Freshwater fungal numbers

Abstract

A comprehensive account of fungal classification from freshwater habitats is outlined and discussed in the present review based on literature of biodiversity studies and recent morpho-phylogenetic analyses. A total of 3,870 freshwater fungal species are listed with additional details on the isolation source, habitat, geographical distribution, and molecular data. The Ascomycota (2,968 species, 1,018 genera) dominated the freshwater fungal taxa wherein Sordariomycetes (823 species, 298 genera) had the largest number, followed by Dothideomycetes (677 species, 229 genera), Eurotiomycetes (276 species, 49 genera), and Leotiomycetes (260 species, 83 genera). Other phyla included in the updated classification of freshwater fungi are: Chytridiomycota (333 species, 97 genera), Rozellomycota (221 species, 105 genera), Basidiomycota (218 species, 100 genera), Blastocladiomycota (47 species, 10 genera), Monoblepharomycota (29 species, 6 genera), Mucoromycota (19 species, 10 genera), Aphelidiomycota (15 species, 3 genera), Entomophthoromycota (6 species, 4 genera), Mortierellomycota (5 species, 3 genera), Olpidiomycota (4 species, 1 genus), Zoopagomycota (3 species, 2 genera), and Sanchytriomycota (2 species, 2 genera). The freshwater fungi belong to 1,361 genera, 386 families and 145 orders. The Pleosporales and Laboulbeniaceae are the largest freshwater fungal order and family comprised of 391 and 185 species, respectively. The most speciose genera are Chitonomyces (87, Laboulbeniomycetes), Verrucaria (50, Eurotiomycetes), Rhizophydium (52, Rhizophydiomycetes), Penicillium (47, Eurotiomycetes), and Candida (42, Saccharomycetes).

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Vol.:(0123456789)
1 3
Fungal Diversity
https://doi.org/10.1007/s13225-022-00503-2
REVIEW
Freshwater fungal numbers
MarkS.Calabon1,2 · KevinD.Hyde1,2,3,4 · E.B.GarethJones5 · Zong‑LongLuo6 · WeiDong4 ·
VedprakashG.Hurdeal1,2 · EleniGentekaki1,2 · WalterRossi7 · MarcoLeonardi7 ·
VinodhiniThiyagaraja1,3,8,9,10 · AnisS.Lestari1,2,11 · Hong‑WeiShen1,2,6 · Dan‑FengBao1,6,10 ·
NattawutBoonyuen12 · MingZeng1,2,13
Received: 14 November 2021 / Accepted: 13 February 2022
© The Author(s) under exclusive licence to Mushroom Research Foundation 2022
Abstract
A comprehensive account of fungal classification from freshwater habitats is outlined and discussed in the present review
based on literature of biodiversity studies and recent morpho-phylogenetic analyses. A total of 3,870 freshwater fungal
species are listed with additional details on the isolation source, habitat, geographical distribution, and molecular data. The
Ascomycota (2,968 species, 1,018 genera) dominated the freshwater fungal taxa wherein Sordariomycetes (823 species, 298
genera) had the largest number, followed by Dothideomycetes (677 species, 229 genera), Eurotiomycetes (276 species, 49
genera), and Leotiomycetes (260 species, 83 genera). Other phyla included in the updated classification of freshwater fungi
are: Chytridiomycota (333 species, 97 genera), Rozellomycota (221 species, 105 genera), Basidiomycota (218 species, 100
genera), Blastocladiomycota (47 species, 10 genera), Monoblepharomycota (29 species, 6 genera), Mucoromycota (19 spe-
cies, 10 genera), Aphelidiomycota (15 species, 3 genera), Entomophthoromycota (6 species, 4 genera), Mortierellomycota
(5 species, 3 genera), Olpidiomycota (4 species, 1 genus), Zoopagomycota (3 species, 2 genera), and Sanchytriomycota
(2 species, 2 genera). The freshwater fungi belong to 1,361 genera, 386 families and 145 orders. The Pleosporales and
Laboulbeniaceae are the largest freshwater fungal order and family comprised of 391 and 185 species, respectively. The
most speciose genera are Chitonomyces (87, Laboulbeniomycetes), Verrucaria (50, Eurotiomycetes), Rhizophydium (52,
Rhizophydiomycetes), Penicillium (47, Eurotiomycetes), and Candida (42, Saccharomycetes).
Keywords Fungal classification· Fungal taxonomy· Freshwater fungi· Higher fungi· Molecular taxonomy· Basal clades
Introduction
It has been over 140years since Saccardo (1880) described
the first freshwater hyphomycetes, Heliscus lugdunensis (≡
Neonectria lugdunensis). This was followed by the discovery
of four novel taxa [Tetracladium marchalianum, Clavari-
opsis aquatica, Lemonniera aquatica, Anguillospora longis-
sima (≡ Amniculicola longissima)] by de Wildeman (1893,
1894, 1895) from different freshwater habitats and on differ-
ent substrates. Ingold’s (1942) documentation of 16 freshwa-
ter hyphomycetous species, of which 10 are novel species,
and identification of the typical habitat of freshwater fungi
served as an impetus for further studies on the occurrence of
fungal taxa, both sexual and asexual, in freshwater habitats
(Ranzoni 1953; Tubaki 1957; Petersen 1962, 1963a, b; Nils-
son 1964; Ingold 1975; Webster and Descals 1981; Tubaki
etal. 1983; Dudka 1985; Nawawi 1985a; Goh and Hyde
1996c; Goh 1997; Sivichai etal. 2002; Shearer etal. 2004,
2007; Lu etal. 2018c; Luo etal. 2019; Dong etal. 2020b).
The classical methods used to document freshwater fun-
gal composition have been significantly improved by recent
developments in fungal identification using molecular tech-
niques. In the classification of freshwater fungi, incorpora-
tion of molecular data in the analysis of extant and novel
species provides a better understanding of the phylogenetic
relationships of freshwater taxa (e.g., generic bounda-
ries delimitation of polyphyletic taxa, resolving species
complexes, linking asexual-sexual morphs, or synasexual
morphs). Furthermore, continuous exploration of fresh-
water habitats led to introduction of many novel taxa (Lu
etal. 2018c; Bao etal. 2020; Dong etal. 2020a; Hsieh etal.
Handling Editor: Jian-Kui Liu.
* Kevin D. Hyde
kdhyde3@gmail.com
Extended author information available on the last page of the article

Fungal Diversity
1 3
2021a). For instance, novel species from China and Thailand
totaled 298 novel species from 2015 to 2020. Unfortunately,
for almost two decades of morpho-phylogenetic taxonomy
of freshwater fungi, there are no published information on
the overall known number of taxa unlike those of fungi in
marine habitats wherein the marine fungal numbers stand
as 1901 species under 769 genera (Jones etal. 2019). The
available published information includes biodiversity studies
in particular geographical areas (Shearer and Crane 1971;
Ho etal. 2001; Pinnoi etal. 2006; Pinruan etal. 2007; Zhang
etal. 2011; Hu etal. 2013a), selected hosts (Gessner and
Van Ryckegem 2003; Pinruan etal. 2007, 2014; Fiuza etal.
2019), or of species within taxonomic (Pinruan etal. 2007,
2014; Luo etal. 2016b, 2019; Zhang etal. 2017a; Lu etal.
2018c; Fiuza etal. 2019; Dong etal. 2020b, 2021a; Hyde
etal. 2021) or morphological groups (Ingold 1975; Webster
and Descals 1981; Bärlocher 1987; Shearer 1993a; Schoen-
lein-Crusius and Piccolo Grandi 2003; Bao etal. 2021a).
The scattered data on freshwater fungi are currently com-
piled in an online database, https:// fresh water fungi. org/, but
since the website only started in early 2020 and in the pro-
cess of collating all the freshwater fungal data, there are no
definite numbers given yet (Calabon etal. 2020b). The only
information in the literature so far is the estimated number of
freshwater fungi to be around 3069–4145, of which, Chytrid-
iomycota (500–1289 species) and Ascomycota (622 species)
have the greatest number of taxa (Jones etal. 2014a). These
figures were considered as an underestimate as there is little
data for many of the aquatic groups, and many microhabi-
tats, substrates, and geographical areas are unexplored or
underexplored and yet to be surveyed.
This paper presents an up-to-date classification of fresh-
water fungi and the number of taxa recorded in freshwater
habitats. The classification scheme follows Wijayawardene
etal. (2020) unless the taxonomy has been updated recently.
The text is divided into three main sections: (1) Classifica-
tion of freshwater Ascomycota, (2) Basidiomycota, and (3)
Freshwater fungal basal clades. For Ascomycota section, a
special section Freshwater lichens is included in order to
discuss and enumerate lichenicolous and lichen-like taxa
from Arthoniomycetes, Candelariomycetes, Coniocybomy-
cetes, Dothideomycetes, Eurotiomycetes, Lecanoromycetes,
Lichinomycetes, and Sordariomycetes. After the section on
the Basidiomycota, the section on Freshwater yeasts was
included with a discussion on their taxonomic classifica-
tion. Furthermore, since Ingold’s contribution in freshwa-
ter mycology is momentous, a special section on Ingoldian
fungi with a taxonomic outline of taxa from freshwater habi-
tats is included in the present review. Every table dealing
with the taxonomic classification of freshwater fungi include
the taxon name, isolation source, and its geographical distri-
bution. Freshwater fungi with molecular data, including the
country where the taxa were isolated, are in bold.
Origin offreshwater fungi
Molecular evidence suggests that fungi first appeared about
1.5 billion years ago and evolved from one common ances-
tor (Heckman etal. 2001). The ancestral fungi may have
lived in water and had flagella, like the members of Chytridi-
omycota (James etal. 2006; Liu etal. 2006; Naranjo-Ortiz
and Gabaldón 2019). Molecular studies of Liu etal. (2006)
showed that the major fungal phyla, Ascomycota, Basidi-
omycota, and Zygomycota formed a well-supported clade,
and they lacked flagella, while Chytridiomycota are mainly
flagellated, and it formed a basal and early diverging clade
within the fungal kingdom. The loss of a flagellum has
occurred only one time, leading to derived fungal phyla
(Zygomycota, Basidiomycota and Ascomycota) to adapt
from the aquatic environment to terrestrial habitats (James
etal. 2006; Liu etal. 2006).
Fungi colonized the land probably during the Cambrian,
over 500 million years ago (MYA), however, the fungal ter-
ritorialization times are still a matter of debate as the land
fungal fossil records are reported from the Devonian (Taylor
and Osborn 1996; Brundrett 2002). Fungal hyphae and spore
cysts in a stem of Asteroxylon mackiei were also discovered
in the Rhynie chert from the Devonian (419.2 to 358.9 MYA).
The earliest land plants appeared around 470 MYA (Kenrick
etal. 2012; Edwards etal. 2014), while oomycetes had already
colonized terrestrial habitats at that time (Taylor etal. 2014;
Selosse etal. 2015). Microorganisms became abundant within
plant fossils since their territorialization (Taylor etal. 2014;
Selosse etal. 2015). A rich terrestrial fungal fossil (e.g., fossil
chytrids, Glomales and fossil hyphae associated with wood
decay) have been found during the Devonian, around 400
MYA (Taylor etal. 1992, 1995, 2014; Hass etal. 1994; Remy
etal. 1994a, b) and terrestrial fungal fossils were reported as
arbuscular mycorrhizal-like symbioses (Simon etal. 1993;
Raghukumar 2017). Simon etal. (1993) investigated the ori-
gin of arbuscular mycorrhizal fungi and their results showed
that these fungi diverged 353–462 MYA, which is consistent
with the fossil records, indicating establishment of arbuscular
mycorrhizal-like symbiosis between fungi and plant roots as
a major event in the colonization of land.
Ascomycota and Basidiomycota diverged at a similar
time with fungal territorialization, around 500–650 MYA,
and mainly saprobic on vascular plants (Lücking etal. 2009;
Gueidan etal. 2011). Studies showed that they had a ter-
restrial origin (Heckman etal. 2001; James etal. 2006; Liu
etal. 2006; Hyde etal. 2021). Fungal adaption to terrestrial
habitats requires a variety of ecological strategies to obtain
nutrients, such as saprotrophism, parasitism, humification,
and the development of mutual relationships. Ascomycota
and Basidiomycota are the largest groups in the fungal king-
dom, and have been reported as endophytes, pathogens, and

Fungal Diversity
1 3
saprobes of plants/animals, indicating rapid diversification
on land through different nutritional modes.
Around 250 MYA fungi rapidly increased in numbers
and became abundant as the result of the Permian–Tri-
assic extinction event. Following this event, numerous
plants and animals evolved and provided abundant hosts
for fungi (Eshet etal. 1995; Schubert and Bottjer 1995).
Molecular analyses showed that the subclasses and some
orders of the two classes Sordariomycetes and Dothideo-
mycetes diverged at 150–250 MYA (Hyde etal. 2017a,
2020d; Liu etal. 2017; Hongsanan etal. 2020). Several
studies indicate that after the Cretaceous-Tertiary extinc-
tion (65 MYA), fungi dramatically increased in numbers
(Ward etal. 2005b). Recent molecular clock analyses also
support this view, with many species diverged after 65
MYA (Hyde etal. 2017a; Liu etal. 2017; Samarakoon
etal. 2019; Hongsanan etal. 2020).
At present, the origin, evolution, and taxonomy of fungi
are better understood with the use of molecular analyses for
their study. Early studies of the fungi placed them in the plant
kingdom, and later separated from the plant kingdom as the
fifth kingdom (Whittaker 1969). Earlier, the true fungi com-
prised four phyla viz. Ascomycota, Basidiomycota Chytridio-
mycota, and Zygomycota (Whittaker 1969). While, presently,
the fungal kingdom comprises 20 phyla (Wijayawardene
etal. 2020; Voigt etal. 2021). Among them, eight phyla viz.
Aphelidiomycota, Ascomycota, Basidiomycota, Blastocladi-
omycota, Chytridiomycota, Monoblepharomycota, Mortierel-
lomycota and Rozellomycota include freshwater fungi (Jones
etal. 2014a; Wijayawardene etal. 2018, 2020). Freshwater
fungi have multiple origins and evolved separately through
different lineages (Shearer 1993a). Chytridiomycota are
likely to have evolved from marine ancestors, while Ascomy-
cota and Basidiomycota originated from terrestrial habitats
(Heckman etal. 2001; Vijaykrishna etal. 2006). Ascomy-
cota are the dominant phylum of freshwater fungi. Freshwater
ascomycetes probably are immigrants from terrestrial envi-
ronments and mainly through three evolutionary pathways
(Shearer 1993a; Vijaykrishna etal. 2006; Hyde etal. 2021).
Classication offreshwater fungi
Our comprehensive review of published data of biodiver-
sity studies and/or single or multi-species taxonomic stud-
ies, resulted in an updated classification of freshwater fungi
under thirteen phyla (45 classes), and documentation of
3,870 species from freshwater habitats: Ascomycota 2,968
(1,018 genera), Chytridiomycota 333 (97 genera), Rozello-
mycota 221 (105 genera), Basidiomycota 218 (100 genera),
Blastocladiomycota 47 (10 genera), Monoblepharomycota
29 (6 genera), Mucoromycota 19 (10 genera), Aphelidio-
mycota 15 (3 genera), Entomophthoromycota 6 (4 genera),
Mortierellomycota 5 (3 genera), Olpidiomycota 4 (1 genus),
Zoopagomycota 3 (2 genera), and Sanchytriomycota 2 (2
genera) (Table1). For Ascomycota, Sordariomycetes has
the highest number of taxa which accounts for 27.68% (823
species in 298 genera), followed by Dothideomycetes (677
species, 229 genera). Chytridiomycetes dominated the taxa
under Chytridiomycota with 181 species recorded while 201
taxa are under Microsporidea (Rozellomycota). The fresh-
water yeasts constitute taxa from Ascomycota and Basidi-
omycota with 163 species each. The total freshwater fungal
number, 3,870 species, included in this review is within
the estimated range of Jones etal. (2014a) which is around
3,069–4,145 species. The present review lists all the fungal
taxa in freshwater habitats. Some species, including their
occurrence and host substrates, will inevitably be missed
in the process of searching the published literatures. All the
updates will be added on the website, www . fresh water fungi.
org (Calabon etal. 2020b).
Classication offreshwater Ascomycota
The freshwater Ascomycota comprises thirteen classes namely
Arthoniomycetes, Candelariomycetes, Coniocybomycetes,
Dothideomycetes, Eurotiomycetes, Laboulbeniomycetes,
Lecanoromycetes, Leotiomycetes, Lichinomycetes, Orbili-
omycetes, Pezizomycetes, Saccharomycetes, and Sordariomy-
cetes. Ascomycota genera incertae sedis totaled to 252 species
(in 141 genera) and two species listed under Pezizomycotina
genera incertae sedis (Pyrenocollema strontianense, Psam-
mina stipitata). Taxonomic classification of freshwater Can-
delariomycetes, Coniocybomycetes, Lecanoromycetes, and
Lichinomycetes were listed in “Freshwater lichens” section as
members of these classes are lichens and lichenicolous taxa.
Arthoniomycetes
Arthoniomycetes is the largest taxonomic group of primarily
lichenized fungi outside of Lecanoromycetes. The taxa are
mostly tropical and subtropical microlichens with bitunicate
asci in ascohymenial apothecia (Henssen and Thor 1994).
Multi-locus phylogenetic analysis shows that Arthoniomy-
cetes is monophyletic and sister class to Dothideomycetes
(Spatafora etal. 2006). On the latest outline of fungi and
fungus-like taxa, Wijayawardene etal. (2020) listed two
orders under Arthoniomycetes: Arthoniales (116 genera) and
Lichenostigmatales (3 genera). Few freshwater Arthoniomy-
cetes have been recorded from freshwater habitats (Björk
etal. 2009; Morse 2013; Khodosovtsev etal. 2016) and the
taxa listed in Table2 accounts for twelve species (in 8 gen-
era) but undoubtedly there are more taxa to be discovered.
The additional information on freshwater lichenicolous taxa
is discussed in “Freshwater Lichens” section.

