, RPB2,
TEF1a and b-tubulin sequence data are available.
Fungal Diversity (2019) 99:451–660
623
Fig. 71 Lepteutypa aquatica (MFLU 15–0077, holotype). a Specimen. b Section through ascomata. c Section of peridium. d, f Asci.
e Paraphyses. g–j Ascospores. k Germinating ascospore. l Colony on MEA. Scale bars: b 100 lm, d, f 50 lm, c 25 lm, e 10 lm, g–k 5 lm
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Fungal Diversity (2019) 99:451–660
b Fig. 72 Phylogram generated from maximum likelihood analysis
based on ITS and LSU sequence data for the selected taxa of
Xylariales (with Chaetosphaeria innumera and Ch. jonesii as
outgroup). The best scoring RAxML tree with a final likelihood
value of - 11696.396327 is presented. RAxML bootstrap support
values equal to or greater than 75% are given before the forward slash
(black). Bayesian posterior probability equal to or higher than 0.95
are given after the forward slash (red). Hyphen (‘-’) indicates a value
lower than 75% for RAxML and Bayesian posterior probability lower
than 0.95. Newly generated sequences are in red. Ex-type strains are
in bold
625
Cuspidatispora xiphiago Shearer & Bartolata was introduced by Miller et al. (2006) with sexual morph which was
collected from freshwater habitats. During our investigation on lignicolous freshwater fungi in China, a
hyphomycetous fungus was collected from decaying wood
submerged in freshwater habitats. It is characterized by
macronematous, mononematous, solitary, erect, unbranched, septate, straight or flexuous, cylindrical conidiophores, polyblastic, integrated, terminal and subterminal
conidiogenous cells and brown to dark brown, obpyriform,
papillate, guttulate, smooth conidia. Phylogenetic analysis
Fig. 73 Arthrinium aquaticum (MFLU 18–1628, holotype). a Colonies on wood. b, d–f Conidia. c Conidiogenous cells and conidia.
g Germinating conidium. h Culture on PDA from surface. Scale bars: b 30 lm, c–g 10 lm
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Fungal Diversity (2019) 99:451–660
b Fig. 74 Phylogram generated from maximum likelihood analysis
based on ITS sequence data for species of Arthrinium (with Seiridium
marginatum and S. phylicae as outgroup). The best scoring RAxML
tree with a final likelihood value of - 4218.246068 is presented.
RAxML bootstrap support values equal to or greater than 75% are
given before the forward slash (black). Maximum parsimony
bootstrap support values equal to or greater than 75% are given after
the forward slash (red). Hyphen (‘-’) indicates a value lower than 75%
for RAxML and maximum parsimony. Newly generated sequences
are in red. Ex-type strains are in bold
shows that our isolate clusters with Cuspidatispora xiphiago with strong support. We therefore identify it as asexual
morph of Cuspidatispora xiphiago based on the one fungus
one name rule. It is the first record for China.
Lockerbia K.D. Hyde, Sydowia 46(1): 23 (1994)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Hyde (1993b), Raja and Shearer
(2008).
Type species: Lockerbia palmicola K.D. Hyde, Sydowia
46(1): 24 (1994)
Notes: The genus Lockerbia, typified by L. palmicola,
was originally found on a terrestrial palm rachis in a
rainforest in Queensland, Australia (Hyde 1993b). Raja and
Shearer (2008) introduced the second species for this genus
and named it as Lockerbia striata Raja & Shearer, it was
originally found on submerged decorticated woody debris
in a creek on an unidentified piece of herbaceous debris in a
lake. Lockerbia striata was also found on submerged
decorticated wood in two lotic habitats in Mississippi (Raja
and Shearer 2008).
Lockerbia striata Raja & Shearer
Distribution: USA, Florida, Blackwater River State
Forest, Horns Creek Swamp/Andrew Lake near Silver
Lake Recreation Area, on submerged herbaceous debris;
Mississippi, Big black River, on submerged decorticated
woody debris (Raja and Shearer 2008).
Asexual morph: Undetermined
Notes: Holotype ILL 40118. Sequence data is not
available.
Sordariomycetidae genera incertae sedis
Hanliniomyces Raja & Shearer, Mycologia 100(3): 471
(2008)
Asexual morph Undetermined. Sexual morph Descriptions and illustration see Raja and Shearer (2008).
Type species: Hanliniomyces hyaloapicalis Raja &
Shearer, Mycologia 100(3): 471 (2008)
Notes: Raja and Shearer (2008) introduced the genus
Hanliniomyces with single species Hanliniomyces
hyaloapicalis Raja & Shearer which was collected from
freshwater habitats. Presently, only one species was
reported in this genus.
627
Hanliniomyces hyaloapicalis Raja & Shearer
Distribution: USA, Florida, Big Cypress National Preserve, Cypress Swamp Loop Road 5/Blackwater River
State Forest, Penny Creek/Blackwater River, north end at
Kennedy Bridge, on submerged partially decorticated
woody debris (Raja and Shearer 2008).
Asexual morph: Undetermined
Notes: Holotype ILL 40116, other specimens collected
from freshwater habitats: F37–1, F37–2. Sequence data is
not available.
Hydromelitis Ferrer et al., Mycologia 104(4): 876 (2012)
Asexual morph Undetermined. Sexual morph Descriptions and illustration see Ferrer et al. (2012).
Type species: Hydromelitis pulchella Ferrer et al.,
Mycologia 104(4): 877 (2012)
Notes: The genus Hydromelitis was introduced by Ferrer
et al. (2012) with one species Hydromelitis pulchella which
was collected from freshwater habitats. The single species
is only known from the type locality.
Hydromelitis pulchella Ferrer et al.
Distribution: Costa Rica, Alajuela and Heredia, on
submerged wood (Ferrer et al. 2012).
Asexual morph: Undetermined
Notes: Holotype ILL, Ferrer & Salazar, Ferrer 284–2,
other specimens collected from freshwater habitats:
AF284–1, A468–1, A468–2, A468–4. LSU and SSU
sequence data are available.
Subclass Xylariomycetidae O.E. Erikss & Winka
Xylariales Nannf.
Hyponectriaceae Petr
Physalospora Niessl, Verh. nat. Ver. Brünn 14: 170 (1876)
Asexual morph Undetermined. Sexual morph Ascomata
scattered, immersed to partially immersed, becoming
erumpent, ostiolate, globose to subglobose, membranous,
dark brown to black, papillate. Peridium composed of
layers of elongated pesudoparenchymatic cells. Paraphyses
abundant, simple, septate, immersed in a gel matrix,
extending above the asci. Asci unitunicate, cylindrical to
clavate, with a apical ring. Ascospores oblong, broadly
obovoid, or subglobose to oval, surrounded by a gelatinous
sheath, aseptate, hyaline, or pale brownish-yellow (Shear
1907; Raja and Shearer 2008).
Type species: Physalospora alpestris Niessl, Verh. nat.
Ver. Brünn 14: 170 (1876)
Notes: Physalospora is a widespread genus and more
than 450 epithets are listed in Index Fungorum (Taylor
et al. 2001; Index Fungorum 2018). Presently, two species
have been reported from freshwater habitats.
Physalospora aquatica Ingold
Distribution: UK, Anglia, Lancashire, on submerged
Eleocharis palustris (Ingold 1955).
Asexual morph: Undetermined
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Fungal Diversity (2019) 99:451–660
b Fig. 75 Peroneutypa lignicola (MFLU 15–0081). a Specimen. b,
c Appearance of stromata on substrate. d Horizontal section through
stroma. e Vertical section through stroma. f Section of peridium. g–
j Asci. k–m Ascospores. Scale bars: c, d 500 lm, e 300 lm, f 25 lm,
g–j 10 lm, k–m 5 lm
Notes: Holotype IMI 51811. Sequence data is not
available.
Physalospora limnetica Raja & Shearer
Distribution: USA, Florida, Apalachicola National Forest, on submerged herbaceous debris (Raja and Shearer
2008).
Asexual morph: Undetermined
Notes: Holotype ILL40115. Sequence data is not
available. This species is only known from the type
locality.
629
consisting of a dark brown, 21–27 lm thick pseudoparenchymatous outer layer and a narrow hyaline to yellowish hyphal inner layer. Paraphyses apically free,
simple, septate, hyaline, 4–6 lm wide, multi-guttulate.
Asci 126–138 9 8–10 lm (
x = 132 9 9 lm, n = 20), 8spored, unitunicate, long cylindrical, thick-walled, hyaline
to
pale
brown.
Ascospores
15–17 9 5–7 lm
(
x = 16 9 6 lm, n = 20), uniseriate, oblong to reniform,
straight to slightly curved, guttulate, pale brown, smooth,
thick-walled.
Material examined: THAILAND, Chiang Rai Province,
saprobic on submerged decaying wood in a freshwater
stream, November 2013, K.D. Hyde, ZL-23 (MFLU
15–0077, holotype), ex-type living culture MFLUCC
14–0045.
Notes: Lepteutypa aquatica resembles L. uniseptata in
having subglobose, black, immersed ascomata, paraphyses
ca 5 lm wide, septate, multiguttulate, asci long cylindrical
and smooth, thick-walled, pale brown ascospores with
similar size (Tsui et al. 2001d). However, Lepteutypa
aquatica differs from L. uniseptata in having thicker
peridium, asci without apical ring and oblong to reniform,
aseptate ascospore. Phylogenetic analysis also shows that
Lepteutypa aquatica is distinct from L. uniseptata
(Fig. 72).
Amphisphaeriaceae G. Winter
Lepteutypa Petr., Ann. Mycol. 21: 276 (1923)
Asexual morph Undetermined. Sexual morph Saprobic
on decaying wood. Ascomata single or in groups,
immersed, globose or subglobose, black. Peridium brown,
pseudoparenchymatous, hyaline inside. Paraphyses
numerous, simple, sometimes multiguttulate. Asci
8-spored, unitunicate, cylindrical, oblong, thick-walled.
Ascospores uniseriate, oblong to ellipsoid, straight, rarely
Apiosporaceae Hyde et al.
curved, normally with a central scarcely constricted
Arthrinium Kunze, Mykologische Hefte (Leipzig) 1: 9
euseptum, guttulate, aseptate or septate, light to yellow- or
(1817)
medium brown, thick-walled.
Holomorph Descriptions and illustration refer to
Type species. Lepteutypa fuckelii (G.H. Otth) Petr.,
Maharachchikumbura et al. (2016).
Annls mycol. 21(3/4): 276 (1923)
Type species: Arthrinium caricicola Kunze & J.C.
Notes: Lepteutypa Petr. was introduced to accommodate
Schmidt, Mykologische Hefte (Leipzig) 1: 9 (1817)
a single species L. fuckelii (Nitschke) Petr. (Petrak, 1923).
Notes: Arthrinium, an asexual typified genus which has
Several papers have since then dealt with Lepteutypa, the
been traditionally linked to the sexual typified genus
most recent one is Jaklitsch et al. (2016).
Apiospora (Ellis 1971; Seifert et al. 2011). Arthrinium was
Lepteutypa uniseptata (Tsui et al.) Jaklitsch & Voglmayr,
confirmed as the asexual morph of Apiospora by molecular
Persoonia 37: 88 (2016)
data (Crous and Groenewald 2013; Senanayake et al. 2015)
Distribution: China, Hong Kong, Tai Po, Lam Tsuen
and Apiospora, Cordella and Pteroconium were reduced to
River, on submerged wood (Tsui et al. 2001d).
synonymy with Arthrinium (Crous and Groenewald 2013).
Asexual morph: Undetermined
Arthrinium species are geographically widely distributed in
Notes: Holotype IFRD 8733. LSU, SSU and RPB2
various hosts (Wang et al. 2018). Many species of
sequence data are available.
Arthrinium are associated with plants as endophytes or
Lepteutypa aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
saprobes, as well as plant pathogens on some important
nov.
ornamentals (Chen et al. 2014; Li et al. 2016b). In this
Index Fungorum number: IF 555687, Facesoffungi
study, we introduced a new Arthrinium species collected
number: FoF 05478, Fig. 71
from submerged wood in freshwater.
Etymology: Referring to the aquatic habitat of this
Arthrinium aquaticum Z.L. Luo, K.D. Hyde & H.Y. Su,
fungus
sp. nov.
Holotype: MFLU 15–0077
Index Fungorum number: IF 555688, Facesoffungi
Saprobic on decaying wood submerged in freshwater
number: FoF 05479, Fig. 73
habitats. Asexual morph Undetermined. Sexual morph
Etymology: Referring to the aquatic habitat of this
Ascomata immersed in wood, subglobose to depressed
fungus
globose, 250–320 lm high, 300–330 lm diam. Peridium
123
630
Holotype: MFLU 18–1628
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on the substratum superficial, dark brown to black. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in
clusters on hyphae, hyaline to pale brown, smooth, doliform to
ampulliform. Conidia 9–11 lm long, 8–10 lm wide
(
x = 10 9 9 lm, n = 20), globose to subglobose, olivaceous
to brown, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April
2015, Z.L. Luo, S-642 (MFLU 18–1628, holotype, HKAS
92855, isotype), ex-type living culture DLUCC 0642.
Notes: Arthrinium aquaticum resembles A. camelliae-sinensis in having conidiophores reduced to conidiogenous
cells, erect, hyaline to pale brown, smooth, doliform to
ampulliform conidiogenous cells, globose to subglobose,
smooth conidia with similar size (Wang et al. 2018). However,
Phylogenetic analysis shows that they are different species in
this genus (Fig. 74). A. aquaticum also shares similar morphological characters with A. saccharicola in having conidiophores reduced to conidiogenous cells, ampulliform
conidiogenous cells, globose to subglobose, smooth conidia
(Crous et al. 2014a, b). However, A. aquaticum differs from A.
saccharicola by its smooth conidiogenous cells, larger conidia
(9–11 9 8–10 lm vs. 8–9 9 4–6 lm).
Diatrypaceae Nitschke
Peroneutypa Berl., Icon. fung. (Abellini) 3(3–4): 80 (1902)
Asexual morph Coelomycetous, forming pycnidia on PDA
medium. Conidiomata superficial, solitary or aggregated,
subconical, globose to subglobose, shiny, with smooth
surface, white, dark brown to black. Pycnidial walls
comprising several layers of dark brown, compressed
hyphae, arranged in a textura intricata. Conidiogenous cells
holoblastic, cylindrical, straight or curved, apically distorted on conidial secession. Conidia lunate to filiform,
curved or rarely straight, with flattened base and blunt
apex, hyaline. Sexual morph Stromata solitary to gregarious, locules, immersed, becoming raised to erumpent by a
long ostiolar canal, black, glabrous, circular to irregular in
shape. Ascomata perithecial, immersed in a stroma, black,
globose to subglobose. Ostioles cylindrical, black. Peridium thick, dark brown to brown of out layer, subhyaline of
inner layer. Asci 8-spored, unitunicate, hyaline. Ascospores
allantoid, elongate, straight to curved, hyaline to pale yellowish, smooth-walled.
Type species: Peroneutypa bellula (Desm.) Berl., Icon.
fung. (Abellini) 3(3–4): 81 (1902)
Notes: Berlese (1902) introduced the genus Peroneutypa
(Diatrypaceae) to accommodate P. bellula, P. corniculata
and P. heteracantha without designating the type species.
Rappaz (1987) proposed P. bellula as the type species
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Fungal Diversity (2019) 99:451–660
because of the good condition of the herbarium of this
species. Carmarán et al. (2006) reinstated Peroneutypa and
accepted eight species from Eutypella and Echinomyces in
this genus. Shang et al. (2018) revised the taxonomy of the
genus based on morphology and combined ITS and TUB2
sequence data, and an updated key to 13 Peroneutypa
species was also provided. In this study, we introduce one
new species Peroneutypa lignicola based on morphology
and phylogeny and it is the first time to collect Peroneutypa
species from freshwater habitats.
Peroneutypa lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555689, Facesoffungi
number: FoF 05480, Fig. 75
Etymology: Referring to this fungus dwelling on wood
Holotype: MFLU 15–0081
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Undetermined. Sexual morph
Stromata 609–685 lm high, 1196–1246 lm diam., solitary
to gregarious, 4–6 locules, immersed, becoming raised to
erumpent by a long ostiolar canal, black, glabrous, circular
to irregular in shape, arranged in longitudinally, with
conspicuous, clustered, cylindrical prominent ostioles on
substrate surface. Ascomata 448–662 lm high, 355–
445 lm diam., perithecial, immersed in a stroma, black,
globose to subglobose. Ostioles cylindrical, black. Peridium 62–92 lm thick, composed of two type layers, outer
layer comprising thick-walled, dark brown to black cells of
textura angularis, inner layer comprising of thin-walled,
hyaline cells of textura angularis to textura prismatica.