Fungal Diversity
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Table 1 Taxonomic classifica-
tion and numbers of freshwater
fungi
Class Order Family Number
of species
Classification of freshwater Ascomycota
Arthoniomycetes Arthoniales Arthoniaceae 3
Chrysotrichaceae 2
Lecanographaceae 1
Opegraphaceae 1
Roccellaceae 5
Candelariomycetes Candelariales Candelariaceae 4
Coniocybomycetes Coniocybales Coniocybaceae 4
Dothideomycetes Asterinales Morenoinaceae 1
Botryosphaeriales Botryosphaeriaceae 3
Phyllostictaceae 1
Capnodiales Teratosphaeriaceae 2
Mycosphaerellaceae 5
Neodevriesiaceae 6
Collemopsidiales Xanthopyreniaceae 3
Dothideales Saccotheciaceae 2
Dyfrolomycetales Pleurotremataceae 1
Hysteriales Hysteriaceae 1
Hysteriales genus incertae sedis 1
Jahnulales Aliquandostipitaceae 37
Kirschsteiniotheliales Kirschsteiniotheliaceae 6
Kirschsteiniotheliales genera incertae sedis 6
Lichenotheliales Lichenotheliaceae 2
Microthyriales Microthyriales genera incertae sedis 1
Minutisphaerales Acrogenosporaceae 13
Minutisphaeraceae 5
Monoblastiales Monoblastiaceae 1
Natipusillales Natipusillaceae 4
Pleosporales Acrocalymmaceae 3
Aigialaceae 27
Amniculicolaceae 2
Anteagloniaceae 2
Aquasubmersaceae 2
Astrosphaeriellaceae 24
Bambusicolaceae 1
Corynesporascaceae 4
Cucurbitariaceae 2
Cyclothyriellaceae 2
Delitschiaceae 3
Dictyosporiaceae 55
Didymosphaeriaceae 6
Dothidotthiaceae 5
Hermatomycetaceae 1
Latoruaceae 2
Lentitheciaceae 15
Leptosphaeriaceae 6
Lindgomycetaceae 25
Longipedicellataceae 5
Lophiostomataceae 20
Lophiotremataceae 2

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Massarinaceae 6
Melanommataceae 20
Morosphaeriaceae 13
Mycoporaceae 1
Neopyrenochaetaceae 3
Nigrogranaceae 2
Occultibambusaceae 5
Parabambusicolaceae 3
Periconiaceae 8
Phaeoseptaceae 2
Phaeosphaeriaceae 16
Pleomassariaceae 4
Pleomonodictydaceae 1
Pleosporaceae 19
Pseudoastrosphaeriellaceae 6
Pseudoberkleasmiaceae 1
Pyrenochaetopsidaceae 1
Roussoellaceae 5
Sporormiaceae 7
Striatiguttulaceae 1
Teichosporaceae 1
Testudinaceae 1
Tetraplosphaeriaceae 13
Thyridariaceae 1
Torulaceae 15
Trematosphaeriaceae 9
Wicklowiaceae 3
Pleosporales genera incertae sedis 10
Trypetheliales Polycoccaceae 1
Trypetheliaceae 1
Tubeufiales Tubeufiaceae 133
Wiesneriomycetaceae 12
Venturiales Venturiaceae 1
Dothideomycetes fami-
lies incertae sedis Endosporiaceae 1
Argynnaceae 2
Trichothyriaceae 1
Pseudorobillardaceae 2
Dothideomycetes gen-
era incertae sedis Dothideomycetes genera incertae sedis 31
Eurotiomycetes Arachnomycetales Arachnomycetaceae 1
Chaetothyriales Cyphellophoraceae 4
Herpotrichiellaceae 40
Chaetothyriales genera incertae sedis 2
Chaetothyriomycetidae
family incertae sedis Rhynchostomataceae 1
Eurotiales Aspergillaceae 67
Thermoascaceae 1
Trichocomaceae 8
Mycocaliciales Mycocaliciaceae 9

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Onygenales Arthrodermataceae 4
Onygenaceae 7
Sclerococcales Dactylosporaceae 5
Verrucariales Adelococcaceae 1
Verrucariaceae 126
Laboulbeniomycetes Laboulbeniales Ceratomycetaceae 73
Laboulbeniaceae 185
Pyxidiophorales Pyxidiophoraceae 1
Lecanoromycetes Acarosporales Acarosporaceae 4
Arctomiales Arctomiaceae 1
Baeomycetales Baeomycetaceae 1
Hymeneliaceae 4
Protothelenellaceae 1
Trapeliaceae 6
Xylographaceae 1
Caliciales Caliciaceae 4
Physciaceae 21
Lecanorales Pilocarpaceae 4
Catillariaceae 1
Cladoniaceae 13
Haematommataceae 1
Lecanoraceae 18
Parmeliaceae 13
Psoraceae 1
Ramalinaceae 10
Scoliciosporaceae 2
Stereocaulaceae 1
Lecideales Lecideaceae 8
Ostropales Coenogoniaceae 1
Graphidaceae 1
Gyalectaceae 2
Porinaceae 8
Peltigerales Collemataceae 13
Lobariaceae 2
Massalongiaceae 1
Peltigeraceae 9
Placynthiaceae 4
Pertusariales Megasporaceae 6
Microcaliciaceae 1
Ochrolechiaceae 1
Pertusariaceae 1
Rhizocarpales Rhizocarpaceae 3
Teloschistales Teloschistaceae 14
Umbilicariales Ophioparmaceae 1
Umbilicariaceae 1
Ostropomycetidae gen-
era incertae sedis Ostropomycetidae genera incertae sedis 1
Leotiomycetes Helotiales Discinellaceae 24

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Gelatinodiscaceae 3
Helotiaceae 19
Helotiales genera incertae sedis 19
Hyaloscyphaceae 18
Hydrocinaceae 13
Lachnaceae 5
Loramycetaceae 3
Mitrulaceae 2
Mollisiaceae 27
Ploettnerulaceae 9
Tricladiaceae 33
Vibrisseaceae 12
Lauriomycetales Lauriomycetaceae 3
Leotiales Leotiales genera incertae sedis 5
Rhytismatales Rhytismataceae 4
Thelebolales Thelebolales genera incertae sedis 2
Leotiomycetes families
incertae sedis Calloriaceae 34
Cenangiaceae 1
Hamatocanthoscyphaceae 3
Pezizellaceae 3
Rutstroemiaceae 1
Sclerotiniaceae 2
Leotiomycetes genera
incertae sedis Leotiomycetes genera incertae sedis 15
Lichinomycetes Lichinales Lichinaceae 21
Peltulaceae 3
Orbiliomycetes Orbiliales Orbiliaceae 19
Pezizomycetes Pezizales Ascobolaceae 2
Pezizaceae 7
Pyronemataceae 2
Rhizinaceae 2
Saccharomycetes Saccharomycetales Debaryomycetaceae 21
Dipodascaceae 7
Lipomycetaceae 1
Metschnikowiaceae 4
Phaffomycetaceae 7
Pichiaceae 12
Saccharomycetaceae 30
Saccharomycetales genera incertae sedis 50
Saccharomycodaceae 6
Saccharomycopsidaceae 5
Trichomonascaceae 5
Wickerhamomycetaceae 10
Sordariomycetes Annulatascales Annulatascaceae 36
Aquapteridosporaceae 4
Atractosporales Atractosporaceae 7
Barbatosphaeriales Barbatosphaeriaceae 1
Cancellidiales Cancellidiaceae 7
Ceratolentales Bullimycetaceae 3

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Conlariales Conlariaceae 4
Diaporthales Diaporthaceae 3
Gnomoniaceae 4
Diaporthales genera incertae sedis 1
Sydowiellaceae 1
Diaporthomycetidae
family incertae sedis Acrodictyaceae 6
Junewangiaceae 21
Papulosaceae 7
Pseudoproboscisporaceae 5
Pseudostanjehughesiaceae 2
Woswasiaceae 1
Diaporthomycetidae
genera incertae sedis Diaporthomycetidae genera incertae sedis 10
Distoseptisporales Distoseptisporaceae 22
Jobellisiales Jobellisiaceae 3
Magnaporthales Ceratosphaeriaceae 5
Magnaporthaceae 5
Ophioceraceae 11
Pseudohalonectriaceae 6
Pyriculariaceae 3
Myrmecridiales Myrmecridiaceae 4
Ophiostomatales Ophiostomataceae 1
Parasympodiellales Parasympodiellaceae 1
Phomatosporales Phomatosporaceae 7
Rhamphoriales Rhamphoriaceae 2
Sporidesmiales Sporidesmiaceae 22
Tirisporellales Tirisporellaceae 2
Togniniales Togniniaceae 4
Trichosphaeriales Trichosphaeriaceae 4
Xenospadicoidales Xenospadicoidaceae 21
Coronophorales Bertiaceae 1
Chaetosphaerellaceae 1
Coronophorales genera incertae sedis 2
Nitschkiaceae 1
Glomerellales Australiascaceae 2
Glomerellaceae 1
Plectosphaerellaceae 2
Reticulascaceae 8
Hypocreales Bionectriaceae 5
Hypocreaceae 4
Hypocreales genera incertae sedis 4
Nectriaceae 57
Niessliaceae 2
Sarocladiaceae 1
Stachybotryaceae 7
Hypocreomycetidae
genera incertae sedis Hypocreomycetidae genera incertae sedis 5
Microascales Gondwanamycetaceae 1

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Halosphaeriaceae 42
Microascaceae 2
Microascales genera incertae sedis 1
Triadelphiaceae 5
Torpedosporales Juncigenaceae 1
Vermiculariopsiellales Vermiculariopsiellaceae 5
Pisorisporiales Pisorisporiaceae 2
Conioscyphales Conioscyphaceae 13
Fuscosporellales Fuscosporellaceae 15
Pleurotheciales Pleurotheciaceae 32
Pseudodactylariales Pseudodactylariaceae 4
Savoryellales Savoryellaceae 34
Sordariomycetes gen-
era incertae sedis Sordariomycetes genera incertae sedis 31
Chaetosphaeriales Chaetosphaeriaceae 131
Helminthosphaeriaceae 10
Linocarpaceae 8
Coniochaetales Coniochaetaceae 8
Cordanales Cordanaceae 5
Phyllachorales Phyllachoraceae 3
Phyllachorales Phyllachorales genera incertae sedis 1
Sordariales Chaetomiaceae 16
Lasiosphaeriaceae 19
Podosporaceae 1
Sordariaceae 1
Sordariales genera incertae sedis 5
Sordariomycetidae
genera incertae sedis Sordariomycetidae genera incertae sedis 3
Amphisphaeriales Amphisphaeriaceae 4
Apiosporaceae 4
Beltraniaceae 7
Pseudomassariaceae 2
Sporocadaceae 7
Xylariales Castanediellaceae 2
Clypeosphaeriaceae 2
Diatrypaceae 1
Hyponectriaceae 7
Hypoxylaceae 2
Microdochiaceae 2
Oxydothidaceae 10
Xylariaceae 5
Vamsapriyaceae 2
Zygosporiaceae 3
Xylariales genera incertae sedis 13
Xylariomycetidae fam-
ily incertae sedis Cainiaceae 1
Myelospermataceae 1
Ascomycota genera
incertae sedis Ascomycota genera
incertae sedis Ascomycota genera incertae sedis 252

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Pezizomycotina gen-
era incertae sedis Pezizomycotina genera
incertae sedis Pezizomycotina genera incertae sedis 2
Total 2968
Classification of freshwater Basidiomycota
Agaricomycetes Agaricales Limnoperdaceae 2
Niaceae 2
Physalacriaceae 2
Agaricales genera incertae sedis 2
Atheliales Atheliaceae 6
Cantharellales Hydnaceae 4
Ceratobasidiaceae 2
Agaricomycetes genera
incertae sedis Agaricomycetes genera incertae sedis 8
Agaricostilbomy-
cetes Agaricostilbales Agaricostilbaceae 1
Atractiellomycetes Atractiellales Phleogenaceae 1
Bartheletiomycetes Polyporales Polyporales genera incertae sedis 1
Sebacinales Sebacinaceae 3
Trechisporales Hydnodontaceae 1
Classiculomycetes Classiculales Classiculaceae 2
Cystobasidiomycetes Cystobasidiales Cystobasidiaceae 7
Erythrobasidiales Erythrobasidiaceae 7
Exobasidiomycetes Doassansiales Doassansiaceae 2
Rhamphosporaceae 1
Exobasidiales Brachybasidiaceae 1
Georgefischeriales Gjaerumiaceae 1
Tilletiariaceae 1
Microstromatales Microstromatales genera incertae sedis 1
Microbotryomycetes Kriegeriales Camptobasidiaceae 1
Kriegeriaceae 1
Leucosporidiales Leucosporidiaceae 6
Microbotryales Ustilentylomataceae 1
Sporidiobolales Sporidiobolaceae 27
Microbotryomycetes
families incertae
sedis
Chrysozymaceae 1
Colacogloeaceae 1
Microbotryomycetes
genera incertae sedis Microbotryomycetes genera incertae sedis 5
Moniliellomycetes Moniliellales Moniliellaceae 1
Tremellomycetes Cystofilobasidiales Cystofilobasidiaceae 5
Mrakiaceae 9
Filobasidiales Filobasidiaceae 15
Holtermanniaceae 1
Piskurozymaceae 4
Tremellales Bulleraceae 1
Bulleribasidiaceae 12
Cryptococcaceae 10
Naemateliaceae 1
Phaeotremellaceae 1
Rhynchogastremaceae 8

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Tremellaceae 1
Trimorphomycetaceae 2
Tremellales genera incertae sedis 1
Trichosporonales Trichosporonaceae 10
Ustilaginomycetes Urocystidales Doassansiopsidaceae 10
Ustilaginales Anthracoideaceae 1
Ustilaginaceae 14
Ustilaginales genera incertae sedis 5
Agaricomycotina
incertae sedis Agaricomycotina
incertae sedis Agaricomycotina incertae sedis 1
Basidiomycota gen-
era incertae sedis Basidiomycota genera
incertae sedis Basidiomycota genera incertae sedis 4
Total 218
Classification of freshwater fungal basal clades
Aphelidiomycota
Aphelidiomycetes Aphelidiales Aphelidiaceae 15
Blastocladiomycota
Blastocladiomycetes Blastocladiales Blastocladiaceae 36
Catenariaceae 3
Paraphysodermataceae 1
Catenomycetales Catenomycetaceae 1
Coelomomycetaceae 5
Blastocladiomycetes
genus incertae sedis Blastocladiomycetes genus incertae sedis 1
Total 47
Chytridiomycota
Chytridiomycetes Chytridiales Asterophlyctaceae 4
Chytridiaceae 35
Chytriomycetaceae 57
Phlyctochytriaceae 21
Pseudorhizidiaceae 1
Scherffeliomycetaceae 1
Chytridiales genus incertae sedis 1
Polyphagales Polyphagaceae 9
Zygophlyctidales Zygophlyctidaceae 4
Zygorhizidiales Zygorhizidiaceae 4
Chytridiomycetes fam-
ily incertae sedis Sparrowiaceae 1
Chytridiomycetes gen-
era incertae sedis Chytridiomycetes genera incertae sedis 43
Cladochytriomycetes Cladochytriales Catenochytridiaceae 2
Cladochytriaceae 15
Endochytriaceae 26
Nowakowskiellaceae 2
Septochytriaceae 2
Cladochytriales genera incertae sedis 4
Lobulomycetes Lobulomycetales Alogomycetaceae 1
Lobulomycetaceae 2
Mesochytriomycetes Gromochytriales Gromochytriaceae 1
Mesochytriales Mesochytriaceae 1
Polychytriomycetes Polychytriales Arkayaceae 1