Asci 15–17 9 5–7 lm (
x = 16 9 6 lm, n = 20), 8-spored,
unitunicate, oblong or cylindrical, sessile, hyaline. Ascospores 5–7 9 1.5–2.5 lm (
x = 6 9 2 lm, n = 20),
seriate, elongate-allantoid, straight to slightly curved,
aseptate, with olive-green guttules at both ends, hyaline,
smooth-walled.
Material examined: THAILAND, Chiang Rai Province,
saprobic on decaying wood submerged in a freshwater
stream, November 2013, Z.L. Luo, ZL-14 (MFLU
15–0081, holotype), ex-type living culture MFLUCC
14–0040.
Notes: Peroneutypa lignicola mostly resembles P.
mackenziei in having solitary to gregarious, immersed,
loculate stromata, perithecial, black, globose to subglobose
ascomata immersed in a stroma, unitunicate, 8-spored,
sessile asci and aseptate, elongate-allantoid ascospores
with similar size (Shang et al. 2017). However, Peroneutypa lignicola differs from P. mackenziei in having thicker
peridium (62–92 vs. 45–65 lm) and ascospores with
obviously olive-green guttules at both ends. Phylogenetic
analysis also shows that Peroneutypa lignicola is distinct
from P. mackenziei (Fig. 72).
Fungal Diversity (2019) 99:451–660
631
Fig. 76 Hypoxylon lignicola (MFLU 18–1629, holotype). a Colonies on wood. b–e Conidiophores, Conidiogenous cells with conidia. f–
j conidia. l, m, n Culture on PDA from above and reverse. Scale bars: b 50 lm, c–e 25 lm, f 10 lm, g–j 5 lm
123
632
Fig. 77 Phylogram generated from maximum likelihood analysis
based on ITS sequence data for species of Hypoxylaceae (with
Xylaria hypoxylon as outgroup). The best scoring RAxML tree with a
final likelihood value of - 30247.954400 is presented. RAxML
bootstrap support values equal to or greater than 75% are given before
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Fungal Diversity (2019) 99:451–660
the forward slash (black). Bayesian posterior probability equal to or
higher than 0.95 are given after the forward slash (red). Hyphen (‘-’)
indicates a value lower than 75% for RAxML and bayesian posterior
probability lower than 0.95. Newly generated sequences are in red.
Ex-type strains are in bold
Fungal Diversity (2019) 99:451–660
633
Fig. 78 Seiridium aquaticum (MFLU 18–1627, holotype). a, b Colonies on wood. c Conidiogenous cells and conidia. d–i Conidia.
g Germinating conidium. h Culture on PDA from above and reverse. Scale bars: c–j 20 lm
123
634
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Fungal Diversity (2019) 99:451–660
b Fig. 79 Phylogram generated from maximum likelihood analysis
based on ITS and RPB2 sequence data for species of Seiridium (with
Neopestalotiopsis protearum as outgroup). The best scoring RAxML
tree with a final likelihood value of - 4999.544181 is presented.
RAxML bootstrap support values equal to or greater than 75% are
given before the forward slash (black). Maximum parsimony
bootstrap support values equal to or greater than 75% are given after
the forward slash (red). Hyphen (‘-’) indicates a value lower than 75%
for RAxML and maximum parsimony. Newly generated sequences
are in red. Ex-type strains are in bold
Hypoxylaceae DC.
Hypoxylon Bull., Hist. Champ. Fr. (Paris) 1: 168 (1791)
Asexual morph Colonies on the substratum, effuse,
superficial, gray to pale brown. Conidiophores macronematous, mononematous, cylindrical, single, straight to
curved, unbranched, septate, pale brown to brown. Conidiogenous cells integrated, intercalary, hyaline to pale
brown, smooth. Conidia globose to subglobose, or ellipsoid, gray to pale brown, smooth-walled. Sexual morph
Descriptions refer to Daranagama et al. (2017).
Type species: Hypoxylon fragiforme (Pers.) J. Kickx f.,
FI. crypt. Louvain (Bruxelles): 116 (1835)
Notes: Hypoxylon is a species rich genus with more than
170 known species (Kuhnert et al. 2014), and it is extensively
studied for its morphology, phylogeny and chemotaxonomy.
With a cosmopolitan distribution, Hypoxylon species inhabit
mainly woody substrates as saprobes (Ju and Rogers 1996;
Stadler et al. 2008; Kuhnert et al. 2014). In this study, we
introduce a new species Hypoxylon lignicola based on morphology and phylogenetic analysis, and it is the first time to
report Hypoxylon species from freshwater habitats.
Hypoxylon lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555690, Facesoffungi
number: FoF 05481, Fig. 76
Etymology: Referring to this fungus dwelling on wood
Holotype: MFLU 18–1629
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on the substratum,
effuse, superficial, gray to pale brown. Conidiophores 258–
320 lm long, 4.5–5.5 lm wide (
x = 289 9 5 lm, n = 20),
macronematous, mononematous, cylindrical, single,
straight to curved, unbranched, septate, pale brown to
brown. Conidiogenous cells polyblastic, integrated, intercalary, hyaline to pale brown, smooth. Conidia 3–5 lm
long, 3–4 lm wide (
x = 4 9 3.5 lm, n = 20), globose to
subglobose, or ellipsoid, gray to pale brown, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April
2015, Z.L. Luo, S-331 (MFLU 18–1629, holotype, HKAS
92856, isotype), ex-type living culture MFLUCC 16–0926;
KUMCC 15–0332.
635
Notes: Hypoxylon lignicola resembles the asexual
morph of H. griseobrunneum in having macronematous
conidiophores, ellipsoid, smooth conidia with similar size
(Kuhnert et al. 2014). However, H. lignicola differs from
H. griseobrunneum in having unbranched, pale brown to
brown conidiophores, intercalary, hyaline to pale brown
conidiogenous cells, gray to pale brown conidia. Phylogenetic analysis shows that H. lignicola is distinct from
other Hypoxylon species (Fig. 77).
Sporocadaceae Corda
Hymenopleella Munk, Dansk bot. Ark. 15(no. 2): 89
(1953)
Holomorph Descriptions and illustration refer to (Liu et al.
2019).
Type species: Hymenopleella hippophaëicola Jaklitsch
& Voglmayr, Persoonia 37: 96 (2016)
Notes: Dyrithiopsis was introduced by Jeewon et al.
(2003) with D. lakefuxianensis as type species which was
collected from a lake in Yunnan, China. Liu et al. (2019)
synonymised Dyrithiopsis under Hymenopleella based on
morphology and phylogenetic studies.
Hymenopleella lakefuxianensis (L. Cai et al.) Liu et al.
: Dyrithiopsis lakefuxianensis L. Cai, Jeewon & K.D.
Hyde
Distribution: China, Yunnan Province, Lake Fuxian, on
submerged wood (Jeewon et al. 2003).
Asexual morph: See Jeewon et al. (2003)
Notes: Holotype IFRD 8807. LSU sequence data is
available.
Seiridium Nees, Syst. Pilze (Würzburg): 22 (1816) [181617]
Holomorph Descriptions and illustrations see Bonthond
et al. (2018).
Type species: Seiridium marginatum Nees, Syst. Pilze
(Würzburg): 23 (1816) [1816-17]
Notes: The genus Seiridium was established based on S.
marginatum (Nees 1817), collected from rose stems in
Germany and recently epitypified by Jaklitsch et al. (2016).
Bonthond et al. (2018) investigated the phylogeny of the
genus Seiridium, and epitypified species for which no extype culture is currently available. Seiridium (Sordariomycetes, Xylariales, Sporocadaceae) comprises a variety
of mainly plant pathogenic fungi, such as S. cardinale, S.
cupressi and S. unicorne are considered responsible for a
pandemic of cypress canker disease, impacting plantations
for wood production and ornamental tree cultivation
(Boesewinkel 1983; Graniti 1986, 1998; Bonthond et al.
2018).
Seiridium aquaticum Z.L. Luo, K.D. Hyde & H.Y. Su, sp.
nov.
Index Fungorum number: IF 555691, Facesoffungi
number: FoF 05482, Fig. 78
123
636
Etymology: Referring to the aquatic habitat of this
fungus
Holotype: MFLU 18–1627
Saprobic on decaying wood submerged in freshwater
habitats. Asexual morph Colonies on the substratum
superficial, dark brown to black. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells subcylindrical,
hyaline, smooth. Conidia 29–35 lm long, 12–14 lm wide
(
x = 32 9 13 lm, n = 20), straight to slightly curved,
5-septate, bearing minute marginal frills, four median cells,
smooth, doliform to ellipsoidal, dark brown to black, septa
darker than the rest of the cells; basal cell obconic with a
truncate base, hyaline, smooth-walled; apical cell conical,
hyaline, smooth-walled. Sexual morph Undetermined.
Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a
freshwater stream, September 2016, S.M. Tang, S-793
(MFLU 18–1627, holotype), ex-type living culture
MFLUCC 17–0474; Ibid., saprobic on decaying wood
submerged in a freshwater stream, April 2016, S.M. Tang,
S-837, living culture MFLUCC 17–1314; Ibid., saprobic on
decaying wood submerged in a freshwater stream, July
2014, L.W. Wang, S-136 (HKAS 84016, paratype), living
culture MFLUCC 18–1308.
Notes: Seiridium aquaticum resembles S. spyridicola in
having subcylindrical, hyaline, smooth conidiogenous
cells, septate conidia bearing marginal frills, with hyaline
cells at both ends (Bonthond et al. 2018). However, S.
aquaticum differs from S. spyridicola in having larger
conidia (29–35 9 12–14 vs. 24–28 9 8–10.5 lm).
Seiridium aquaticum is also distinct from other Seridium
species by its minute marginal frills at both ends of conidia.
Phylogenetic analysis shows that Seiridium aquaticum is
distinct from other Seridium species (Fig. 79).
Xylariaceae Tul. & C. Tul.
Anthostomella Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
Padova, Sér. 4 4: 84 (1875)
Holomorph Descriptions and illustrations refer to Daranagama et al. (2017).
Type species: Anthostomella tomicoides Sacc., Atti Soc.
Veneto-Trent. Sci. Nat., Padova, Sér. 4 4: 101 (1875)
Notes: Many Anthostomella species are distributed in
tropics, subtropics and even temperate regions (Daranagama et al. 2017). A world monograph of Anthostomella was published by Lu and Hyde (2000). The
phylogenetic placement of Anthostomella was established
by Daranagama et al. (2015, 2016) based on molecular
data. Only one species has been reported from freshwater
habitats.
Anthostomella aquatica K.D. Hyde & Goh
Distribution: Australia, north Queensland, Lake Barrine, on submerged wood (Hyde and Goh 1998b).
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Fungal Diversity (2019) 99:451–660
Asexual morph: see Hyde and Goh (1998b)
Notes: Holotype IFRD 8653. SSU sequence data is
available.
Sordariomycetes genera incertae sedis
Aquadulciospora Fallah & Shearer, Mycologia 93(3): 570
(2001)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Fallah and Shearer (2001).
Type species: Aquadulciospora rhomboidia Fallah &
Shearer, Mycologia 93(3): 571 (2001)
Notes: Aquadulciospora was introduced by Fallah and
Shearer (2001) to accommodate A. rhomboidia which is
characterized by immersed, scattered or in rows, globose or
obpyriform, papillate ascomata, nitunicate, numerou, basal,
fasciculate, cylindric-clavate asci and ellipsoid-rhomboid
to fusoid, multiguttulate, hyaline, smooth ascospores.
Currently, there is only one species accepted in this genus.
Aquadulciospora rhomboidia Fallah & Shearer
Distribution: USA, Wisconsin, Sparkling Lake, on submerged decomposed stems of Scirpus validus (Fallah and
Shearer 2001).
Asexual morph: Undetermined
Notes: Holotype ILLS 53999, other specimens collected
from freshwater habitats: ILLS 54001, ILLS 54002, ILLS
54003, ILLS 54004, ILLS 54006, ILLS 54007. Sequence
data is not available.
Aquasphaeria K.D. Hyde, Nova Hedwigia 61: 122 (1995)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Hyde (1995a).
Type species: Aquasphaeria dimorphospora K.D. Hyde,
Nova Hedwigia 61(1–2): 123 (1995)
Notes: The genus Aquasphaeria was introduced by
Hyde (1995a) based on the collection from submerged
wood in freshwater habitats. Currently, this genus includes
only one species.
Aquasphaeria dimorphospora K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Hyde 1995b).
Asexual morph: Undetermined
Notes: Holotype BRIP 21482. Sequence data is not
available.
Ascoyunnania L. Cai & K.D. Hyde, Fungal Divers 18: 2
(2005)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Cai et al. (2005).
Type species: Ascoyunnania aquatica L. Cai & K.D.
Hyde, Fungal Divers 18: 3 (2005)
Notes: The genus Ascoyunnania was introduced by Cai
et al. (2005) based on the collection from submerged wood
in freshwater habitats in China. Currently, this genus
includes only one species.
Ascoyunnania aquatica L. Cai & K.D. Hyde
Fungal Diversity (2019) 99:451–660
Distribution: China, Yunnan Province, Jinghong City,
on submerged bamboo in a small forest stream (Cai et al.
2005).
Asexual morph: Undetermined
Notes: Holotype PDD 75039. Sequence data is not
available. This species is only known from the type
locality.
Paoayensis Cabanela et al., Cryptog. Mycol. 28(4): 303
(2007)
Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Cabanela et al. (2007).
Type species: Paoayensis lignicola Cabanela et al.,
Cryptog. Mycol. 28(4): 304 (2007)
Notes: Ascoyunnania was introduced by Cabanela et al.
(2007) with single species which was collected from
freshwater habitats. One species was accepted in this
genus.
Paoayensis lignicola Cabanela et al.
Distribution: Phillipines, Ilocos Norte, on submerged
wood in Paoay Lake (Cabanela et al. 2007).
Asexual morph: Undetermined
Notes: Holotype HKU (M) 17516 (now in IFRD). LSU
and SSU sequence data are available.
Rivulicola K.D. Hyde, Nova Hedwigia 64(1–2): 186
(1997)
Asexual morph Undetermined. Sexual morph Ascomata
scattered to gregarious, hyaline to light brown, membranous; venter globose to subglobose, ostiolate. Neck hyaline,
periphysate. Peridium comprising several layers of hyaline
to light brown, elongated, flattened cells. Parphyses simple, septate, filamentous. Asci unitunicate, numerous, basal,
fasciculate, cylindrical, with apical ring. Ascospores septate, ellipsoidal, hyaline, smooth-walled.
Type species: Rivulicola incrustata K.D. Hyde, Nova
Hedwigia 64(1–2): 186 (1997)
Notes: Rivulicola was introduced for a freshwater
ascomycete with hyaline ascomata, cylindrical asci with a
discoid refractive apical ring, hyaline, multiseptate, and
ellipsoidal ascospores (Hyde et al. 1997). Two more species were introduced by Ranghoo et al. (2000) and Raja
et al. (2009a) respectively. All species of this genus were
collected from freshwater habitats.
Rivulicola aquatica Ranghoo & K.D. Hyde
Distribution: China, Hong Kong, Tai Po, on submerged
wood in freshwater (Ranghoo et al. 2000).
Asexual morph: Undetermined
Notes: Holotype IFRD 8843. Sequence data is not
available. This species is only known from the type
locality.
Rivulicola cygnea Raja & Shearer
637
Distribution: USA, Florida, Apalachicola National Forest, on submerged decorticated wood in a unnamed lake
(Raja et al. 2009a).
Asexual morph: Undetermined
Notes: Holotype ILL 40111. Sequence data is not
available. This species is only known from the type
locality.
Rivulicola incrustata K.D. Hyde
Distribution: Australia, Queensland, on submerged
wood (Hyde et al. 1997).
Asexual morph: Undetermined
Notes: Holotype BRIP 23340. Sequence data is not
available. This species is only known from the type
locality.
Saccardoella Speg., Michelia 1(no. 5): 461 (1879)
Asexual morph Undetermined. Sexual morph Ascomata
perithecioid, solitary or gregarious, immersed or erumpent,
globose to subglobose, coriaceous or carbonaceous, dark
brown to black, papillate. Ostiole mostly central, conical,
short, brown to black, periphysate. Peridium medium,
composed of two strata. Paraphyses hyphae-like, filiform,
numerous, septate, not embedded in a gelatinous matrix.
Asci 8-spored, long-cylindrical or filiform, short-pedicellate, with apical ring. Ascospores fusiform, multi-septate,
euseptate or distoseptate, sometimes slightly constricted at
the septum, guttulate, some are surrounded by mucilaginous sheath, hyaline, smooth, thick-walled.
Type species: Saccardoella montellica Speg., Michelia
1(no. 5): 461 (1879)
Notes: Saccardoella is a widespread genus and comprises 22 epithets in Index Fungorum (December 2018).
Among these species, four of them have been reported
from freshwater habitats.
Saccardoella aquatica Tsui et al.
Distribution: China, Hong Kong, New Territories, Lam
Tsuen River, on submerged wood (Tsui et al. 1998b);
South Africa, Durban, Palmet River, on submerged wood
(Tsui et al. 1998b).
Asexual morph: Undetermined
Notes: Holotype IFRD 8846. Sequence data is not
available.
Saccardoella horizontalis Fallah & Shearer
Distribution: Philippines, Liput River, on submerged
wood (cai et al. 2003b); USA, Wisconsin, on submerged
wood (Fallah and Shearer 2001).
Asexual morph: Undetermined
Notes: Holotype ILLS 54025. Sequence data is not
available.
Saccardoella lacustris Fallah & Shearer
Distribution: USA, Wisconsin, on submerged wood
(Fallah and Shearer 2001).
Asexual morph: Undetermined
123
638
Notes: Holotype ILLS 54026. Sequence data is not
available.
Saccardoella minuta L. Cai & K.D. Hyde
Distribution: Philippines, Liput River, on submerged
bamboo and wood (Cai et al. 2002c; cai et al. 2003b).
Asexual morph: Undetermined
Notes: Holotype PDD 75037, isotype IFRD 8847.
Sequence data is not available.
Stanjehughesia Subram., Proc. Indian natn Sci. Acad., Part
B. Biol. Sci. 58(4): 184 (1992)
Asexual morph Colonies on natural substrate effuse,
black. Mycelium superficial, composed of branched, septate, pale brown to brown, smooth-walled hyphae. Conidiophores absent or short, septate, brown to dark brown.
Conidiogenous cells holoblastic, monoblastic, determinate,
simple, brown. Conidia solitary, acrogenous, straight or
curved, obclavate to obclavate-rostrate, distoseptate, septate, brown to dark brown, smooth-walled. Sexual morph
Undetermined.
Type species: Stanjehughesia hormiscioides (Corda)
Subram., Proc. Indian natn Sci. Acad., Part B. Biol. Sci.
58(4): 184 (1992)
Notes: Subramanian (1992) established Stanjehughesia
to accommodate five Sporidesmium species. The genus is
mainly characterized by very reduced or absent conidiophores and solitary, euseptate conidia seceding schizolytically from monoblastic, determinate, lageniform or
ampulliform, short conidiogenous cells.
Stanjehughesia clavata D.A.C.Almeida & Gusmão
Distribution: Brazil, Bahia, on submerged decaying
bark (de Almeidala et al. 2014).
Sexual morph: Undetermined
Notes: Holotype HUEFS 155078. Sequence data is not
available.
Discussion
Our morphological and phylogenetic study on freshwater
Sordariomycetes has resulted in taxa belonging to different
orders and genera. These are: Annulatascales (10 genera),
Atractosporales (2 genera), Chaetosphaeriales (16 genera),
Coniochaetales (1 genus), Conioscyphales (1 genus), Cordanales (1 genus), Coronophorales (1 genus), Diaporthales
(5 genera), Distoseptisporales (2 genera), Fuscosporellales
(6 genera), Glomerellales (2 genera), Hypocreales (19
genera), Jobellisiales (1 genus), Magnaporthales (4 genera), Microascales (10 genera), Myrmecridiales (1 genus),
Ophiostomatales (1 genus), Phomatosporales (1 genus),
Phyllachorales (2 genera), Pisorisporiales (2 genera),
Pleurotheciales (5 genera), Savoryellales (4 genera), Sordariales (13 genera), Sporidesmiales (1 genus),
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Tirisporellales (1 genus), Togniniales (1 genus), Torpedosporales (1 genus), Trichosphaeriales (2 genera),
Xenospadicoidales (3 genera) and Xylariales (8 genera).
Diaporthomycetidae
Annulatascales
The order Annulatascales was introduced by
Maharachchikumbura et al. (2015) and includes the single
family Annulatascaceae which was established by Wong
et al. (1998a) to accommodate species of Annulatascus
(typified by A. velatispora) and related genera. Members of
Annulatascaceae are primarily characterized by the presence of relatively massive, wedge-shaped, bipartite asci
with J-apical ascal ring (Wong et al. 1998a). However,
Réblová and Winka (2001) showed that species characterized by these characters are polyphyletic. To clarify the
taxonomy of these polyphyletic species, even new genera,
such as Annulusmagnus has been established to accommodate the distantly related species Annulatascus triseptatus. Dayarathne et al. (2016) provided the epitype for the
type species of Annulatascus. Réblová et al. (2018) synonymized Annulatascus biatriisporus under Torrentispora
biatriispora based on morphology and phylogeny. The
relationship and placement of several species in Annulatascus are more resolved now and the taxonomic problems of polyphyly were addressed.
Zhang et al. (2017a, b) studied Annulatascaceae-like taxa
and introduced one new order and six new families based on
morphology and phylogenetic analyses. In their study,
Aquaticola, Dictyosporella, Fusoidispora and Pseudoannulatascus are excluded from Annulatascaceae and placed in
Diaporthomycetidae genera incertae sedis. Conlariaceae
was introduced for the genus Conlarium, Pseudoproboscisporaceae includes Pseudoproboscispora and Diluviicola,
while Atractosporaceae includes the genera Rubellisphaeria
and Atractospora. Cyanoannulus was assigned to the new
family Woswasiaceae in Diaporthomycetidae families incertae sedis. Later, Réblová et al. (2018) synonymized Fusoidispora under Torrentispora.
In our phylogenetic analysis, the Annulatascales clade
(Fig. 1, clade 17) includes seven genera, viz. Annulatascus,
Annulusmagnus, Ascitendus, Longicollum, Paoayensis,
Pseudoproboscispora and Submersisphaeria. This clade
appears to be related to Myrmecridiales, though the statistical support is low. The genera Pseudoproboscispora
and Submersisphaeria appear in distinct lineages (Fig. 1).
Zhang et al. (2017a, b) introduced a new species Pseudoproboscispora thailandensis and transferred Pseudoproboscispora to the new family Pseudoproboscisporaceae.
However, in our phylogenetic analysis, Pseudoproboscispora caudae-suis does not cluster with the new described
Fungal Diversity (2019) 99:451–660
species. This needs to be further verified by more taxon
sampling especially the inclusion of type-derived DNA
data.
Although Annulusmagnus has been established to
accommodate Annulatascus triseptatus which is phylogenetically distant to A. velatisporus, the remaining members
in Annulatascus still deserves some taxonomic consideration. For instance, while A. saprophyticus, A. velatisporus,
A. hongkongensis and A. nilensis constitute a moderate
supported monophyletic group, the type strain of Annulatascus aquatorba is phylogenetically distant to the former species (Fig. 1, clade 17). We presume that only
further collections of Annulatascus species will clarify
whether we should still consider it as a polyphyletic genus.
Atractosporales
The order Atractosporales was established by Zhang et al.
(2017a, b) comprising three families, viz. Atractosporaceae, Conlariaceae and Pseudoproboscisporaceae. In
morphology, species in this three families share similar
characters in having perithecial, non-stromatic, immersed
to semi-immersed ascomata with neck, unitunicate, eightspored asci with apical ring and hyaline, fusiform ascospores (Hyde et al. 1998c; Liu et al. 2012; Réblová et al.
2016b; Zhang et al. 2017a, b). However, in the phylogenetic analyses of Zhang et al. (2017a, b), these three
families cluster together but without bootstrap support. Our
phylogenetic analysis shows that these three families do
not cluster together as Pseudoproboscisporaceae appears
related to Junewangiaceae (Fig. 1, clade 1 and 2); species
of Atractosporaceae cluster together with strong bootstrap
support (Fig. 1, clade 3) and Conlariaceae formed a distinct
clade as well (Fig. 1, clade 5). There is however, no support to conclusively interpret phylogenetic relationships
among these families. Given the uncertain phylogenetic
placement of Conlariaceae and Pseudoproboscisporaceae,
we therefore suggest that Atractosporales only accepts for
the type family Atractosporaceae, while Conlariaceae and
Pseudoproboscisporaceae
be
assigned
as
Diaporthomycetidae family incertae sedis.
Papulosaceae
Maharachchikumbura et al. (2015) placed two freshwater
genera, Brunneosporella and Fluminicola in this family.