Fungal Diversity
1 3
Table 1 (continued) Class Order Family Number
of species
Polychytriaceae 7
Rhizophlyctidomy-
cetes Rhizophlyctidales Borealophlyctidaceae 1
Rhizophlyctidaceae 3
Rhizophydiomycetes Rhizophydiales Alphamycetaceae 3
Angulomycetaceae 1
Aquamycetaceae 1
Collimycetaceae 1
Coralloidiomycetaceae 1
Globomycetaceae 2
Gorgonomycetaceae 1
Halomycetaceae 3
Kappamycetaceae 1
Pateramycetaceae 1
Protrudomycetaceae 1
Rhizophydiaceae 52
Staurastromycetaceae 2
Terramycetaceae 2
Spizellomycetes Spizellomycetales Spizellomycetaceae 1
Synchytriomycetes Synchytriales Synchytriaceae 4
Synchytriales genus incertae sedis 4
Total 333
Entomophthoromycota
Entomophthoromy-
cetes Entomophthorales Ancylistaceae 2
Entomophthoraceae 4
Monoblepharomycota
Hyaloraphidiomy-
cetes Hyaloraphidiales Hyaloraphidiaceae 1
Monoblepharidomy-
cetes Monoblepharidales Gonapodyaceae 4
Harpochytriaceae 12
Monoblepharidaceae 10
Oedogoniomycetaceae 1
Telasphaerulaceae 1
Sanchytriomycota
Sanchytriomycetes Sanchytriales Sanchytriaceae 2
Mortierellomycota
Mortierellomycetes Mortierellales Mortierellaceae 5
Mucoromycota
Endogonomycetes Endogonales Endogonaceae 1
Mucoromycetes Mucorales Choanephoraceae 1
Cunninghamellaceae 3
Lichtheimiaceae 1
Mucoraceae 10
Rhizopodaceae 2
Syncephalastraceae 1
Total 19
Olpidiomycota
Olpidiomycetes Olpidiales Olpidiaceae 4

Fungal Diversity
1 3
Dothideomycetes
Freshwater Dothideomycetes is a large taxonomic unit refer-
ring to fungal species having bitunicate asci, often with fis-
situnicate dehiscence, with many collected from freshwater.
The ecology of freshwater fungi has not been systematically
studied due to the variability and uncertainty of the host
habitat, especially the specimens collected from the rainy
or drought seasons. Thus, the numbers of freshwater Doth-
ideomycetes are counted based on the fungi reported from
biodiversity studies and/or single or multi-species taxonomic
studies from freshwater habitats (Dong etal. 2020b). Dong
etal. (2020b) is the key reference for a comprehensive taxo-
nomic account of freshwater Dothideomycetes, including
morphological descriptions, multi-locus phylogenetic analy-
sis, notes with dichotomous keys of taxa, and mapped fresh-
water distributions with references for each species.
According to recent statistics, freshwater dothideomycet-
ous species reside in six orders, 43 families and 145 genera,
while 46 genera are exclusively documented from freshwater
environments (Dong etal. 2020b). In this paper, freshwa-
ter Dothideomycetes are distributed in 18 orders, 75 fami-
lies and 229 genera. Pleosporales is the largest freshwater
order with 391 freshwater species, followed by Tubeufiales
and Jahnulales with 145 and 37 freshwater species, respec-
tively (Table3, Fig.1). Few taxa are reported in Minuti-
sphaerales, Capnodiales, and Kirschsteiniotheliales with 18,
13, and 12 freshwater species, respectively. Representative
Table 1 (continued) Class Order Family Number
of species
Rozellomycota
Microsporidea Amblyosporida Amblyosporidae 49
Caudosporidae 5
Gurleyidae 24
Amblyosporida genera incertae sedis 2
Glugeida Glugeidae 12
Spragueidae 7
Thelohaniidae 10
Unikaryonidae 6
Glugeida genus incertae sedis 1
Neopereziida Berwaldiidae 4
Neopereziidae 12
Neopereziida genera incertae sedis 3
Nosematida Enterocytozoonidae 1
Mrazekiidae 6
Nosematidae 6
Ordosporidae 1
Nosematida genera incertae sedis 4
Microsporidea families
incertae sedis Fibrillasporidae 2
Cougourdellidae 2
Duboscqiidae 3
Microfilidae 2
Neonosemoidiidae 1
Pleistophoridae 10
Toxoglugeidae 1
Tuzetiidae 7
Microsporidea genera
incertae sedis Microsporidea genera incertae sedis 20
Rozellomycota gen-
era incertae sedis Rozellomycota genera
incertae sedis Rozellomycota genera incertae sedis 20
Total 221
Zoopagomycota
Zoopagomycetes Zoopagales Zoopagaceae 3
Overall total 3870

Fungal Diversity
1 3
Dothideomycetes taxa are illustrated in Fig.2. Natipusillales
is the only order in which all species are from freshwater
habitats. In total, 677 dothideomycetous species are reported
in this study from freshwater habitats.
The taxonomic studies on freshwater fungi have greatly
advanced with molecular data resulting in a rapid increase in
fungal numbers (Zhang etal. 2017a; Bao etal. 2018, 2019c,
2020). However, large numbers of fungi are yet to be docu-
mented in poorly studied regions as shown in Dong etal.
(2020b). Furthermore, the identities of some earlier proposed
fungi need to be reappraised with molecular data to confirm
their taxonomical status. Fungal numbers are often related
to the numbers of researchers working on specific fungal
groups. It is a challenge indeed for mycologists and/or tax-
onomists to classify and document this huge number of fungi
before they become extinct.
Eurotiomycetes
Eurotiomycetes was established by Eriksson and Winka
(1997) and presently consists of six subclasses: Chaetothy-
riomycetidae, Coryneliomycetidae, Cryptocaliciomycetidae,
Eurotiomycetidae, Mycocaliciomycetidae, and Sclerococ-
comycetidae (Hibbett etal. 2007; Wood etal. 2016; Geiser
etal. 2017; Réblová etal. 2017; Prieto etal. 2021). The
microcolonies of Eurotiomycetes formed under natural con-
ditions do not have much morphological differences leading
to difficulty morphologically in distinguishing species. The
incorporation of molecular techniques in the fungal taxon-
omy made the classification of fungi more objective, accu-
rate, and comprehensive (Wood etal. 2016; Réblová etal.
2017; Luo etal. 2019; Wijayawardene etal. 2020; Dong
etal. 2020b; Prieto etal. 2021). Wijayawardene etal. (2020)
provided an outline of the classification of the fungi and
fungus-like taxa, and counted all legally published species,
with a total of 3,994 Eurotiomycetes species.
The Eurotiomycetes are medically and industrially important
group (e.g., producers of toxic and potentially useful metabo-
lites, fermentation agents in food and enzymes production,
pathogens of humans and animals) (Alexopoulos etal. 1996;
Jaklitsch etal. 2016; Quan etal. 2020). Eurotiomycetes includes
mostly saprophytic species occurring in various substrates (e.g.,
wood, compost, dung, decaying plant material, foodstuffs, soil),
and some have parasitic and/or commensal associations with
other fungi (Yang etal. 2014, 2018a; Garrido-Benavent and
Pérez-Ortega 2015; Jaklitsch etal. 2016; Le etal. 2020).
Most species of Eurotiomycetes are reported in terrestrial
environments, and freshwater environments account for only
a small part (Liu etal. 2015b; Dong etal. 2020b; Wan etal.
2021) (Fig.3). Currently known freshwater Eurotiomycetes
are mostly saprophytes on submerged wood, decaying leaves,
branches, and plant debris in lakes and streams (Liu etal.
2015b, 2018b; Dong etal. 2018; Wang etal. 2019). Some
taxa grow on inundated stones and isolated from sediments
Table 2 Classification of freshwater Arthoniomycetes with information on the type of habitats, substrates, and geographical distribution of fun-
gal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
ARTHONIOMYCETES
Arthoniales
Arthoniaceae
Arthonia caudata River watershed Canada Maloles etal. (2018)
Arthonia helvola River watershed Canada Maloles etal. (2018)
Arthonia radiata River watershed Canada Maloles etal. (2018)
Chrysotrichaceae
Chrysothrix caesia River watershed Canada Maloles etal. (2018)
Chrysothrix granulosa River Canada Björk etal. (2009)
Lecanographaceae
Alyxoria varia River watershed Canada Maloles etal. (2018)
Opegraphaceae
Cresponea chloroconia River watershed Canada Maloles etal. (2018)
Roccellaceae
Enterographa osagensis Creek Central North America Morse (2013)
Enterographa zonata Waterfall Canada Halda (2011)
Gyrographa gyrocarpa North America Earland-Bennett and Seaward (1974)
Pseudoschismatomma rufescens River watershed Canada Maloles etal. (2018)
Schismatomma cretaceum Nature reserve Ukraine Khodosovtsev etal. (2016)

Fungal Diversity
1 3
in freshwater habitats, and even from drinking water (Iwatsu
etal. 1991; de Hoog etal. 2011; Crous etal. 2013).
Table4 lists 276 species of freshwater Eurotiomycetes
fungi, but this number is undoubtedly underestimated. The
data includes common taxa in freshwater environments, i.e.,
Aspergillus, Penicillium, Thelidium, Verrucaria., and most of
this data comes from cultivable fungi. In this paper, freshwater
Eurotiomycetes are documented from seven orders, thirteen
families and 49 genera. Verrucariales is the largest freshwater
order with 127 freshwater species, followed by Eurotiales and
Chaetothyriales with 76 and 46 freshwater species, respectively
(Table4). Representative freshwater Eurotiomycetes are illus-
trated in Fig.3. The most speciose genus is Verrucaria with
50 freshwater species, followed by Penicillium (47 species),
Thelidium (22 species), Aspergillus (18 species), Exophiala (17
species), and Staurothele (16 species). Many freshwater Euro-
tiomycetes are uncultivable, and some groups are misidentified
without the help of molecular data as morphologically, they
are indistinguishable. In addition, some species will inevitably
be missed in the process of searching the published literature.
Laboulbeniomycetes
The Laboulbeniomycetes includes about 2100 species, most of
which belong to the order Laboulbeniales (Weir and Hammond
1997; Goldmann and Weir 2018). The latter are all obligate
biotrophs associated with arthropods, therefore it is possible
to define “Freshwater Laboulbeniales” as species occurring on
arthropods (which in this case are all insects) that live or spend
most of their time in the aquatic environment. Insect ecology
is not the “questionable” subject of this paper, and we rely on
the definitions contained in a paper dealing with this topic (Jäch
1998). After this clarification, we can report that the “freshwater”
Laboulbeniales total about 258 species and represent the 12% of
the described species in this order of fungi. These 258 species
are found in 25 genera, of which 20 include only aquatic species,
while the other 4 are mainly composed of species that live in
other ecosystems (Table5). Figure4 illustrates Laboulbeniomy-
cetes from freshwater habitats.
These numbers are entirely provisional. Freshwater
Laboulbeniales have received little attention in recent years,
with only 7 species described during the last six years (Rossi
etal. 2016; Das etal. 2018; Santamaria etal. 2020; Boon-
mee etal. 2021; Sundberg etal. 2021). On the other hand,
molecular analysis confirmed the presence of dimorphic
species in the largest genus Chitonomyces (Goldmann and
Weir 2012): which is something that will certainly affect the
taxonomy (and the numbers) of this and other genera.
Leotiomycetes
Leotiomycetes, the most diverse ascomycetous class in Pezi-
zomycotina, encompasses hyphomycetes with undetermined
sexual morphs and apothecial or cleistothecial-bearing fun-
gal members that are either distinctively growing on the
substrates or immersed in stromatic tissues (Berbee 2001;
Baschien etal. 2013; Johnston etal. 2019). Freshwater
Leotiomycetes thrives on various submerged substrates or
floating objects either in static or running waters (Shearer
1993a; Goh and Hyde 1996c; Thomas 1996). Freshwater
Leotiomycetes have been found in various environments:
ponds, rainfall, melting ice (Czeczuga and Orłowska 1999),
lakes (Ingold 1954), springs (Czeczuga etal. 2007), swamps
(Shearer and Crane 1986; Raja etal. 2008b), a polluted lake
(Luo etal. 2004), rivers (Funck etal. 2015), water distri-
bution system (Grabińska-Łoniewska etal. 2007) and on
various substrates: herbaceous and woody substrates (Tsui
etal. 2000; Wong and Hyde 2001), dead cryptogamic plants
(Ingold 1954), scums trapped by woody debris in streams
(Ingold 1974), and isolated from stream spume (Magnes and
Hafellner 1991; Baschien etal. 2013).
In previous studies on fungal discoveries, freshwater
Leotiomycetes were listed along with other classes under
freshwater ascomycetes (Shearer 1993a; Cai etal. 2003d;
Vijaykrishna etal. 2006; Shearer and Raja 2013) or docu-
mented exclusively as discomycetes (Ingold 1954; Kjuller
1960; Beaton and Weste 1976, 1977), as hyphomycetes
(Dudka 1963; Marvanová and Descals 1987; Yu etal. 1988;
Zhu and Yu 1992; Czeczuga and Orłowska 1999; Baschien
etal. 2013), Ingoldian fungi (Ingold 1942; Chan etal. 2000),
and aero-aquatic fungi (Abdullah and Webster 1983; Sri-
indrasutdhi etal. 2015). Studies documenting the overall
freshwater fungal members within Leotiomycetes are lack-
ing (Baschien etal. 2013).
Previous discoveries on new taxa and sexual morph asso-
ciation of freshwater Leotiomycetes were solely based on
morphological observation (Ingold 1942; Beaton and Weste
1977; Muller etal. 1979; Shearer 1993a, b; Webster etal.
1993, 1995). However, the advance of sequencing technol-
ogy introduced deeper morpho-phylogenetic insights on
novel species and holomorph revelation of freshwater Leo-
tiomycetes from various regions and substrates (Belliveau
and Bärlocher 2005; Campbell etal. 2006, 2009; Baschien
etal. 2013; Duarte etal. 2015; Sri-indrasutdhi etal. 2015;
Baudy etal. 2019). Additionally, evolutionary lineages of
freshwater fungi began to unfold. Studies on evolutionary
freshwater fungi based on 18S rRNA data revealed that
terrestrial fungi were the common ancestors of freshwater
Leotiomycetes along with other classes (Dothideomycetes,
Orbiliomycetes, Sordariomycetes) (Belliveau and Bär-
locher 2005; Vijaykrishna etal. 2006). Hence, providing
freshwater fungi with DNA molecular data is important not
only for basic taxonomic justification but also to answer
unsolved evolutionary links within Leotiomycetes. This
study provides a list of freshwater Leotiomycetes including

Fungal Diversity
1 3
Table 3 Classification of freshwater Dothideomycetes with information on the type of habitats, substrates, and geographical distribution of fungal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
DOTHIDEOMYCETES
Asterinales
Morenoinaceae
Morenoina palmicola Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Botryosphaeriales
Botryosphaeriaceae
Dothiorella uruguayensis Submerged decaying wood (stream) China Hu etal. (2013b)
Lasiodiplodia theobromae Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Tiarosporella paludosa Submerged decaying wood (reservoir, river, stream, lake) Australia, Brunei, China, Malaysia,
Philippines Hyde (1993a), Goh and Hyde (1999), Ho
etal. (2001), Cai etal. (2003a), Luo
etal. (2004)
Neodevriesiaceae
Tripospermum camelopardus Foam (rivulet, river, stream), submerged leaf litter of Alnus
glutinosa (stream) Austria, Canada, France, Ireland, UK Ingold etal. (1968a), Bärlocher (1987),
Gessner etal. (1993), Voglmayr (1996),
Harrington (1997), Marvanová (2000)
Tripospermum infalcatum Rainwater: building roofs, needles of Pinus densiflora Japan, Poland Ando and Tubaki (1984a), Czeczuga and
Orłowska (1997)
Tripospermum limneticum Submerged decaying leaves India Patil etal. (2019)
Tripospermum myrti Foam (rivulet, stream), submerged leaf litter of Alnus gluti-
nosa (stream), stemflow and throughfall of tree canopies,
water-filled treeholes
Austria, Brazil, Canada, France, Hun-
gary, India, Malaysia Nawawi (1985b), Bärlocher (1987),
Gessner etal. (1993), Voglmayr (1996),
Marvanová (2000), Gönczöl and Révay
(2003), Gulis and Suberkropp (2003),
Schoenlein-Crusius etal. (2009), Srid-
har and Karamchand (2009)
Tripospermum prolongatum Stemflow and throughfall of tree canopies, submerged
decaying leaves (stream), foam (stream) India, USA Sinclair and Morgan-Jones (1979a), Srid-
har and Karamchand (2009)
Tripospermum sinense Submerged decaying leaves of dicotyledon (stream) China Qiao etal. (2017)
Phyllostictaceae
Guignardia manokwaria Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Capnodiales
Mycosphaerellaceae
Mycosphaerella aspidii Equisetum fluviatile Austria Magnes and Hafellner (1991)
Mycosphaerella equisetina Equisetum fluviatile Austria Magnes and Hafellner (1991)
Mycosphaerella equiseticola Equisetum fluviatile Austria Magnes and Hafellner (1991)
Mycosphaerella perexigua Carex rostrata, Eriophorum angustifolium Austria Magnes and Hafellner (1991)
Stigmidium rivulorum River Bulgaria Shivarov etal. (2018)
Teratosphaeriaceae

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Apenidiella foetida Submerged plant debris (river) Spain Crous etal. (2019)
Apenidiella strumelloidea Leaf of Carex sp. (stagnant water) Russia Quaedvlieg etal. (2014)
Collemopsidiales
Xanthopyreniaceae
Collemopsidium japonicum Nature reserves Japan Harada (1999)
Collemopsidium monense Stream UK Gilbert and Giavarini (1997)
Zwackhiomyces lacustris Stream and lake margin Europe Orange (2002)
Dothideales
Saccotheciaceae
Aureobasidium melanogenum Ground- and tapwater Slovenia Novak Babič etal. (2016)
Aureobasidium pullulans Water (lake, pond), glacial meltwater river Slovakia, Argentina Sláviková etal. (1992), Sláviková and
Vadkertiová (1993), de García etal.
(2012)
Dyfrolomycetales
Pleurotremataceae
Melomastia aquatica Submerged decaying wood (river) China, South Africa Tsui etal. (1998b)
Hysteriales
Hysteriaceae
Gloniopsis calami Submerged decaying wood (pond) Thailand Hyde etal. (2020a)
Hysteriales genus incertae sedis
Graphyllium planispora Scirpus, Typha USA Cavaliere (1975)
Jahnulales
Aliquandostipitaceae
Aliquandostipite crystallinus Submerged decaying wood (river) Costa Rica, Panama, Peru, USA, Ven-
ezuela Raja etal. (2005, 2009b), Shearer etal.
(2015)
Aliquandostipite khaoyaiensis Submerged decaying wood (stream) Costa Rica, USA, Thailand Raja etal. (2005), Raja etal. (2009b),
Dong etal. (2020b),
Aliquandostipite manochiae Submerged decaying wood (lake) Thailand Liu etal. (2015a)
Aliquandostipite minuta Submerged decorticated wood (swamp) USA Raja and Shearer (2007), Raja etal.
(2009b)
Aliquandostipite separans Submerged decaying branches Egypt Pang etal. (2002)
Aliquandostipite siamensiae Submerged decaying wood Thailand, USA Pang etal. (2002), Raja etal. (2009b)
Ascagilis bipolaris Submerged decaying wood (river, stream, waterfall) Australia, China, Costa Rica, Malaysia,
Peru, Thailand, USA Hyde (1992a), Hyde and Wong (1999b),
Tsui etal. (2000), Ho etal. (2001), Tsui
and Hyde (2004), Raja and Shearer
(2006), Hyde (2007), Raja etal.
(2009b), Suetrong etal. (2011), Shearer
etal. (2015)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Ascagilis guttulaspora Submerged decaying wood (stream) China Hyde etal. (2017b)
Ascagilis queenslandica Submerged decaying wood (stream) Australia Hyde etal. (2019)
Ascagilis seychellensis Submerged decaying wood (river) Brazil, Costa Rica, Seychelles Hyde and Wong (1999b), Raja and
Shearer (2006), Barbosa etal. 2013a)
Ascagilis submersa Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Ascagilis sunyatsenii Submerged decaying wood (stream) China Inderbitzin etal. (2001), Pang etal. 2002)
Ascagilis thailandensis Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Brachiosphaera jamaicensis Submerged decaying wood Jamaica, China, Poland Crane and Dumont (1975), Chang (1994)
Brachiosphaera tropicalis Submerged decaying herbaceous, wood (river, stream,
waterfall), foam (river), water Brazil, China, Malaysia, Panama, Peru,
Thailand, USA, Venezuela Descals etal. (1976), Tubaki etal. (1983),
Chang (1994), Cai etal. (2002b),
Campbell etal. (2007). Smits etal.
(2007), Raja etal. (2009b), Suetrong
etal. (2011), Shearer etal. (2015)
Jahnula apiospora Submerged decorticated wood (creek) Canada Raja etal. (2008a)
Jahnula appendiculata Submerged decaying wood (e.g. (Azadirachta indica var.
siamensis, Erythrophleum teysmannii, Melaleuca cajuputi,
Shorea obtusa, S. roxburghii, Wrightia tomentosa, Xylia
xylocarpa, Zollingeria dongnaiensis (river), Licuala longi-
calycata and Eleiodoxa conferta (peat swamp)
Peru, Thailand Pinruan etal. (2002), Pinnoi etal. (2006),
Pinruan etal. (2007), Sivichai and
Boonyuen (2010), Shearer etal. (2015)
Jahnula aquatica Submerged decaying wood (e.g., Salix sp.) (creek, lake,
stream, waterfall) China, France, Germany, Malaysia,
South Africa, Thailand, USA Kirschstein (1936), Hawksworth (1984),
Ho (1998), Hyde and Wong (1999b),
Ho etal. (2001), Raja and Shearer
(2006), Raja etal. (2009b), Fournier
etal. (2010), Sivichai etal. (2011)
Jahnula bipileata Submerged decorticated wood (lake) Ecuador, USA Raja and Shearer (2006)
Jahnula granulosa Submerged decaying wood (lake) China, South Africa, Thailand Hyde and Wong (1999b), Cai etal.
(2002b), Suetrong etal. (2011)
Jahnula morakotii Submerged Azadirachta indica test block (peat swamp) Thailand Sivichai and Boonyuen (2010)
Jahnula poonythii Submerged decaying wood (lake) China, Mauritius, Mexico Hyde and Wong (1999b), Cai etal.
(2002b), Luo etal. (2004), González
and Chavarría (2005)
Jahnula purpurea Submerged decorticated branch (river) Martinique Fournier etal. (2015)
Jahnula rostrata Submerged decaying wood (lake, stream) China, USA Raja and Shearer (2006), Raja etal.
(2009b), Dong etal. (2020b),
Jahnula sangamonensis Submerged decorticated wood (river) USA Raja and Shearer (2006), Raja etal.
(2009b)
Jahnula seychellensis Submerged decaying wood and twig (stream, river) Brazil, Philippines, Seychelles Hyde and Wong (1999b), Cai etal.
(2003a), Barbosa etal. (2013b)
Jahnula systyla Submerged decaying wood (river) Australia, China, Malaysia Ho (1998), Hyde and Wong (1999b), Ho
etal. (2001)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Megalohypha aqua-dulces Submerged decaying wood (river) Panama, Thailand Ferrer etal. (2007)
Neojahnula australiensis Submerged decaying wood (river) Australia, Brunei, China, Peru, Thai-
land, USA Hyde (1993b), Ho etal. (2001), Raja etal.
(2009b), Suetrong etal. (2011), Shearer
etal. (2015), Dong etal. (2020b)
Pseudojahnula potamophila Submerged decaying wood (river) Australia, Costa Rica, USA Hyde and Wong (1999b), Raja and
Shearer (2006), Raja etal. (2009b),
Xylomyces acerosisporus Submerged decaying leaves (river) Brazil Oliveira etal. (2015b)
Xylomyces aquaticus Submerged decaying woods (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) and leaves, foam (stream),
water (river), submerged decaying Licuala longicaly-
cata and Eleiodoxa conferta (peat swamp), water-filled
treeholes
Austria, Brazil, Hungary, Poland, Spain,
Thailand, Ukraine, UK, USA Dudka (1966), Czeczuga etal. (1990),
Hyde and Goh (1999a), Marvanová
(2000), Sivichai etal. (2002), Gönczöl
and Révay (2003), Pinnoi etal. (2006),
Pinruan etal. (2007), Barbosa etal.
(2013b), Hernández-Restrepo etal.
(2017)
Xylomyces chlamydosporus Submerged decaying wood (lake, river, stream, waterfall) Australia, Brazil, Brunei, China, Sey-
chelles, Thailand, USA Goos etal. (1977), Lamore and Goos
(1978), Goh etal. (1997a), Hyde and
Goh (1997), Hyde and Goh (1998c),
Tsui etal. (2001d), Fryar etal. (2004),
Luo etal. (2004), Raja etal. (2009b),
Kurniawati etal. (2010), Barbosa etal.
(2013b)
Xylomyces foliicola Submerged decaying wood and leaves (e.g., Calophyllum
brasiliense) (stream) Brazil Barbosa etal. (2013b), Fiuza etal. 2018)
Xylomyces giganteus Submerged decaying wood (river) Australia, China, South Africa Goh etal. (1997a), Hyde etal. (1998a),
Jiang etal. (2008)
Xylomyces punctatus Submerged decaying wood (river) China Goh etal. (1997a), Tsui etal. (2001d)
Xylomyces pusillus Submerged decaying wood and leaf (lake, river) Brazil, China Goh etal. (1997a), Tsui etal. (2001d),
Cai etal. (2002b), da Silva and Gusmão
(2015)
Kirschsteiniotheliales
Kirschsteiniotheliaceae
Kirschsteiniothelia aquatica Submerged decaying wood (stream) China Barbosa etal. (2013b), Bao etal. (2018)
Kirschsteiniothelia atra Submerged decaying wood Brazil, China Su etal. (2016a)
Kirschsteiniothelia cangshanensis Submerged decaying wood (river) China Bao etal. (2018)
Kirschsteiniothelia fluminicola Submerged decaying wood (river) China Bao etal. (2018)
Kirschsteiniothelia rostrata Submerged decaying wood (stream) China, Thailand Hyde etal. (2017b), Bao etal. (2018)
Kirschsteiniothelia submersa Submerged decaying wood (stream) China Su etal. (2016a)
Kirschsteiniotheliales genera incertae sedis