Khemmuk et al. (2016) added Wongia to Papulosaceae
based on the phylogenetic analysis and morphological
characters. Later, Zhang et al. (2017a, b) showed that the
genus Platytrachelon is affiliated to Papulosaceae in their
phylogenetic analysis thus added Platytrachelon to Papulosaceae. However, in our multi-gene phylogenetic analysis, Platytrachelon did not cluster in Papulosaceae and it is
639
more related to Acrodictyaceae but with weak support
(Fig. 1, clade 11).
Pseudoproboscisporaceae
The genus Cateractispora has never been validly published
(Zhang et al. 2017a, b). Ranghoo (1998) provided the LSU
sequence data for the ex-type strain HKUCC3710 and
illustrations for C. recepticuli, but without any descriptions. Zhang et al. (2017a, b) showed that C. recepticuli
clusters
with
Pseudoproboscispora
thailandensis
(MFLUCC 15–0989) with strong support and this is supported in our study (Fig. 1, clade 2). Morphologically,
Cateractispora recepticuli shares similar characters with
Pseudoproboscispora thailandensis as well. According to
the morphology and phylogeny analysis, Cateractispora is
likely to be an invalidly published synonym of Pseudoproboscispora. However, Ranghoo (1998) did not provide
the information for herbarium of Cateractispora recepticuli and the specimen is unavailable for loan to verify the
morphology. Therefore, we follow Zhang et al. (2017a, b)
to keep the name in our phylogenetic tree. Fresh collections
are needed in future study to clarify the classification of
this species.
Diaporthales and Tirisporellales
Seven freshwater species in five genera are listed for the
order Diaporthales in this study. The order Tirisporellales
was introduced by Jones et al. (2015) in the class Sordariomycetes, subclass Diaporthomycetidae and includes a
single family Tirisporellaceae based on SSU and LSU
rDNA sequence analysis. However, Hongsanan et al.
(2017) showed that the order Tirisporellales had a divergence time at 112 MYA and shared a common ancestor
with Sydowiellaceae and Pseudovalsariaceae in their MCC
analysis and Tirisporellales was also closely related to
Pseudovalsaceae in the phylogenetic tree. Our phylogenetic
analysis based on LSU, SSU, RPB2 and TEF1a sequence
data shares corroborates those of Hongsanan et al. (2017)
in that the placement of Tirisporellales is not well supported, but falls within Diaporthales. Tirisporellaceae is
probably a family of the Diaporthales. However, we following Hongsanan et al. (2017) to keep Tirisporellales as
an order under Diaporthomycetidae and further study is
needed to clarify the placement of Tirisporellales.
Distoseptisporales
Su et al. (2016) introduced the family Distoseptisporaceae
typified by Distoseptispora with two species. Yang et al.
(2018a, b) emended description for the genus Distoseptispora. In the past 3 years, thirteen species have been
123
640
Fungal Diversity (2019) 99:451–660
introduced in the genus (Hyde et al. 2016b; Su et al. 2016;
Yang et al. 2018a; Luo et al. 2018a). In our phylogenetic
analysis, all Distoseptisporaceae species cluster together
with high support value (98% ML) and formed distinct
clade with other orders or families in Diaporthomycetidae.
We therefore introduce a new order Distoseptisporales to
accommodate the family Distoseptisporaceae.
In our phylogenetic analysis, four newly generated isolates
forms a distinct subclade within Xenospadicoidales with
strong support (Fig. 1, clade 6). Here, we introduce
Neospadicoides as a new genus in the family Xenospadicoidaceae with three new species.
Magnaporthales
Hypocreales
In Magnaporthales, the freshwater fungi mainly include
members from Ophioceraceae and Pseudohalonectriaceae
with eleven and seven known species, respectively. Magnaporthales has been shown to be monophyletic and
including four families Magnaporthaceae, Ophioceraceae,
Pseudohalonectriaceae and Pyriculariaceae (Thongkantha
et al. 2009; Maharachchikumbura et al. 2015, 2016; Hongsanan et al. 2017). In our phylogenetic analysis, and that
of Thongkantha et al. (2009) and Cai et al. (2014), one
strain of Ophioceras tenuisporum (SMH 1643) was distinct
from other Ophioceras species and clustered with Ceratosphaeria species. However, ex-type strain of Ophioceras tenuisporum (ATTC 56671) clusters with other
Ophioceras species (Fig. 1, clade 20). The LSU sequence
data of Ophioceras tenuisporum (SMH 1643) was provided
by Huhndorf et al. (2004), but without any illustrations,
descriptions and herbarium information for this strain.
Therefore, further studies are needed to resolve the placement of this strain. On the other hand, most of Ceratosphaeria species were published at 1880s to 1990s,
sequence data are not available for those species. The
specimen needs to be relooked into and sequenced to
clarify its taxonomy.
Wijayawardene et al. (2018) accepted 23 genera in the
family Magnaporthaceae. In this study, the genus Ceratosphaeria is excluded from Magnaporthaceae based on
morphology and phylogeny. In our phylogenetic analyses,
the genera Ceratosphaerella and Mycoleptodiscus do not
group with other genera in Magnaporthaceae. Ceratosphaerella clusters with Ophioceras species and species
of Mycoleptodiscus formed a separate clade and basal to
other families in Magnaporthales (Fig. 15). Therefore,
further studies on Ceratosphaerella and Mycoleptodiscus
are needed in order to resolve the placement of these two
genera.
35 species in 19 genera of freshwater fungi for Hypocreales
are listed in this study. In our phylogenetic analysis, species of Hypocreales cluster together with good support
value (Fig. 1, clade 38, 39).
Sordariomycetidae
Xenospadicoidales
Chaetosphaeriales
The order Xenospadicoidales was established by Réblová
et al. (2018) with a single family Xenospadicoidaceae
including four genera, viz. Calyptosphaeria, Lentomitella,
Spadicoides and Torrentispora. Species in Spadicoides and
Torrentispora have been reported from freshwater habitats.
The order Chaetosphaeriales was established in Sordariomycetidae based on molecular analysis of LSU sequence
data (Huhndorf et al. 2004). In our phylogenetic analysis
(Fig. 1, clade 26), Chaetosphaeriales is close to Phyllachorales.
Presently,
the
order
comprises
123
Hypocreomycetidae
Microascales
Microascales proposed by Luttrell (1951) and validly
introduced by Benny and Kimbrough (1980) to accommodate Chadefaudiellaceae, Microascaceae and Pithoascaceae, and later expanded to include Ceratocystidaceae,
Gondwanamycetaceae, Halosphaeriaceae and Graphiaceae
(Réblová et al. 2011). Maharachchikumbura et al. (2016)
and Hongsanan et al. (2017) accepted Ceratocystidaceae,
Chadefaudiellaceae, Gondwanamycetaceae, Graphiaceae,
Halosphaeriaceae and Microascaceae in Microascales. 32
species of freshwater fungi have been reported for
Microascales. Halosphaeriaceous species are one of the
most typical and common freshwater Sordariomycetes on
submerged wood. In Fig. 1 (clade 40), 15 freshwater
strains cluster in the family Halosphaeriaceae. The genus
Triadelphia is currently placed in the order Microascales as
genera incertae sedis (Wijayawardene et al. 2017, 2018; Lu
et al. 2018b). In our phylogenetic analysis, strains of Triadelphia formed a distinct clade with strong bootstrap
support (Fig. 1, clade 41) and close to the family Graphiaceae. Triadelphia can be easily separated from other
genera in Ceratocystidaceae, Chadefaudiellaceae, Gondwanamycetaceae, Graphiaceae, Halosphaeriaceae and
Microascaceae in having 2–5 forms of conidia (Shearer and
Crane 1971; Li and Ye 2017; Lu et al. 2018b). Based on
morphological characters and phylogenetic evidence, we
introduce a new family Triadelphiaceae in this study.