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Brachysporiella gayana Submerged decaying wood (reservoir, stream, waterfall),
submerged decaying Eleiodoxa conferta (peat swamp) Brazil, China, Thailand Goh and Hyde (1999), Pinnoi etal.
(2006), Kurniawati etal. (2010), Bar-
bosa etal. (2013b)
Brachysporiella pulchra Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Brachysporiella rhizoidea Submerged decaying Licuala longicalycata and Eleiodoxa
conferta (peat swamp) China, Thailand Pinruan etal. (2007), Tsui etal. (2001b),
Pinnoi etal. (2006), Ranghoo etal.
(1999), Tsui etal. (2001b)
Brachysporiella setosa Submerged decaying wood (stream) Brunei, China Ho etal. (2001)
Taeniolella longissima Submerged Scots pine test block (water-cooling tower) UK Jones etal. (2002)
Taeniolella phialosperma Sediment Korea Goh etal. (2021a)
Lichenotheliales
Lichenotheliaceae
Endococcus rugulosus Stream and lake margin Europe Orange (2002)
Endococcus verrucisporus Stream and lake margin Europe Orange (2002)
Lichenotheliaceae
Endococcus rugulosus Stream and lake margin Europe Orange (2002)
Monoblastiales
Monoblastiaceae
Acrocordia cavata River watershed Canada Maloles etal. (2018)
Microthyriales
Microthyriales genera incertae sedis
Heliocephala elegans Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Minutisphaerales
Acrogenosporaceae
Acrogenospora aquatica Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora basalicellularispora Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora ellipsoidea Submerged decaying wood (stream) China Hu etal. (2010a)
Acrogenospora guttulatispora Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora obovoidspora Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora olivaceospora Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora ovalis Submerged decaying wood (reservoir) China Goh etal. (1998c)
Acrogenospora sphaerocephala Submerged decaying wood (river, stream, waterfall),
bamboo, leaves (stream), decaying Metroxylon sagu (peat
swamp), submerged Phragmites (river)
Brazil, Brunei, China, Philippines,
Seychelles, South Africa, Thailand,
USA, UK
Lamore and Goos (1978), Shearer and
Crane (1986), Hyde and Goh (1998c),
Goh and Hyde (1999), Sivichai etal.
(2000b), Tsui etal. (2000), Cai etal.
(2003a), Barbosa etal. (2013b), Hyde
etal. (2019)
Acrogenospora submersa Submerged decaying wood (river) China Bao etal. (2020)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Acrogenospora subprolata Submerged decaying wood (river, stream) China Goh etal. (1998c), Tsui etal. (2001d),
Bao etal. (2020)
Acrogenospora thailandica Submerged decaying wood (stream) Thailand Hyde etal. (2019)
Acrogenospora verrucispora Submerged decaying wood (stream) China Bao etal. (2020)
Acrogenospora yunnanensis Submerged decaying wood (stream) China Bao etal. (2020)
Minutisphaeraceae
Minutisphaera aquaticum Submerged decorticated wood river) Thailand Bao etal. (2019a)
Minutisphaera aspera Submerged decorticated wood (stream) USA Raja etal. (2015)
Minutisphaera fimbriatispora Submerged decorticated wood (pond) USA Ferrer etal. (2011), Raja etal. (2013b)
Minutisphaera japonica Submerged decorticated Fraxinus excelsior, Salix sp.
(brook) Japan, France Raja etal. (2013b, 2015)
Minutisphaera parafimbriatispora Submerged decorticated wood (swamp) USA Raja etal. (2015)
Monoblastiales
Monoblastiaceae
Acrocordia cavata River watershed Canada Maloles etal. (2018)
Natipusillales
Natipusillaceae
Natipusilla bellaspora Submerged decaying wood (river, stream) Peru Raja etal. (2012b), Shearer etal. (2015)
Natipusilla decorospora Submerged corticated wood (swamp) Ecuador Ferrer etal. (2011)
Natipusilla limonensis Submerged decaying wood (river, stream) Costa Rica, Peru Ferrer etal. (2011), Shearer etal. (2015)
Natipusilla naponensis Submerged corticated wood (swamp) Ecuador Ferrer etal. (2011)
Pleosporales
Acrocalymmaceae
Acrocalymma aquaticum Submerged decaying wood (stream) Thailand Zhang etal. (2012a)
Acrocalymma bipolare Submerged decaying wood (river) Egypt Dong etal. (2020b)
Acrocalymma medicaginis Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Aigialaceae
Fissuroma neoaggregatum Submerged decaying wood Thailand Dong etal. (2020b)
Neoastrosphaeriella aquatica Submerged decaying wood Thailand Bao etal. (2019b)
Amniculicolaceae
Amniculicola aquatica Submerged decaying wood China Hyde etal. (2019)
Amniculicola guttulata Submerged decaying wood China Hyde etal. (2019), Dong etal. (2020b)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Amniculicola longissima Submerged decaying wood (Alnus glutinosa, Acer pseu-
doplatanus, Corylus avellana, Fraxinus excelsior, Fagus
sylvatica, Ulmus sp.) (stream), foam (stream, spring, river,
lake), submerged decaying leaves of Salix spp., Populus
spp., Betula glandulosa, Alnus tenuifolia, A. glutinosa
(stream, river), water (river, stream), scum (stream),
sediment (dam, stream), stemflow and throughfall of tree
canopies and palm species
Austria, Brazil, Canada, Czech Repub-
lic, France, Germany, India, Ireland,
Jamaica, Nigeria, Poland, UK, USA,
Venezuela
Ingold (1942), Ingold (1943a), Ranzoni
(1953), Hudson and Ingold (1960)ngold
(1960, Baxter (1960), Ingold (1960),
Petersen (1962), Crane (1968), Alasoa-
dura (1968a), Iqbal and Webster (1973),
Abdullah and Webster (1980), Bär-
locher (1987), Czeczuga etal. (1990),
Gessner etal. (1993), Voglmayr (1996),
Harrington (1997), Marvanová (2000),
Gulis and Suberkropp (2003), Rajashek-
har and Kaveriappa (2003), Smits
etal. (2007), Baschien etal. (2008),
Schoenlein-Crusius etal. (2009), Srid-
har and Karamchand (2009), Sudheep
and Sridhar (2012), Fiuza and Gusmão
(2013a)
Amniculicola immersa Submerged Salix sp. wood Denmark Zhang etal. (2009a)
Amniculicola lignicola Partly submerged wood of Fraxinus excelsior France Zhang etal. (2008b)
Amniculicola parva Submerged wood of Alnus glutinosa France Zhang etal. (2009a)
Anguillospora crassa Scum (stream), foam (stream, river), submerged decaying
leaves of Alnus glutinosa, A. tenuifolia Salix spp., Populus
spp., Betula glandulosa (stream), sediment (dam), stem-
flow and throughfall of tree canopies, water (river, stream)
Austria, Brazil. Canada, Czech Repub-
lic, France, Germany, India, Ireland,
Japan, Jamaica, Poland, Portugal,
Slovakia, UK, USA Venezuela
Ingold (1958a), Dixon (1959), Hudson
and Ingold (1960), Baxter (1960),
Ingold (1960), Petersen (1962), Anasta-
siou (1964), Bärlocher (1987), Gessner
etal. (1993), Sridhar and Bärlocher
(1993), Voglmayr (1996), Harrington
(1997), Marvanová (2000), Gulis and
Suberkropp (2003), Rajashekhar and
Kaveriappa (2003), Belliveau and
Bärlocher (2005), Hosoya and Tanaka
(2007), Smits etal. (2007), Baschien
etal. (2008), Schoenlein-Crusius
etal. (2009), Sridhar and Karamchand
(2009), Sudheep and Sridhar (2012),
Ghate and Sridhar (2015), Raposeiro
etal. (2018), Pietryczuk etal. (2018),
Johnston and Baschien 2020)
Anguillospora curvula Submerge decayed leaves of Poaceae (river), foam (river) Ireland, UK Iqbal (1972a), Harrington (1997)
Anguillospora filiformis Foam (stream), submerged decaying leaves (stream), water
(river, stream) Australia, Austria, Brazil, Canada,
Italy, Poland, Portugal, Spain¸ USA,
Venezuela
Sridhar and Bärlocher (1993), Marvanová
(2000), Gulis and Suberkropp (2003),
Belliveau and Bärlocher (2005), Smits
etal. (2007), Schoenlein-Crusius etal.
(2009), Duarte etal. (2012), Pietryczuk
etal. (2018)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Anguillospora furtiva Foam (stream, river) Austria, Ireland, Slovakia, UK Harrington (1997), Marvanová (2000),
Belliveau and Bärlocher (2005),
Baschien etal. (2008)
Anguillospora gigantea Water (river, spring), decaying unidentified leaves (spring),
submerged decaying leaves of Populus sp. (irrigation
ditch)
Cuba, Poland, USA Ranzoni (1953), Petersen (1962),
Marvanová and Marvan (1969), Czec-
zuga etal. (1990)
Anguillospora mediocris Submerged decaying leaves of Alnus glutinosa (stream) Hungary Gönczöl and Marvanová (2002), Bel-
liveau and Bärlocher (2005)
Anguillospora pseudolongissima Submerged decaying leaves of Salix (rivulet), foam (river) Brazil, USA Ranzoni (1953), Petersen (1962), Fiuza
and Gusmão (2013a)
Anguillospora pulchella Foam USA Parker and Roane (1976)
Anguillospora rosea Foam (stream, river, waterfalls), apothecium of Orbilia on
submerged branch of tree, submerged decorticated twigs Austria, Ireland, UK Harrington (1997), Descals etal. (1998),
Marvanová (2000), Belliveau and Bär-
locher (2005)
Anguillospora rubescens Submerged decaying grass blades (river) Belarus Gulis and Marvanová (1999)
Anguillospora virginiana Foam USA Parker and Roane (1976)
Fouskomenomyces cupreorufescens Submerged decaying leaves of Alnus glutinosa (oligotropic
peat bog) Austria Voglmayr (2004), Magaña-Dueñas etal.
(2020)
Fouskomenomyces mimiticus Submerged decaying plant debris Spain Magaña-Dueñas etal. (2020)
Murispora aquatica Submerged decaying wood (stream) China Bao etal. (2019d)
Murispora cicognanii Submerged decaying wood (stream) China Hyde etal. (2019)
Murispora fagicola Submerged decaying wood (river) China Bao etal. (2019d)
Murispora fissilispora Submerged decaying plant debris Spain Magaña-Dueñas etal. (2020)
Murispora asexualis Submerged decaying plant debris Spain Magaña-Dueñas etal. (2020)
Murispora navicularispora Submerged decaying plant debris Spain Magaña-Dueñas etal. (2020)
Murispora rubicunda Submerged dead herbaceous stem (Dipsacus?) Austria, France Magnes and Hafellner (1991), Zhang
etal. (2012b)
Neomassariosphaeria typhicola Submerged culm of Phragmites and stems of Typha latifolia
(lake), submerged decaying branches Denmark, Pakistan, USA Fallah and Shearer (2001), Zhang etal.
(2009b), Tanaka etal. (2010)
Anteagloniaceae
Flammeascoma bambusae Submerged bamboo stem Thailand Liu etal. (2015a)
Purpureofaciens aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Aquasubmersaceae
Aquasubmersa japonica Submerged decaying wood (river) Japan Ariyawansa etal. (2015)
Aquasubmersa mircensis Submerged decaying wood (stream) Thailand Zhang etal. (2012a)
Astrosphaeriellaceae
Aquatospora cylindrica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Astrosphaeriella angustispora Submerged decaying Licuala longicalycata and Eleiodoxa
conferta (peat swamp) Thailand Pinnoi etal. (2006), Pinruan etal. (2007)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Astrosphaeriella aquatica Submerged rachides of Livistona (river), submerged decay-
ing Eleiodoxa conferta and Licuala longicalycata (peat
swamp)
Papua New Guinea, Ecuador, Thailand Hyde (1994a), Hyde and Fröhlich (1998),
Pinruan etal. (2014)
Astrosphaeriella exorrhiza Submerged roots of Iriartea (stream) Ecuador Hyde and Fröhlich (1998)
Astrosphaeriella floridana Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Astrosphaeriella frondicola Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Astrosphaeriella lophiostomopsis Decaying Licuala longicalycata and Metroxylon sagu (peat
swamp) Thailand Pinruan etal. (2014)
Astrosphaeriella malayensis Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Astrosphaeriella maquilingiana Submerged decaying wood (stream) Australia Vijaykrishna and Hyde (2006)
Astrosphaeriella papuana Submerged decaying wood and bamboo (river) Papua New Guinea, Philippines Hyde and Fröhlich (1998), Cai etal.
(2003a)
Astrosphaeriella seychellensis Submerged decaying wood (river) Seychelles Hyde and Goh (1998c)
Astrosphaeriella stellata Submerged decaying wood and bamboo (lake, river, stream,
waterfall), decaying culms of Thysanolaena maxima China, Philippines, Thailand Tsui etal. (2001f), Wong and Hyde
(2001), Cai etal. (2003a), Luo etal.
(2004), Shearer etal. (2007), Kurni-
awati etal. (2010)
Astrosphaeriella thailandensis Submerged decaying wood (stream) Thailand Ren etal. (2013)
Caryospora aquatica Submerged decaying wood (stream, waterfall) China, Thailand Ariyawansa etal. (2015), Dong etal.
(2020b)
Caryospora minima Submerged decaying wood and bamboo (lake, stream,
reservoir) China, Philippines, South Africa Hyde etal. (1998a), Goh and Hyde
(1999), Tsui etal. (2000), Cai etal.
(2003a), Luo etal. (2004)
Caryospora obclavata Submerged decorticated woody debris USA Raja and Shearer (2008)
Caryospora quercus Submerged decaying wood Thailand Dong etal. (2020b)
Caryospora submersa Submerged decaying wood Thailand Dong etal. (2020b)
Pithomyces elaeidicola Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Pithomyces flavus Submerged decaying wood (lake) China Cai etal. (2002b), Hu etal. (2013a)
Pithomyces niger Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Pithomyces obscuriseptatus Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Xenoastrosphaeriella tornata Submerged decaying wood and bamboo (river), decaying
Metroxylon sagu (peat swamp) Philippines, Thailand Cai etal. (2003a), Pinruan etal. (2014)
Xenoastrosphaeriella trochus Submerged decaying wood (stream) Australia, China, South Africa Tsui etal. (2000), Cai etal. (2002b),
Vijaykrishna and Hyde (2006)
Bambusicolaceae
Bambusicola aquatica Submerged decaying wood (stream) China Dong etal. (2020b)
Corynesporascaceae
Corynespora aquatica Submerged decaying leaves (stream) Mexico Castañeda-Ruiz etal. (2004)
Corynespora cubensis Rainwater from building roofs Poland Czeczuga and Orłowska (1997)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Corynespora lignicola Submerged decaying wood (river) China Hyde etal. (2020a)
Corynespora submersa Submerged decaying wood (river) China Hyde etal. (2020a)
Cucurbitariaceae
Neocucurbitaria aquadulcis Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Neocucurbitaria variabilis Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Cyclothyriellaceae
Massariosphaeria scirpina Typha leaves USA Cavaliere (1975)
Massariosphaeria websteri Submerged decaying culms of cf. Phragmites karka Pakistan Tanaka etal. (2010)
Delitschiaceae
Delitschia bispora Water-cooling towers UK Eaton and Jones (1970, 1971b), Eaton
(1972)
Delitschia fasciatispora Submerged branch (river) Mauritius Hyde and Steinke (1996)
Delitschia palmietensis Submerged bark (river) Mauritius Hyde and Steinke (1996)
Dictyosporiaceae
Aquadictyospora lignicola Submerged decaying wood (stream) China Li etal. (2017)
Aquaticheirospora lignicola Submerged decaying wood (stream) Thailand Kodsueb etal. (2007)
Cheirosporium triseriale Submerged decaying wood (stream) China Cai etal. (2008)
Cheirosporium vesiculare Submerged decaying wood (river) Egypt Abdel-Aziz (2016a)
Dendryphiella vinosa Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Dictyocheirospora aquadulcis Submerged decaying wood (stream) Thailand Hyde etal. (2019)
Dictyocheirospora aquatica Submerged decaying wood (river) China Wang etal. (2016)
Dictyocheirospora garethjonesii Submerged decaying wood (river) China Wang etal. (2016)
Dictyocheirospora gigantica Submerged decaying wood (e.g., submerged test block of
Anisoptera oblonga) (river, stream) South Africa, Thailand Goh etal. (1999)
Dictyocheirospora heptaspora Submerged decaying wood (lake) and bamboo (river) China, Philippines Cai etal. (2003a), Luo etal. (2004)
Dictyocheirospora indica Submerged decaying wood (stream) Thailand Yang etal. (2018b)
Dictyocheirospora musae Submerged decaying wood (lake) China Luo etal. (2004)
Dictyocheirospora rotunda Submerged decaying wood (river, stream) China, Thailand Boonmee etal. (2016), Wang etal.
(2016), Dong etal. (2020b)
Dictyocheirospora tetraploides Submerged decaying wood (stream) China Cai etal. (2003b)
Dictyocheirospora thailandica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Dictyocheirospora vinaya Submerged decaying wood (stream) China, Thailand Boonmee etal. (2016), Fu etal. (2021)
Dictyosporium alatum Submerged decaying wood (stream), decaying Metroxylon
sagu (peat swamp) Brunei, Thailand Ho etal. (2001), Pinruan etal. (2014)
Dictyosporium aquaticum Submerged decaying date palm (river) Egypt Liu etal. (2015a)
Dictyosporium biseriale Submerged decaying wood (stream) China Hu etal. (2010a)
Dictyosporium brahmaswaroopii Submerged decaying wood (river) China Goh etal. (1999)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Dictyosporium canisporum Submerged decaying wood (stream) China Cai etal. (2003b)
Dictyosporium digitatum Submerged decaying wood (e.g., Anisoptera oblonga, Dip-
terocarpus alatus, Machilus velutina, Pinus massoniana,
Xylia dolabriformis test blocks) (reservoir, river, stream)
and leaves (stream)
Australia, Brazil, Brunei, China, Thai-
land Goh etal. (1999), Goh and Hyde (1999)
Tsui etal. (2000), Tsui etal. (2001e),
Sivichai etal. (2002)
Dictyosporium elegans Submerged decaying leaves and petiole (stream), submerged
test blocks of Anisoptera oblonga, Dipterocarpus alatus,
Machilus velutina, Pinus massoniana (stream)
Brazil, China, Thailand Ho etal. (2001), Sivichai etal. (2002),
Barbosa etal. (2013b)
Dictyosporium lakefuxianense Submerged decaying wood (lake) China Cai etal. (2003c), Luo etal. (2004)
Dictyosporium muriformis Submerged decaying wood (stream) China Hyde etal. (2020b)
Dictyosporium nigroapice Submerged decaying wood (e.g., Dipterocarpus alatus test
block) (stream) China, Thailand Goh etal. (1999), Sivichai etal. (2002),
Yang etal. (2018a)
Dictyosporium olivaceosporum Submerged decaying twigs (stream) Japan Boonmee etal. (2016)
Dictyosporium palmae Submerged decaying fronds of Phoenix dactylifera (river) Egypt Abdel-Aziz (2016a)
Dictyosporium pelagicum Submerged decaying wood (river) USA Shearer (1972)
Dictyosporium polystichum Submerged decaying wood (lake) China Luo etal. (2004)
Dictyosporium stellatum Submerged decaying wood (lake) Ontario Crous etal. (2011)
Dictyosporium subramanianii Submerged decaying wood (e.g., Dipterocarpus alatus test
block) (river, stream, waterfall) Thailand Sivichai etal. (2002), Kurniawati etal.
(2010)
Dictyosporium tetraseriale Submerged decaying wood (e.g., Machilus velutina, Pinus
massonaina wood baits) (stream) Brunei, China Goh etal. (1999)
Dictyosporium tetrasporum Submerged decaying wood (stream) China Cai and Hyde (2007a)
Dictyosporium thailandicum Submerged decaying wood (stream) Thailand Liu etal. (2015a)
Dictyosporium tratense Submerged decaying wood (stream) Thailand Yang etal. (2018b)
Dictyosporium triramosum Submerged decaying stem (river) Buenos Aires Arambarri (2001)
Dictyosporium tubulatum Submerged decaying wood (stream) Thailand Yang etal. (2018b)
Dictyosporium yunnanense Submerged decaying wood China Cai etal. (2003c)
Dictyosporium zhejiangense Submerged decaying wood (stream) China Wongsawas etal. (2009b)
Digitodesmium bambusicola Submerged bamboo culms (river) Philippines Cai etal. (2002c)
Digitodesmium heptasporum Submerged decaying wood (stream) China Cai etal. (2003b)
Digitodesmium recurvum Submerged decaying wood (stream) China Ho etal. (1999a)
Jalapriya apicalivaginatum Submerged decaying wood (stream) China Fu etal. (2021)
Jalapriya aquatica Submerged decaying wood (stream) China Fu etal. (2021)
Jalapriya pulchra Submerged decaying wood (stream) China Boonmee etal. (2016)
Pseudocoleophoma typhicola Submerged stems of Typha latifolia (lake) UK Hyde etal. (2016b)
Pseudodictyosporium wauense Submerged decaying wood (stream) China Li etal. (2017)
Vikalpa lignicola Submerged decaying wood (stream) China Boonmee etal. (2016)
Didymellaceae

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Calophoma complanata Submerged decaying wood (stream) China Ho etal. (2001)
Didymella castillejae Submerged decaying wood (river) Malaysia Ho etal. (2001), Hyde (2007)
Didymella glacialis Carex rostrata Austria Magnes and Hafellner (1991)
Didymella hellebori Submerged decaying wood (river) Malaysia Hyde (2007)
Microsphaeropsis olivacea Submerged decaying wood (stream) Brunei Ho etal. (2001)
Nothophoma spiraeae Submerged plant litter Korea Goh etal. (2021a)
Didymosphaeriaceae
Didymosphaeria bisphaerica Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Didymosphaeria pittospora Submerged beech test blocks in storage tank and water in
cooling tower India Udaiyan and Hosagoudar (1991)
Paraphaeosphaeria michotii Typha leaves UK Pugh and Mulder (1971)
Paraphaeosphaeria schoenoplecti Senescent culms of Schoenoplectus litoralis China Wong etal. (2000)
Paraconiothyrium fuckelii Submerged decaying wood USA Shearer (1972)
Pseudotrichia allequashensis Submerged, partially decomposed grass stem USA Fallah and Shearer (2001)
Dothidotthiaceae
Mycocentrospora acerina Foam (lake, spring, stream, river), submerged decaying
leaves of Alnus sp. (stream), stemflow and throughfall of
tree canopies
Austria, Brazil, Canada, India, Spain,
Switzerland, USA, Venezuela Ingold (1949), Anastasiou (1964), Crane
(1968), Bärlocher (1987), Marvanová
(2000), Smits etal. (2007), Schoenlein-
Crusius etal. (2009), Sridhar and Kara-
mchand (2009), Menéndez etal. (2012)
Mycocentrospora angulata Submerged decaying leaves of Prunus and unidentified
angiosperm leaves, foam (stream, lake, spring, river) USA Petersen (1962), Crane (1968)
Mycocentrospora aquatica Foam (stream), submerged decaying wood and
leaves(stream) Canada, UK Iqbal (1972a), Bärlocher (1987), Sridhar
and Bärlocher (1993)
Mycocentrospora clavata Submerged decaying leaves (stream) Canada Sridhar and Bärlocher (1993)
Mycocentrospora varians Submerged decaying leaves (stream) USA Sinclair and Morgan-Jones (1979a)
Hermatomycetaceae
Hermatomyces sphaericus Submerged decaying stream (stream) Brazil Barbosa etal. (2013b)
Latoruaceae
Pseudoasteromassaria aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Pseudoasteromassaria spadicea Submerged decaying wood (stream) Thailand Tibpromma etal. (2017)
Lentitheciaceae
Halobyssothecium bambusicola Sumerged decaying wood (stream) Thailand Calabon etal. (2021b)
Halobyssothecium cangshanense Submerged decaying wood (stream) China Su etal. (2016b)
Halobyssothecium carbonneanum Submerged decaying branch of Populus sp. (lake) France Crous etal. (2018)
Halobyssothecium kunmingense Submerged decaying wood (stream) China Dong etal. (2020b)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Halobyssothecium obiones Phragmites australis culms and stems, Typha leaves (river,
swamps) UK, USA Pugh and Mulder (1971), Apinis etal.
(1972a), Taligoola etal. (1972), Shearer
and Crane (1986)
Halobyssothecium unicellulare Submerged decaying wood (river) Egypt Hyde etal. (2016b)
Keissleriella linearis Submerged dead stems of Phragmites sp. (lake) France Zhang etal. (2009c)
Lentithecium clioninum Submerged decaying wood (river) Japan Tanaka etal. (2005b)
Lentithecium pseudoclioninum Submerged twigs of woody plant (pond) Japan Tanaka etal. (2015)
Poaceascoma aquaticum Submerged decaying bamboo (stream) Thailand Luo etal. (2016a)
Setoseptoria arundinacea Submerged decaying wood, submerged stems of Phragmites
sp. (lake) China, Denmark, UK Apinis etal. (1972a), Luo etal. (2004),
Zhang etal. (2009c)
Setoseptoria magniarundinacea Submerged dead herbaceous stems (river) Japan Tanaka etal. (2004)
Tingoldiago clavata Submerged decaying wood Thailand Xu etal. (2020)
Tingoldiago hydei Submerged decaying wood Thailand Xu etal. (2020)
Tingoldiago graminicola Submerged culms of Phragmites australis and P. japonica
(river) Japan, UK Hirayama etal. (2010)
Leptosphaeriaceae
Leptosphaeria acuta Scirpus, Typha (lake) USA Cavaliere (1975)
Leptosphaeria albopunctata Submerged decaying wood (river), Phragmites australis
stems and leaves USA, UK Lamore and Goos (1978), Dudka (1963),
Apinis etal. (1972b), Taligoola etal.
(1972), Lamore and Goos (1978)
Leptosphaeria culmifraga Phragmites australis culms UK Apinis etal. (1972a)
Leptosphaeria eustomoides Phragmites australis leaves UK Apinis etal. (1972b)
Leptosphaeria modesta Typha leaves UK Pugh and Mulder (1971)
Leptosphaeria sepalorum Juncus filiformis (lake) Austria Magnes and Hafellner (1991)
Lindgomycetaceae
Aquimassariosphaeria kunmingensis Submerged decaying wood (stream) China Dong etal. (2020b)
Aquimassariosphaeria typhicola Submerged, partially decomposed stems of Typha latifolia
and unidentified grass stem (lake) USA Fallah and Shearer (2001)
Arundellina typhae Submerged dead stem of Typha sp. (river) UK Hyde etal. (2016b)
Clohesyomyces aquaticus Submerged decaying wood and bamboo (river, stream) Australia, China, Thailand Hyde (1993a), Cai etal. 2006, Zhang
etal. (2012a), Dong etal. (2020b)
Hongkongmyces aquaticus Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Hongkongmyces brunneisporus Submerged decaying wood (stream, lake) China Bao etal. (2021b)
Hongkongmyces pedis Freshwater (river) Korea Pangging etal. (2021b)
Hongkongmyces snookiorum Submerged detritus (fen) Korea, USA Crous etal. (2018), Pangging etal.
(2021b)
Lindgomyces angustiascus Submerged decorticated wood (lake) USA Raja etal. (2013a)
Lindgomyces apiculatus Submerged twigs of woody plant (pond) Japan Raja etal. (2011)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Lindgomyces aquaticus Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Lindgomyces breviappendiculatus Submerged decaying wood (river) Japan Tanaka etal. (2005b), Hirayama etal.
(2010)
Lindgomyces carolinensis Submerged decaying wood (lake) USA Raja etal. (2017)
Lindgomyces cigarosporus Submerged decaying wood (lake) USA Raja etal. (2017)
Lindgomyces cinctosporus Submerged decaying wood of Salix sp. and branchlet of
Fagus sylvatica (brook, stream) France, Spain, USA Hirayama etal. (2010), Zhang etal.
(2014d)
Lindgomyces griseosporus Submerged wood and twig of Alnus glutinosa, Fraxinus
excelsior, Rhamnus cathartica, Frangula alnus, Salix sp.,
and submerged dead root of Platanus sp. (stream)
France, Spain Zhang etal. (2014d)
Lindgomyces ingoldianus Submerged woody debris (e.g., Dipterocarpus alatus,
Platanus sp., Xylia dolabriformis test blocks) (reservoir,
stream), submerged herbaceous stem (Scirpus brevicauda-
tus, S. validus, S. atrovirens, Typha latifolia) (lake)
Australia, Brunei, China, France, Japan,
Malaysia, Thailand, USA Shearer and Hyde (1997), Hyde and Apt-
root (1998a), Tsui etal. (2000), Sivichai
etal. (2002), Hirayama etal. (2010),
Goh and Hyde (1999) Fallah and
Shearer (2001), Zhang etal. (2014d)
Lindgomyces lemonweirensis Submerged, decorticated wood (river) USA Raja etal. (2011)
Lindgomyces madisonensis Submerged decaying wood (creek) USA Crous etal. (2015)
Lindgomyces okinawaensis Submerged dead twigs (river) Japan Li etal. (2016)
Lindgomyces pseudomadisonensis Submerged dead twigs of woody plant (river) Japan Hyde etal. (2016b)
Lindgomyces rotundatus Submerged decaying wood (e.g., Salix sp.) (pond, river) Japan, USA Hirayama etal. (2010)
Lolia aquatica Decayed stem of Phragmites australis (irrigation canal) Egypt Abdel-Aziz and Abdel-Wahab (2010),
Abdel-Aziz (2016c)
Lolia dictyospora Submerged decayed rachis of Phoenix dactylifera (river) Egypt Abdel-Aziz (2016c)
Neolindgomyces submersus Submerged decaying wood (river) Australia Hyde and Goh (1999b)
Longipedicellataceae
Longipedicellata aptrootii Submerged decaying wood and bamboo (river, stream, river) China, Malaysia, Philippines, Thailand Hyde and Wong (1999a), Cai etal.
(2003a), Kurniawati etal. (2010), Zhang
etal. (2016), Dong etal. (2020b), Goh
and Hyde (1999)
Longipedicellata aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Pseudoxylomyces aquaticus Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Pseudoxylomyces elegans Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks), twigs, bark Australia, Brazil, China, India, Japan,
Seychelles, Thailand, USA Goh etal. (1997a), Sivichai etal. (2000b,
2002), Tsui and Hyde (2004), Raja etal.
(2007), Barbosa and Gusmão (2011),
Patil and Borse (2015), Tanaka etal.
(2015), Dong etal. (2020b), Chethana
etal. (2021)
Submersispora variabilis Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Lophiostomataceae

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Biappendiculispora japonica Submerged decaying wood (stream) China Bao etal. (2019c)
Crassiclypeus aquaticus Submerged decaying wood (pond) Japan Hashimoto etal. (2018)
Pseudocapulatispora longiappendiculata Submerged decaying wood (stream) China Dong etal. (2020b)
Flabellascoma aquaticum Submerged decaying wood (river) China Bao etal. (2019c)
Lentistoma aquaticum Submerged decaying wood Thailand Dong etal. (2020b)
Lentistoma bipolare Submerged decaying wood (lake, river, reservoir, stream,
waterfall), submerged decaying Licuala longicalycata and
Metroxylon sagu (peat swamp)
Australia, Brunei, China, Japan, Malay-
sia, Philippines Seychelles, South
Africa, Thailand, USA
Hyde (1995a,b), Hyde and Aptroot
(1998a), Hyde and Goh (1998c), Goh
and Hyde (1999) Sivichai etal. (2000b),
Tsui etal. (2000), Ho etal. (2001,
2002), Sivichai etal. (2002), Cai etal.
(2003a), Luo etal. (2004), Pinruan
etal. (2007, 2014), Raja etal. (2009b),
Hashimoto etal. (2018)
Lophiostoma amphibium Equisetum fluviatile, Carex rostrata, C. nigra Austria Magnes and Hafellner (1991)
Lophiostoma appendiculatum Submerged Salix wood (lake) USSR Dudka (1963)
Lophiostoma aquaticum Submerged decaying wood (river, stream) Brunei, Malaysia, South Africa. UK Webster (1965), Hyde and Aptroot
(1998a), Hyde etal. (1998a), Ho etal.
(2001)
Lophiostoma frondisubmersum Submerged rachides of Livistona, submerged decaying Lic-
uala longicalycata and Eleiodoxa conferta (peat swamp),
submerged decaying wood (stream, river, waterfall) and
bamboo (river)
New Guinea, Philippines, Thailand,
USA Hyde (1994a), Cai etal. (2003a), Pinnoi
etal. (2006), Pinruan etal. (2007), Raja
etal. (2009b), Kurniawati etal. (2010)
Lophiostoma lunisporum Submerged decaying wood (lake) Australia Hyde and Goh (1998a)
Lophiostoma maquilingense Submerged decaying wood (stream) Philippines Hyde and Aptroot (1998a)
Lophiostoma proprietunicatum Submerged decaying wood (lake) China Tsui etal. (1999b), Luo etal. (2004)
Lophiostoma purpurascens Submerged decaying wood (river, stream) Australia, China, New Guinea Hyde and Aptroot (1998a), Tsui etal.
(2000)
Neotrematosphaeria biappendiculata Submerged decaying wood (pond) Japan Tanaka etal. (2005a)
Neovaginatispora fuckelii Submerged decaying wood (stream) China Bao etal. (2019c)
Sigarispora arundinis Phragmites australis USSR Dudka (1963)
Sigarispora clavata Submerged decaying wood China Bao etal. (2019c)
Vaginatispora aquatica Submerged decaying wood (stream) Australia, China, Thailand Hyde (1995c), Tsui etal. (2000), Zhang
etal. (2014a)
Vaginatispora armatispora Submerged decaying wood (waterfall) Thailand Hu etal. (2010b), Bao etal. (2019c),
Hyde etal. (2019)
Lophiotremataceae
Lophiotrema culmifragum Schoenoplectus lacustri USSR Dudka (1963)
Lophiotrema rubi Submerged, partially decomposed wood (lake) USA Fallah and Shearer (2001)
Massarinaceae

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Byssothecium circinans Submerged decaying wood (stream) Hungary Révay and Gönczöl (1990), Gönczöl and
Revay (1993)
Byssothecium flumineum Twigs of Prunus serotina (creek) USA Crane etal. (1992)
Helminthosporium aquaticum Submerged decaying wood (stream) China Zhu etal. (2016)
Helminthosporium velutinum Submerged decaying wood (stream) China Zhu etal. (2016)
Massarina australiensis Submerged decaying wood (waterfalls) Australia Hyde (1992a)
Massarina eburnea Submerged decaying wood (lake) USA Fallah and Shearer (2001)
Melanommataceae
Bertiella ellipsoidea Submerged decaying wood (stream) Thailand Hyde etal. (2018)
Byssosphaeria schiedermayeriana Submerged decaying wood (river) Seychelles Hyde and Goh (1998c)
Byssosphaeria siamensis Submerged decaying wood (stream) Thailand Hyde etal. (2018)
Camposporium appendiculatum Submerged decaying wood (river) China Hyde etal. (2020b)
Camposporium antennatum Foam (stream), water (stream) Hungary, Venezuela Rajashekhar and Kaveriappa (2003),
Smits etal. (2007)
Camposporium cambrense Submerged decaying wood (river, stream, waterfall) Thailand Kurniawati etal. (2010)
Camposporium dulciaquae Submerged decaying wood (stream) Thailand Calabon etal. (2021a)
Camposporium fusisporum Submerged decaying wood (river) Philippines Cai etal. (2003a)
Camposporium hyalinum Submerged cupules of Fagus sylvatica (stagnant water) UK Abdullah (1980)
Camposporium marylandicum Submerged decaying wood (river) USA Shearer (1974)
Camposporium multiseptatum Submerged decaying wood (river) China Hyde etal. (2020b)
Camposporium pellucidum Submerged decaying wood (stream, reservoir), foam
(stream), water-filled treeholes Austria, Brazil, China, Hungary, India,
Ireland, UK, Venezuela Ingold (1975), Gönczöl (1976b), Abdul-
lah and Webster (1980), Voglmayr
(1996), Marvanová (2000), Gönczöl
and Révay (2003), Rajashekhar and
Kaveriappa (2003), Smits etal. (2007),
Schoenlein-Crusius etal. (2009), Hyde
etal. (2020b)
Camposporium quercicola Submerged decaying bamboo (river) Philippines Cai etal. (2003a)
Camposporium septatum Submerged decaying wood (stream) Thailand Hyde etal. (2020b)
Herpotrichia dalisayi Submerged decaying wood (stream) Philippines Hyde and Aptroot (1998b)
Mamillisphaeria dimorphospora Submerged decaying wood (stream) Australia, Brunei, Malaysia Bareen and Iqbal (1994), Hyde etal.
(1996b), Ho etal. (2001), Vijaykrishna
and Hyde (2006), Hyde (2007)
Melanomma australiense Submerged decaying wood (stream) Australia Hyde and Goh (1999b), Vijaykrishna and
Hyde (2006)
Phragmocephala atra Submerged decaying wood (stream) China Su etal. (2015)
Phragmocephala garethjonesii Submerged decaying wood (stream) China Su etal. (2015)
Phragmocephala glanduliformis Water-filled treeholes Hungary Gönczöl and Révay (2003)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Tumularia aquatica Foam (rivulet, stream), submerged decaying leaves of broad-
leaved trees and shrubs, water (river), submerged leaf litter
of Alnus glutinosa (stream)
Austria, Brazil, Canada, Cuba, Czech
Republic, France, India, Poland, UK,
USA
Ingold (1943a), Ranzoni (1953),
Marvanová and Marvan (1969), Iqbal
and Webster (1973), Bärlocher (1987),
Marvanová and Descals (1987), Sridhar
and Kaveriappa (1992), Gessner etal.
(1993), Voglmayr (1996), Marvanová
(2000), Belliveau and Bärlocher (2005),
Schoenlein-Crusius etal. (2009), Pietry-
czuk etal. (2018)
Tumularia tuberculata Foam (stream), submerged leaves of Alnus glutinosa, Alnus
sp., Quercus, and Fagus sylvatica, stemflow and through-
fall of tree canopies, water (stream)
Austria, Canada, France, Hungary,
India, Spain Gönczöl (1976a), Bärlocher (1987), Gess-
ner etal. (1993), Marvanová (2000),
Sridhar and Karamchand (2009),
Menéndez etal. (2012), Cornut etal.
(2014)
Morosphaeriaceae
Aquihelicascus songkhlaensis Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Aquihelicascus thalassioideus Submerged decaying wood (lake, river, stream, waterfall),
bamboo (river), twigs Australia, Brunei, China, Martinique,
Peru, Japan, Philippines, Thailand Hyde and Aptroot (1998a), Tsui etal.
(2000), Ho etal. (2001), Cai etal.
(2003a), Luo etal. (2004) Kurniawati
etal. (2010), Zhang etal. (2013a),
Tanaka etal. (2015), Shearer etal.
(2015), Dong etal. (2020b)
Aquihelicascus yunnanensis Submerged decaying wood (stream) China Dong etal. (2020b)
Neohelicascus aquaticus Submerged decaying wood (pond, stream) China, Japan, Thailand Zhang etal. (2013a), Tanaka etal. (2015),
Zeng etal. (2018), Dong etal. (2020b)
Neohelicascus chiangraiensis Submerged decaying wood (pond) Thailand Luo etal. (2016c)
Neohelicascus egyptiacus Submerged decaying dicotyledonous wood (river) Egypt Zhang etal. (2013a)
Neohelicascus elaterascus Submerged decaying wood (stream, river) and bamboo
(river), twigs (lake) Brunei, Chile, China, Japan, Peru, Phil-
ippines, South Africa, Thailand, USA Shearer (1993b), Hyde etal. (1998a), Goh
and Hyde (1999), Tsui etal. (2000), Ho
etal. (2001), Cai etal. (2003a), Luo
etal. (2004), Raja etal. (2009b), Tanaka
etal. (2015), Shearer etal. (2015), Hyde
etal. (2020a), Dong etal. (2020b)
Neohelicascus gallicus Submerged wood of Alnus glutinosa, Corylus avellana, Rob-
inia pseudoacacia, Fraxinus excelsior, Salix sp. (brook) France Zhang etal. (2014c)
Neohelicascus submersus Submerged decaying wood (river) China Dong etal. (2020b)
Neohelicascus unilocularis Submerged decaying wood (river) Martinique Zhang etal. (2014c), Zhang etal. (2015)
Neohelicascus uniseptatus Submerged decaying wood (stream) Thailand Luo etal. (2016c)
Mycoporaceae
Mycoporum antecellens Twig (waterfall) Canada Björk etal. (2009)
Neopyrenochaetaceae