Fungal Diversity (2019) 99:451–660
641
Table 1 Checklist of the freshwater species currently assigned as Ascomycota genera incertae sedis
Genus
Species
Distribution
Holotype
References
Angulospora
A. aquatica Sv. Nilsson
India, Venezuela
Not indicated
Nilsson (1962)
Arachnophora
A. combuensis Monteiro et al.
Brazil
UEFS 196432
Monteiro et al. (2014)
A. longa Fryar & K.D. Hyde
Australia
BRI
AQ522463
Fryar and Hyde (2018)
Arbusculina
A. fragmentans Marvanová
Slovenia
IMI 317746
Marvanová (1988)
Arthrotaeniolella
A. aquatica Conç. et al.
Brazil
HUEFS
224988
Monteiro et al. (2017a)
Ascagilis
A. bipolaris K.D. Hyde
Australia
BRIP 17155
Hyde (1992c)
Ascomauritiana
A. lignicola Ranghoo & K.D. Hyde
Mauritius
IFRD 8694
Ranghoo and Hyde (1999)
Blastoheterospora
B. catenata Monteiro et al.
Brazil
HUEFS
216679
Monteiro et al. (2017b)
Brachydesmiella
B. anthostomelloidea Goh & K.D. Hyde
Australia
BRIP 23200
Goh and Hyde (1996b)
B. biseptata G. Arnaud
Thailand
Sivichai et al. (1998b)
B. caudata V. Rao & de Hoog
China, India,
Thailand
BIOTEC
SS285
CBS-H 3847
Sivichai et al. (1998b)
B. orientalis (V. Rao & de Hoog) Goh
Thailand
Not indicated
Sivichai et al. (1998b)
B. verrucosa Goh et al.
Malaysia
IFRD 8712
Sivichai et al. (1998b)
Brevicatenospora
B. enteroproliferata Castañeda et al.
Cuba
INIFAT C04/
111
Castañeda-Ruiz et al.
(2006)
Cacumisporium
C. rugosum Tsui et al.
China
HKU
(M) 5548
Tsui et al. (2001e)
C. uniseptatum Wongs.et al.
China
HMAS
196815
Wongsawas et al. (2009)
Candelosynnema
C. ranunculosporum K.D. Hyde & Seifert
Australia
BRIP 19446
Hyde and Seifert (1992)
Cheiromyces
Ch. lignicola Ho et al.
China
HKU
(M) 5937
Ho et al. (2000)
Clavariana
C. aquatica Nawawi
Malaysia
IMI 195181
Descal et al. (1976)
Cryptophiale
Delortia
C. multiseptata Goh & K.D. Hyde
D. aquatica Goh & K.D. Hyde
Australia
Australia
BRIP 23150
BRIP 23323
Goh and Hyde (1996c)
Goh and Hyde (1997b)
D. tumidiapicis Goh & K.D. Hyde
Australia
BRIP 23322
Goh and Hyde (1997b)
Dictyoaquaphila
D. appendiculata Monteiro et al.
Brazil
HUEFS
216013
Monteiro et al. (2016c)
Dictyotrichocladium
D. aquaticum Fiuza et al.
Brazil
HUEFS
215954
Fiuza et al. (2017)
Diplocladiella
aquatica Lee et al.
Brunei
HKU
(M) 8260
Lee et al. (1998)
Filosporella
D. appendiculata Nawawi
Malaysia
IMI 309529
Nawawi (1987)
D. tricladioides Nawawi
Malaysia
IMI 289094
Nawawi (1985)
F. annelidica (Shearer & J.L. Crane) J.L. Crane &
Shearer
USA
ILLS 36352
Crane and Shearer (1977)
F. aquatica Nawawi
Malaysia
IMI 177454
Nawawi (1976)
F. exilis Gulis & Marvanová
Belarus
IMI 378148
F. pinguis Marvanová & Bärl.
New Brunswick
DAOM
220746
Gulis and Marvanová
(1998)
Marvanová and Bärlocher
(1998)
F. versimorpha Marvanová et al.
UK
IMI 344098
Marvanová et al. (1992)
Frigidispora
F. colnensis K.D. Hyde & Goh
UK
IFRD 8759
Hyde and Goh (1999)
Janetia
J. curviapicis Goh & K.D. Hyde
Australia
BRIP 23223
Goh and Hyde (1996d)
Linkosia
L. aquatica Conç. et al.
Brazil
HUEFS
216022
Conceição et al. (2016)
Melanocephala
M. triseptata (Shearer et al.) S. Hughes
USA
ILLS 36154
Shearer et al. (1976)
123
642
Fungal Diversity (2019) 99:451–660
Table 1 (continued)
Genus
Species
Distribution
Holotype
References
Mirandina
M. uncinata Fiuza et al.
Brazil
HUEFS
216003
Fiuza et al. (2016)
Neta
N. angliae K.D. Hyde & Goh
UK
HKU
(M) 3243
Hyde and Goh (1999)
N. patuxentica Shearer & J.L. Crane
USA
ILLS 34520
Shearer and Crane (1971)
lignicola Shearer
USA
ILLS 35536
Shearer and Crane (1971)
Paraceratocladium
P. malaysianum Goh & K.D. Hyde
Malaysia
IFRD 8825
Goh and Hyde (2000a, b)
Phaeomonilia
P. pleiomorpha Castañeda et al.
Mexico
XAL CB746
Castañeda-Ruı́z et al.
(2007)
Phalangispora
P. constricta Nawawi & J. Webster
Malaya
IMI 256650
Potamomyces
P. armatisporus K.D. Hyde
Australia
BRIP 21490
Nawawi and Webster
(1982)
Hyde (1995b)
Proboscispora
P. aquatica S.W. Wong & K.D. Hyde
Australia
HKU
(M) 2792
Wong and Hyde (1999a, b)
Pseudofuscophialis
P. lignicola Sivan. & H.S. Chang
China
IMI 356633
Cai and Hyde (2007)
Pyramidospora
P. quadricellularis Oliveira et al.
Brazil
URM 87706
Oliveira et al. (2015b)
Sirothecium
S. triseriale Hu et al.
China
YMF 6.00017
Hu et al. (2007)
Tricellula
T. aquatica J. Webster
UK
K(M) 235168
Webster (1959)
Triscelophorus
T. acuminatus Nawawi
Malaysia
IMI 184587
Nawawi (1975)
T. magnificus R.H. Petersen
USA
NY
Petersen (1962)
T. monosporus Ingold
UK
Not indicate
Ingold (1943)
Vanakripa
V. fasciata Castañeda et al.
Cuba
MUCL 45276
Castañeda-Ruı́z et al.
(2005)
V. menglensis Hu et al.
China
IFRDC 8882
Hu et al. (2010)
Yinmingella
Y. mitriformis Goh et al.
China
HKU
(M) 8042
Goh et al. (1998c)
Ypsilina
Y. graminea (Ingold et al.) Descals et al.
UK
IMI 123908
Ingold et al. (1968)
Zelotetraploa
Z. aquatica Monteiro et al.
Brazil
HUEFS
216680
Monteiro et al. (2017b)
Chaetosphaeriaceae, Helminthosphaeriaceae, Leptosporellaceae and Linocarpaceae (Huhndorf et al. 2004;
Maharachchikumbura et al. 2016; Konta et al. 2017). There
are several studies have been reported many freshwater
species in Chaetosphaeriaceae in the past 3 years (Liu et al.
2016; Lu et al. 2016; Luo et al. 2016; Yang et al. 2016b;
Wei et al. 2018). In this study, we provide the phylogenetic
tree for Chaetosphaeriales based on combined ITS and
LSU sequence data (Fig. 51).
Morphological characters of the sexual morph of
Chaetosphaeria are similar and hardly distinguishable,
while the asexual morph characters are considered as distinctive characters to identify the species of Chaetosphaeria (Gams and Holubova-Jechova 1976; Huhndorf
et al. 2004). The asexual morph of Chaetosphaeria is
hyphomycetous with macronematous, mononematous
conidiophores, monophialidic or polyphialidic, hyaline,
123
conidiogenous cells with a distinct funnel-shaped collarette
and hyaline to brown, aseptate to multi-septate, guttulate or
eguttulate conidia, with or without appendages (Maharachchikumbura et al. 2016). Three newly obtained isolates are morphologically identical to Catenularia.
However, phylogenetic analysis showed that they cluster
with Chaetosphaeria species with good support value
(Fig. 51). Previous studies have reported that some Chaetosphaeria species with Catenularia asexual morph, such
as Ch. cubensis, Ch. cupulifera and Ch. novaezelandiae
(Holubova-Jechova 1982; Réblová 2004). We therefore
accommodate our isolates in the genus Chaetosphaeria.
Catenularia appears to be a synonym of Chaetosphaeria
and further studies on these groups are needed to resolve
the placement of these two genera.
In our phylogenetic analysis, species of Chaetosphaeria
forms seven distinct clades within Chaetosphaeriaceae
Fungal Diversity (2019) 99:451–660
(Fig. 51). Further study incorporating herbarium loaning,
sampling and DNA sequencing are needed to resolve the
species placement of this genus.