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Neopyrenochaeta asexualis Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Neopyrenochaeta glabra Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Neopyrenochaeta submersa Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Pyrenochaetopsidaceae
Pyrenochaetopsis aquatica Submerged plant debris Spain Magaña-Dueñas etal. (2021)
Nigrogranaceae
Nigrograna aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Nigrograna cangshanensis Submerged decaying wood (stream) China Tibpromma etal. (2017)
Occultibambusaceae
Brunneofusispora hyalina Submerged decaying wood (stream) Thailand Calabon etal. (2021a)
Occultibambusa aquatica Submerged decaying bamboo (waterfall) Thailand Hyde etal. (2016b)
Occultibambusa kunmingensis Submerged decaying bamboo (stream) China Dong etal. (2020b)
Occultibambusa pustula Submerged decaying wood (stream) China Dong etal. (2020b)
Seriascoma didymosporum Submerged decaying wood (stream) China Dong etal. (2020b)
Parabambusicolaceae
Aquastroma magniostiolatum Submerged decaying wood (pond, stream) Japan, Thailand Tanaka etal. (2015), Dong etal. (2020b)
Parabambusicola aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Paramonodictys solitarius Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Periconiaceae
Periconia aquatica Submerged decaying wood (river) China Hyde etal. (2017b)
Periconia byssoides Submerged decaying wood (lake) USA, China Shearer (1972), Luo etal. (2004)
Periconia cookei Submerged decaying wood (stream) China Hyde etal. (2018)
Periconia digitata Submerged decaying wood (lake) China Luo etal. (2004)
Periconia minutissima Submerged decaying wood (stream, river) Brazil, China, Philippines Cai etal. (2002b), Cai etal. (2003a) Luo
etal. (2004), Barbosa etal. (2013b),
Hyde etal. (2017b)
Periconia pseudobyssoides Submerged decaying wood (stream) China Hyde etal. (2018)
Periconia pseudodigitata Dead culms of Phragmites australis, submerged stems of
unidentified herbaceous plant (river) Japan Tanaka etal. (2015)
Periconia submersa Submerged decaying wood (river) China Hyde etal. (2017b)
Phaeoseptaceae
Phaeoseptum aquaticum Submerged branch of Robinia pseudoacacia, driftwood of
Salix sp France Zhang etal. (2013b)
Pleopunctum pseudoellipsoideum Submerged decaying wood China Dong etal. (2020b)
Phaeosphaeriaceae
Didymocyrtis pseudeverniae Nature Reserve Ukraine, UK Khodosovtsev etal. (2016)
Juncaceicola alpina Carex rostrata, Equisetum fluviatile Austria Magnes and Hafellner (1991)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Leptospora rubella Submerged decaying wood (lake) UK Hyde etal. (2016b)
Ophiobolus typhae Submerged decaying leaves of Typha latifolia UK Ingold (1951)
Ophiosphaerella herpotricha Typha sheath UK Pugh and Mulder (1971)
Phaeosphaeria caricis Submerged decaying grass (lake) USA Fallah and Shearer (2001)
Phaeosphaeria caricinella Carex rostrata, C. paupercula subspec. irrigua, Juncus
filiformis Austria Magnes and Hafellner (1991)
Phaeosphaeria culmorum Carex rostrata, Equisetum fluviatile, Glyceria fluitans Austria Magnes and Hafellner (1991)
Phaeosphaeria eustoma Submerged decaying wood (river), Typha leaves, Scirpus UK, USA Eaton and Jones (1971a), Pugh and
Mulder (1971), Cavaliere 1975)
Phaeosphaeria halima Phragmites australis stem UK Taligoola etal. (1972)
Phaeosphaeria herpotrichoides Submerged beech wood, Carex rostrata Austria, UK Eaton and Jones (1971b), Magnes and
Hafellner (1991)
Phaeosphaeria juncicola Scirpus, Typha USA Cavaliere (1975)
Phaeosphaeria licatensis Typha leaves UK Pugh and Mulder (1971)
Phaeosphaeria microscopica Phragmites australis leaves and steam UK Apinis etal. (1972b), Taligoola etal.
(1972)
Phaeosphaeria sowerbyi Typha leaves USA Cavaliere (1975)
Phaeosphaeria vilasensis Submerged decaying Juncus stem (lake) USA Fallah etal. (1999b)
Pleomassariaceae
Beverwykella cerebriformis Submerged decaying angiospermic leaves (stream) Malaysia Nawawi and Kuthubutheen (1988b)
Beverwykella clathrata Submerged decaying leaves of Pinus serotina and evergreen
broad-leaved shrubs (puddles), Calophyllum brasiliense
(stream)
Brazil, USA Voglmayr and Delgado-Rodríguez (2003),
Fiuza etal. (2019)
Beverwykella pulmonaria Decaying leaves (tin can filled with rainwater), rainwater
from building roofs, blackened leaf (pond), submerged
Fagus leaves (stream), water-filled treeholes
Hungary, Ireland, Netherlands, Poland,
UK van Beverwijk (1954), Abdullah and
Webster (1980), Czeczuga and
Orłowska (1997), Gönczöl and Révay
(2003)
Splanchnonema britzelmayrianum Submerged decaying wood (stream) Hungary Révay and Gönczöl (1990)
Pleomonodictydaceae
Pleomonodictys capensis Submerged decaying wood (stream) Brunei, China Ho etal. (2001), Bao etal. (2021a)
Pleosporaceae
Alternaria alternata Water (river) Poland Pietryczuk etal. (2018)
Clathrospora permunda Scirpus, Typha (lake) USA Cavaliere (1975)
Comoclathris typhicola Submerged Typha leaves UK Pugh and Mulder (1971)
Curvularia eragrostidis Submerged decaying wood (stream) China Su etal. (2015)
Curvularia geniculata Submerged leaf Brazil da Silva and Gusmão (2015)
Curvularia muehlenbeckiae Submerged decaying twig Korea Nam etal. (2020)
Curvularia pseudobrachyspora Submerged decaying plant material Korea Nam etal. (2020)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Curvularia verruculosa Submerged decaying wood (stream) China Su etal. (2015)
Decorospora gaudefroyi Submerged decaying wood UK Webster and Lucas (1961)
Platysporoides tirolensis Carex rostrata (lake) Austria Magnes and Hafellner (1991)
Pleospora hydrophila Salix wood USSR Dudka (1963)
Pleospora incerta Carex rostrata Austria Magnes and Hafellner (1991)
Pleospora palustris Scirpus maritimus UK Webster and Lucas (1961)
Pleospora pelagica Submerged decaying wood (stream) China Hu etal. (2013b)
Pleospora phaeospora Equisetum fluviatile (lake) Austria Magnes and Hafellner (1991)
Pleospora scirpicola Submerged decaying Scirpus validus stems (lake), Schoeno-
plectus lacustris stalks (la ke) UK, USA, USSR Ingold (1955), Dudka (1963), Fallah and
Shearer (2001)
Pleospora submersa Submerged decaying Salix twigs UK Webster and Lucas (1961)
Stemphylium majusculum Submerged decaying wood (stream) Brunei Ho etal. (2001)
Typhicola typharum Submerged decaying Typha latifolia stem and leaves (lake) USA, UK Webster (1955), Pugh and Mulder (1971),
Cavaliere (1975), Fallah and Shearer
(2001)
Pseudoastrosphaeriellaceae
Carinispora nypae Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Pseudoastrosphaeriella africana Dead bamboo stems (waterfall) Thailand Phookamsak etal. (2015)
Pseudoastrosphaeriella aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Pseudoastrosphaeriella bambusae Submerged dead branches of bamboo and wood (stream) Thailand Phookamsak etal. (2015), Dong etal.
(2020b)
Pseudoastrosphaeriella longicolla Submerged dead branches of bamboo Thailand Phookamsak etal. (2015)
Pseudoastrosphaeriella papillata Submerged decaying wood and bamboo (lake), decaying
Licuala longicalycata (peat swamp) Brunei, China, Philippines, Thailand Hyde and Fröhlich (1998), Cai etal.
(2003a), Luo etal. (2004), Pinruan etal.
(2014)
Pseudoberkleasmiaceae
Pseudoberkleasmium chiangmaiense Submerged decaying wood (river) China Bao etal. (2021a)
Roussoellaceae
Roussoella aquatica Submerged decaying wood (stream) China Dong etal. (2020b)
Roussoella intermedia Submerged, partially decomposed bark (lake) USA Fallah and Shearer (2001)
Roussoella minutella Submerged decaying wood and bamboo (river) Philippines Cai etal. (2003a)
Neoroussoella bambusae Submerged decaying wood (stream) China Dong etal. (2020b)
Neoroussoella leucaenae Submerged decaying wood (stream) China Dong etal. (2020b)
Sporormiaceae
Sporormiella minima Submerged wood (water-cooling towers) UK Eaton and Jones (1971b)
Sporormiella minimoides Submerged decaying wood (lake) China Luo etal. (2004)
Westerdykella aquatica Endophytic in Acorus calamus (pond), mud (rice field),
sediment China, Korea Song etal. (2020), Goh etal. (2021a)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Westerdykella aurantiaca Sediment Korea Goh etal. (2021a)
Westerdykella dispersa Sediment Korea Goh etal. (2021a)
Westerdykella multispora Beech wood UK Eaton (1972)
Westerdykella purpurea Submerged decaying wood (stream) Thailand Chethana etal. (2021)
Striatiguttulaceae
Longicorpus striatisporus Submerged decaying wood (river, stream, waterfall) Thailand Kurniawati etal. (2010)
Teichosporaceae
Chaetomastia clavigera Schoenoplectus lacustris USSR Dudka (1963)
Testudinaceae
Angustospora nilensis Submerged decaying wood (river) Egypt Li etal. (2016)
Tetraplosphaeriaceae
Aquatisphaeria thailandica Submerged decaying wood (stream) Thailand Li etal. (2021b)
Ernakulamia cochinensis Submerged decaying wood and leaf (stream) Brazil, Thailand da Silva and Gusmão (2015), Dong etal.
(2020b)
Shrungabeeja aquatica Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Shrungabeeja vadirajensis Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Tetraploa abortiva Decaying leaf litter (river) Argentina Schoenlein-Crusius and Piccolo Grandi
(2003)
Tetraploa aquatica Submerged decaying wood (river) China Li etal. (2020)
Tetraploa aristata Decaying palm leaflets, submerged branches, roots of
Ctenanthe oppenheimiana and Stromanthe sanguinea,
stemflow and throughfall of tree canopies, submerged
decaying woods (stream, waterfall, river), foam (stream),
decaying Metroxylon sagu (peat swamp)
Argentina, Brazil, China, Hungary,
India, Japan, Malaysia, Peru, Thailand Révay (1993), Tsui etal. (2001d),
Schoenlein-Crusius and Piccolo Grandi
(2003), Hosoya etal. (2006), Hosoya
and Tanaka (2007), Karamchand and
Sridhar 2008, Sridhar and Karamchand
(2009), Kurniawati etal. (2010), Pin-
ruan etal. (2014)
Tetraploa ellisii Leaf lesions of Oryza sativa (river) Peru Schoenlein-Crusius and Piccolo Grandi
(2003)
Tetraploa puzheheiensis Submerged decaying wood (wetland) China Dong etal. (2020b)
Tetraploa scheueri Decaying leaves, Carex acutiformis UK Scheuer (1991)
Tetraploa thailandica Submerged decaying wood Thailand Bao etal. (2021a)
Tetraploa yunnanensis Submerged decaying wood (werland) China, Thailand Dong etal. (2020b)
Triplosphaeria acuta Submerged bamboo culms (stream) Japan Tanaka etal. (2009)
Thyridariaceae
Thyridaria triseptata Submerged decaying wood (lake) USA Fallah and Shearer (2001)
Torulaceae
Dendryphion aquaticum Submerged decaying wood (stream) China Su etal. (2016a)
Dendryphion comosum Submerged leaves Brazil da Silva and Gusmão (2015)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Dendryphion fluminicola Submerged decaying wood China Su etal. (2018)
Dendryphion nanum Submerged decaying wood (river, stream) China, USA Su etal. (2016a)
Dendryphion submersum Submerged decaying wood (stream) China Su etal. (2016a)
Neotorula aquatica Submerged decaying wood (stream) China Su etal. (2016a)
Neotorula submersa Submerged decaying wood (river) China Hyde etal. (2016b)
Rostriconidium aquaticum Submerged decaying wood (river) China Su etal. (2018)
Rostriconidium cangshanense Submerged decaying wood (stream) China Shen etal. (2021)
Rostriconidium pandanicola Submerged decaying wood (river) China Shen etal. (2021)
Torula aquatica Submerged decaying wood (lake, river) China Su etal. (2018)
Torula herbarum Submerged decaying wood (stream, lake) Brazil, China Luo etal. (2004), Barbosa etal. (2013b)
Torula fici Submerged decaying wood (river) China Su etal. (2018)
Torula gaodangensis Submerged decaying twig (river) China Hyde etal. (2018)
Torula masonii Submerged decaying wood (lake) China Su etal. (2018)
Trematosphaeriaceae
Falciformispora aquatica Submerged decaying wood (waterfall) Thailand Hyde etal. (2019)
Falciformispora lignatilis Submerged decaying wood USA Raja etal. (2009b)
Falciformispora uttaraditensis Submerged decaying twigs (creek) Thailand Hyde etal. (2020b)
Hadrospora fallax Submerged decaying wood (lake, river) China, USA Shearer and Crane (1971), Luo etal.
(2004
Trematosphaeria callicarpa Submerged decaying wood (lake) and bamboo (river) China, Philippines, USA Cai etal. (2003a), Luo etal. (2004)
Trematosphaeria confusa Submerged decaying wood (stream) Australia, Malaysia Hyde (1995b), Ho etal. (2001)
Trematosphaeria hydrela Decaying wood Union of Soviet Socialist Republics Petrak (1925)
Trematosphaeria lineolatispora Submerged decaying wood USA Raja etal. (2009b)
Trematosphaeria pertusa Submerged decaying wood (water-cooling tower) Hungary, Japan, UK Eaton and Jones (1971b), Révay and
Gönczöl (1990), Kane etal. (2002),
Tanaka etal. (2015)
Wicklowiaceae
Wicklowia aquatica Submerged decorticated debris (river) USA Raja etal. (2010b)
Wicklowia phuketensis Submerged decaying wood (stream) Thailand Calabon etal. (2020a)
Wicklowia submersa Submerged decaying wood (stream) Thailand Boonmee etal. (2019)
Pleosporales genera incertae sedis
Ascorhombispora aquatica Submerged bamboo (stream) China Cai and Hyde (2007b)
Fusiformiseptata crocea Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Isthmosporella pulchra Submerged decorticated wood (lake) USA Shearer and Crane (1999)
Mycoenterolobium aquadictyosporium Submerged decaying wood (stream) Thailand Calabon etal. (2020c)
Mycoenterolobium platysporum Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Scolecobasidium anellii Foam and submerged decaying leaf litter (stream) India Rajashekhar and Kaveriappa (2003)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Scolecobasidium dendroides Submerged decaying wood (stream) Brunei Ho etal. (2001)
Scolecobasidium flagelliferum Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Rebentischia unicaudata Submerged decaying wood (swamp) USA Shearer and Crane (1986)
Repetophragma fasciatum Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Trypetheliales
Polycoccaceae
Polycoccum sp. Stream and lake margin Europe Orange (2002)
Trypetheliaceae
Arthopyrenia fallaciosa River Watershed Canada Maloles etal. (2018)
Tubeufiales
Tubeufiaceae
Acanthohelicospora aurea Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)
Acanthohelicospora guianensis Submerged decaying wood (stream, reservoir), stemflow China, India, Thailand Goh and Hyde (1999), Chuaseeharon-
nachai etal. (2013), Ghate and Sridhar
(2015)
Acanthophiobolus helicosporus Carex paniculata Denmark Munk (1957)
Aquaphila albicans Submerged decaying wood (lake, river, stream, waterfall),
bamboo (river)submerged rotting wood of angiosperm Australia, Brunei, China, Malaysia,
Philippines, Thailand Goh etal. (1998b), Ho etal. (2001), Cai
etal. (2003a), Luo etal. (2004), Kurni-
awati etal. (2010), Hyde etal. (2016b),
Dong etal. (2020b)
Berkleasmium aquaticum Submerged decaying wood (stream) China Lu etal. (2017a)
Berkleasmium corticola Submerged decaying wood (stream) Brazil, China Ho etal. (2001), Barbosa etal. (2013b)
Berkleasmium guangxiense Submerged decaying wood (stream) China Lu etal. (2017a)
Berkleasmium latisporum Submerged decaying wood (stream) Thailand Lu etal. (2017a), Dong etal. (2020b)
Berkleasmium longisporum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Berkleasmium micronesiacum Submerged decaying Eleiodoxa conferta and Nenga pumila
(peat swamp) Thailand Pinnoi etal. (2006), Pinruan etal. (2014)
Berkleasmium monilicellulare Submerged decaying bamboo (river) Philippines Cai etal. (2003a)
Berkleasmium moriforme Submerged decaying wood (stream) Malaysia Ho etal. (2001), Hyde (2007)
Berkleasmium thailandicum Submerged decaying wood (stream) Thailand Lu etal. (2017a)
Berkleasmium typhae Submerged decaying Licuala longicalycata and Eleiodoxa
conferta (peat swamp) Thailand Pinruan etal. (2007), (Pinnoi etal. (2006)
Berkleasmium zhejiangense Submerged decaying wood (stream) China Wongsawas etal. (2009a)
Boerlagiomyces grandisporus Submerged decaying wood (river) Philippines Stanley and Hyde (1997), Cai etal.
(2003a)
Boerlagiomyces lacunosisporus Submerged decaying wood (river) Australia Hyde (1992b)
Boerlagiomyces websteri Submerged decorticated wood (river) USA, Thailand Shearer and Crane (1995), Sivichai and
Boonyene (2004)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Camporesiomyces vaccinii Stemflow and throughfall of palm species India Ghate and Sridhar (2015)
Chlamydotubeufia chlamydospora Submerged twig (creek) Panama Shearer (1987)
Chlamydotubeufia cylindrica Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Chlamydotubeufia huaikangplaensis Submerged decaying wood (stream) Thailand Hyde etal. (2017b)
Chlamydotubeufia krabiensis Submerged decaying wood (stream) Thailand Hyde etal. (2017b)
Dematiohelicoma perelegans Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)
Dematiohelicomyces helicosporus Submerged decaying wood (stream) Thailand Hyde etal. (2016b), Lu etal. (2018c)
Dictyospora thailandica Submerged decaying wood Thailand Brahmanage etal. (2017), Lu etal.
(2018c)
Helicoarctatus aquaticus Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicoarctatus thailandicus Submerged decaying wood (stream) Thailand Yuan etal. (2020)
Helicodochium amazonicum Submerged decaying wood Brazil Monteiro etal. (2014c)
Helicodochium aquaticum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicohyalinum aquaticum Submerged decaying wood Thailand Brahmanage etal. (2017)
Helicoma ambiens Stemflow and throughfall of tree canopies India Sridhar and Karamchand (2009)
Helicoma atroseptatum Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)
Helicohyalinum infundibulum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicoma aquaticum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicoma gigasporum Submerged decaying wood, decaying Licuala longicalycata
(peat swamp) China. Thailand Tsui etal. (2001b), Pinruan etal. (2014)
Helicoma guttulatum Submerged decaying wood, bamboo (river) Philippines, Thailand Hyde etal. (2016b)
Helicoma hongkongense Submerged decaying wood (stream) China, Thailand Tsui etal. (2001b), Lu etal. (2018c)
Helicoma longisporum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicoma multiseptatum Submerged decaying wood (stream) China Lu etal. (2018c)
Helicoma nematosporum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicoma pannosum Submerged decaying wood (stream, river) Malaysia, Thailand Ho etal. (2001), Hyde (2007), Chuasee-
haronnachai etal. (2013)
Helicoma resinae Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)
Helicoma septoconstrictum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicomyces ambiguus Submerged decaying wood (river) USA Lamore and Goos (1978)
Helicomyces bellus Water-filled treeholes Hungary Gönczöl and Révay (2003)
Helicomyces chiayiensis Submerged decorticated wood (stream) China Kuo and Goh (2018a)
Helicomyces colligatus Submerged decaying wood, stemflow and throughfall of tree
canopies, foam (stream) India, Thailand, Venezuela Smits etal. (2007), Sridhar and Karam-
chand (2009), Lu etal. (2018c)
Helicomyces geniculatus Submerged decaying wood (stream) China Kuo and Goh (2018a)
Helicomyces hyalosporus Submerged decaying wood (stream) China Lu etal. (2018c)
Helicomyces macrofilamentosus Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)