643
2016a). Thirteen strains including eleven freshwater strains
of Fuscosporellales represent a monophyletic group and
sister to Conioscyphales, Pleurotheciales and Savoryellales
within the Savoryellomycetidae (Fig. 1, clade 46).
Coniochaetales
Pleurotheciales
Shearer and Raja (2018) listed eight freshwater Coniochaeta species belonging to this order with C. gigantospora and C. renispora originally described from
freshwater habitats (Crane and Shearer 1995; Raja et al.
2012). However, among these eight species, Coniochaeta
kellermanii is an invalid name (Index Fungorum 2018) and
another record is named as Coniochaeta sp. 4022 (Shearer
and Raja 2018), we therefore only accepted six freshwater
Coniochaeta species.
Cordanales
The order Cordanales was introduced by HernándezRestrepo et al. (2015b) to accommodate the family Cordanaceae with a single genus Cordana Preuss. Previous
studies described two freshwater Cordana species, viz.
Cordana abramovii and C. uniseptata, which were collected from submerged wood in Perú and China respectively (Cai et al. 2004a; Zelski et al. 2014). In this study,
we introduce two new species C. aquatica and C. lignicola,
and provide descriptions, illustrations and DNA sequence
data for C. abramovii and C. terrestris. Four Cordana
species have been reported from freshwater habitats.
Sordariales
In this study, we list 26 species of Sordariales that have
been reported from freshwater habitats based on the
available literature. In our phylogenetic analysis, species of
Chaetomiaceae and Sordariaceae form monophyletic lineages respectively with good support (Fig. 1, clade 29).
However, further study is needed to resolve the species
placement of genera in Lasiosphaeriaceae.
Savoryellomycetidae
Fuscosporellales
Fuscosporellales is a recently established order with most
of its members from freshwater environment (Yang et al.
2016a, 2017). Fuscosporellales is characterized by astromatic, immersed to semi-immersed ascomata, fragile,
several layered peridium, cylindrical to cylindricfusiform
asci, fusiform, uniseriate, transversely septate ascospores
and sporodochial conidiomata, micronematous or
macronematous conidiophores usually reduced to conidiogenous cells, obovate to obpyriform conidia (Yang et al.
Twenty-four species in five genera of freshwater fungi have
been reported for the Pleurotheciales, mostly were known
as new species growing on submerged wood (Réblová et al.
2016a; Hyde et al. 2018; Luo et al. 2018b). In our phylogenetic analysis, 34 freshwater strains cluster in Pleurotheciales with well support (99% ML). In the
phylogenetic analysis of Xia et al. (2017), Rhexoacrodictys
species clustered in Savoryellaceae with strong support
values (100% ML, 0.99 PP), and Rhexoacrodictys was
assigned to Savoryellaceae. However, they did not include
species of Conioscyphales, Fuscosporellales and Pleurotheciales in their study. Species of Conioscyphales,
Fuscosporellales, Pleurotheciales and Savoryellales are
included herein and results show that Rhexoacrodictys
species cluster in Pleurotheciellaceae (Fig. 1, clade 43).
Therefore we accommodate Rhexoacrodictys as a genus of
Pleurotheciellaceae.
Savoryellales
Boonyuen et al. (2011) established the order Savoryellales
based on morphology and combined LSU, SSU and RPB2
DNA sequence data. Freshwater species in this order are
accommodated in Ascotaiwania, Canalisporium and Savoryella. In this study, we introduce a new genus Dematiosporium to accommodate an asexual fungus collected
from freshwater habitats. 26 freshwater species have been
reported in the order Savoryellales.
Xylariomycetidae
Xylariales
The classification of Xylariomycetidae is rather controversial. Senanayake et al. (2015) re-validated the order
Amphisphaeriales based on both sequence data and morphology. However, Maharachchikumbura et al. (2016)
showed that there was little support to validate the order
Amphisphaeriales and the placement of the families within
the subclass Xylariomycetidae were not stable. However,
Hongsanan et al. (2017) provided evidence to maintain the
Amphisphaeriales and Xylariales as distinct orders in
Xylariomycetidae based on molecular clock evidence, but
still with weak support. In our phylogenetic analysis, the
placement of families within Xylariomycetidae was also
We
therefore
follow
not
stable
(Fig. 1).
123
644
Maharachchikumbura et al. (2016) to keep the order
Xylariales to represent the taxa in the Xylariomycetidae
until further data are available to support to validate the
order Amphisphaeriales.
We introduce five new freshwater species in Xylariales
based on morphology and DNA sequence data. Eleven
species of freshwater fungi are accepted in the order
Xylariales.
The use of DNA sequence data coupled with phylogenetic analyses has significantly improved our understanding on the phylogeny and evolution of freshwater
ascomycetes (Vijajkrishna et al. 2006). Most of the new
genera and species of freshwater fungi published in the past
ten years are provided with molecular data, for example:
Aquapteridospora, Baipadisphaeria, Conlarium, Distoseptispora and Mucispora (Pinruan et al. 2010; Liu et al.
2012; Yang et al. 2015, 2017, 2018a; Su et al. 2016; Luo
et al. 2018a). However, there are still some new genera and
species of freshwater fungi published without molecular
data, such as Arthrotaeniolella, Dictyoaquaphila, Linkosia
and Zelotetraploa (Conceição et al. 2016; Monteiro et al.
2016c, 2017a, b). The placement of many freshwater Sordariomycetes are currently uncertain and suffers from lack
of living cultures and type sequences. Therefore, to obtain
a better understanding of phylogeny in freshwater Sordariomycetes in a natural classification system, further
investigations are needed. Future studies should put high
priority to obtain living cultures. We also provide a
checklist of the freshwater species which are currently
assigned to Ascomycota genera incertae sedis (Table 1).
Further studies incorporating herbarium materials, more
sampling and DNA sequencing are needed to resolve the
placement of these genera.
Acknowledgements This work was funded by the grants of the
National Natural Science Foundation of China (NSFC 31660008,
31860006) and Fungal diversity conservation and utilization innovation team of Dali University (ZKLX2019213). The authors would
like to thank Dr. Shaun Pennycook (Landcare Research Manaaki
Whenua, New Zealand) for advising on the fungal names. Professor
Yong-Chang Zhao and Professor Shu-Hong Li (Institute of Biotechnology and Gerplamic Resources, Yunnan Academy of Agricultural
Sciences) are appreciated for kindly support on molecular work. JianKui Liu thanks the National Natural Science Foundation of China
(NSFC 31600032). Zong-Long Luo thanks Xiao-Ying Liu, Qi Dai,
Xing-Chen Tao, Hong-Wei Shen, Xi-Jun Su and Song-Ming Tang
(College of Agriculture & Biological Science, Dali University) for the
great assistance on sample collection. Rajesh Jeewon thanks Mae Fah
Luang University and University of Mauritius for research support.
K.D. Hyde thanks the Thailand Research Funds for the grant entitled
Impact of climate change on fungal diversity and biogeography in the
Greater Mekong Subregion, grant number: RDG6130001.
Open Access This article is distributed under the terms of the Creative
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Affiliations
Zong-Long Luo1,2 • Kevin D. Hyde2 • Jian-Kui (Jack) Liu3 • Sajeewa S. N. Maharachchikumbura3,4 •
Rajesh Jeewon5 • Dan-Feng Bao1,6 • Darbhe Jayarama Bhat7 • Chuan-Gen Lin2 • Wen-Li Li1 • Jing Yang2
Ning-Guo Liu2 • Yong-Zhong Lu8 • Ruvishika S. Jayawardena2 • Jun-Fu Li2 • Hong-Yan Su1
•
1
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, People’s Republic of China
5
Department of Health Sciences, Faculty of Science,
University of Mauritius, Reduit 80837, Mauritius
2
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
6
3
School of Life Science and Technology, University of
Electronic Science and Technology of China,
Chengdu 611731, People’s Republic of China
Department of Entomology and Plant Pathology, Faculty of
Agriculture, Chiang Mai University, Chiang Mai 50200,
Thailand
7
No. 128/1-J, Azad Co-Op Housing Society, Curca,
Goa Velha 403108, India
8
School of Pharmaceutical Engineering, Guizhou Institute of
Technology, Guiyang 550003, People’s Republic of China
4
Department of Crop Sciences, College of Agricultural and
Marine Sciences, Sultan Qaboos University, P.O. Box 34,
Alkhoud, 123 Muscat, Oman
123