Fungal Diversity
1 3
Taxon Substrate Distribution References
Helicomyces roseus Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (lake, stream, river, water-
fall), stemflow and throughfall of tree canopies and palm
species, leaves (stream), foam (stream)
Australia, Brazil, Brunei, China, India,
Malaysia, Thailand Hyde and Goh (1997), Ho etal. (2001,
2002), Sivichai etal. (2002), Rajashek-
har and Kaveriappa (2003), Luo etal.
(2004), Sridhar and Karamchand
(2009), Kurniawati etal. (2010), Chua-
seeharonnachai etal. (2013), Barbosa
etal. (2013b), Ghate and Sridhar (2015)
Helicomyces scandens Stemflow and throughfall of tree canopies India Sridhar and Karamchand (2009)
Helicomyces torquatus Submerged decaying wood (stream), twig, foam (stream),
stemflow and throughfall of palm species Brunei, China, India, Malaysia,
Panama, Thailand, Venezuela Lane and Shearer (1984), Ho etal.
(2001), Smits etal. (2007), Ghate and
Sridhar (2015), Lu etal. (2018c), Kuo
and Goh (2018b)
Helicosporium aquaticum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicosporium flavidum Submerged decaying culms of Miscanthus floridulus
(stream) China Hsieh etal. (2021a)
Helicosporium flavisporum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicosporium flavum Submerged decaying wood Thailand Brahmanage etal. (2017)
Helicosporium gracile Submerged decaying wood (wood) Thailand Chuaseeharonnachai etal. (2013)
Helicosporium lumbricopsis Submerged decaying wood (wood) Thailand Chuaseeharonnachai etal. (2013)
Helicosporium murinum Stemflow and throughfall of palm species India Ghate and Sridhar (2015)
Helicosporium panachaeum Submerged decaying wood (wood) Thailand Chuaseeharonnachai etal. (2013)
Helicosporium setiferum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicosporium thailandense Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Helicosporium vesicarium Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Helicotruncatum palmigenum Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Helicotubeufia guangxiensis Submerged decaying wood (stream) China Liu etal. (2018a)
Helicotubeufia jonesii Submerged decaying wood (stream) China Liu etal. (2018a)
Muripulchra aquatica Submerged decaying wood (stream) China Luo etal. (2017)
Neochlamydotubeufia fusiformis Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neochlamydotubeufia khunkornensis Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicomyces aquaticus Submerged decaying wood (river) China Luo etal. (2017)
Neohelicomyces dehongensis Submerged decaying wood (stream) China Dong etal. (2020b)
Neohelicomyces grandisporus Submerged decaying wood (river) China Luo etal. (2017)
Neohelicomyces hyalosporus Submerged decaying wood (stream) China Lu etal. (2018c)
Neohelicomyces longisetosus Submerged decaying culms of Miscanthus floridulus
(stream) China Hsieh etal. (2021a)
Neohelicomyces submersus Submerged decaying wood (stream) China Luo etal. (2017)
Neohelicomyces thailandicus Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Table 3 (continued)

Fungal Diversity
1 3
Table 3 (continued)
Taxon Substrate Distribution References
Neohelicosporium acrogenisporum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicosporium aquaticum Submerged decaying wood (stream) China Lu etal. (2018a)
Neohelicosporium astrictum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicosporium ellipsoideum Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicosporium guangxiense Submerged decaying wood (stream) China Lu etal. (2018a)
Neohelicosporium griseum Submerged decaying wood (wood, reservoir, river) Brunei, China, Malaysia, Thailand Goh and Hyde (1999), Ho etal. (2001),
Hyde (2007), Chuaseeharonnachai etal.
(2013)
Neohelicosporium hyalosporum Submerged decaying wood (stream) China Lu etal. (2018a)
Neohelicosporium irregulare Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicosporium krabiense Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Neohelicosporium parvisporum Submerged decaying wood (stream) China Lu etal. (2018a)
Neohelicosporium submersum Submerged decaying wood (stream) Thailand Dong etal. (2020b)
Neohelicosporium sympodiophorum Submerged decaying culm of Miscanthus floridulus (stream) China Hsieh etal. (2021a)
Neohelicosporium taiwanense Submerged decaying wood (stream) China Kuo and Goh (2018a)
Neohelicosporium thailandicum Submerged decaying wood (stream) Thailand Lu etal. (2018a)
Neotubeufia krabiensis Submerged decaying wood (stream) Thailand Chaiwan etal. (2017)
Parahelicomyces talbotii Submerged decaying wood (stream)` China, Thailand Lu etal. (2018c), Hsieh etal. (2021a)
Parahelicomyces aquaticus Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Parahelicomyces hyalosporus Submerged decaying wood (stream) China Luo etal. (2017)
Pseudohelicoon gigantisporum Submerged decaying wood (stream), water (river) Australia, Poland, Thailand Goh and Hyde (1996a), Chuaseeharon-
nachai etal. (2013), Pietryczuk etal.
(2018)
Pseudohelicoon subglobosum Submerged decaying wood (stream) China Goh and Kuo (2018)
Thaxteriellopsis lignicola Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia abundata Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia aquatica Submerged decaying wood (stream) China, Thailand Luo etal. (2017), Lu etal. (2018c), Dong
etal. (2020b)
Tubeufia brevis Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia brunnea Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia cerea Foam, submerged decaying leaf litter (stream) India Rajashekhar and Kaveriappa (2003)
Tubeufia chlamydospora Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia claspisphaeria Submerged decaying wood, submerged decaying Licuala
longicalycata and Eleiodoxa conferta (peat swamp) China, Thailand Kodsueb etal. (2004), Pinnoi etal.
(2006), Pinruan etal. (2007)
Tubeufia cylindrothecia Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (stream) China, Thailand Sivichai etal. (2002), Luo etal. (2017),
Lu etal. (2018c), Dong etal. (2020b)
Tubeufia dictyospora Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia eccentrica Submerged decaying wood (stream) China Lu etal. (2018c)

Fungal Diversity
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Table 3 (continued)
Taxon Substrate Distribution References
Tubeufia fangchengensis Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia filiformis Submerged decaying wood (stream) Thailand Lu etal. (2017b), Lu etal. (2018c)
Tubeufia geniculata Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia guangxiensis Submerged decaying wood (stream) China Chaiwan etal. (2017), Lu etal. (2018c)
Tubeufia hechiensis Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia hyalospora Submerged decaying wood (stream) Thailand Hyde etal. (2016b)
Tubeufia inaequalis Submerged decaying wood (stream) China, Thailand Lu etal. (2018c)
Tubeufia krabiensis Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia latispora Submerged decaying wood (stream) Thailand Lu etal. (2017b)
Tubeufia laxispora Submerged decaying wood (stream) Thailand Lu etal. (2017b), Lu etal. (2018c)
Tubeufia mackenziei Submerged decaying wood (stream) Thailand Lu etal. (2017b)
Tubeufia palmarum Submerged decaying wood (stream, river) Brunei, Malaysia Ho etal. (2001), Hyde (2007)
Tubeufia parvispora Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia rubra Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia sympodihylospora Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia sympodilaxispora Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia sympodiophora Submerged decaying wood China Ho etal. (2002)
Tubeufia taiwanensis Submerged decaying wood (stream) China Lu etal. (2018c)
Tubeufia tectonae Submerged decaying wood (stream) Thailand Hyde etal. (2016b), Lu etal. (2018c),
Dong etal. (2020b)
Tubeufia tratensis Submerged decaying wood (stream) Thailand Lu etal. (2018c)
Tubeufia xylophila Submerged decaying wood (stream) China Lu etal. (2018c)
Xenosporium africanum Submerged decaying culms of Miscanthus floridulus
(stream) China Kuo and Goh (2021)
Xenosporium formosiforme Submerged decaying culms of Miscanthus floridulus
(stream) China Kuo and Goh (2021)
Xenosporium parvulum Submerged decaying culms of Miscanthus floridulus
(stream) China Kuo and Goh (2021)
Wiesneriomycetaceae
Phalangispora bharathensis Throughfall of palm species India Ghate and Sridhar (2015)
Phalangispora constricta Foam (stream, river), sediment (dam, stream), stemflow and
throughfall of tree canopies India, Malaysia, Thailand, Venezuela Nawawi and Webster (1982a), Sridhar
and Kaveriappa (1992), Sakayaroj etal.
(2005), Smits etal. (2007), Sridhar and
Karamchand (2009), Sudheep and Srid-
har (2012), Ghate and Sridhar (2015)
Phalangispora nawawii Submerged decaying leaves (river), foam (stream) India, Malaysia, Venezuela Kuthubutheen (1987a), Sridhar and Kave-
riappa (1992), Smits etal. (2007)
Phalangispora sinensis Submerged decaying leaves (stream) China Guo etal. (2019)

Fungal Diversity
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Table 3 (continued)
Taxon Substrate Distribution References
Setosynnema isthmosporum Submerged decaying leaves (stream) Papua New Guinea, Thailand Shaw and Sutton (1985), Marvanová and
Hywel-Jones (2000)
Setosynnema limnetica Submerged leaves (river) India Borse etal. (2019)
Setosynnema yunnanense Submerged decaying dicotyledonous leaves (river) China Bai etal. (2013)
Speiropsis hyalospora Submerged leaves of Coffea arabica (stream), foam (stream) India Sridhar and Kaveriappa (1988, 1992),
Rajashekhar and Kaveriappa (2003)
Speiropsis pedatospora Rainwater from building roofs, submerged leaves of Coffea
arabica (stream), foam (stream, rivulet), stemflow and
throughfall of tree canopies
Austria, India, Poland, Thailand Sridhar and Kaveriappa (1988, 1992),
Voglmayr (1996), Czeczuga and
Orłowska (1997), Sivichai and Boony-
ene (2004), Sridhar and Karamchand
(2009)
Speiropsis scopiformis Submerged decaying angiospermous leaves (stream) Brazil, Malaysia Kuthubutheen and Nawawi (1987a),
Barbosa etal. (2013b)
Wiesneriomyces conjunctosporus Submerged decaying wood Malaysia Kuthubutheen and Nawawi (1988c)
Wiesneriomyces laurinus Rainwater from building roofs, submerged decaying Licuala
longicalycata (peat swamp), submerged decaying leaves
(stream), foam (stream)
Brazil, India, Malaysia, Poland, Thai-
land Czeczuga and Orłowska (1997),
Rajashekhar and Kaveriappa (2003),
Pinruan etal. (2007), Barbosa etal.
(2013b), Pinruan etal. (2014), Hosoya
etal. (2019)
Venturiales
Venturiaceae
Magnohelicospora fuscospora Submerged decaying leaves and twigs of Salix (reservoir) Ireland Abdullah and Webster (1980)
Dothideomycetes families incertae sedis
Argynnaceae
Lepidopterella palustris Submerged decaying wood (swamp) USA Shearer and Crane (1980), Raja etal.
(2009b)
Lepidopterella tangerina Submerged decaying wood (river) USA Raja and Shearer (2008), Raja etal.
(2009b)
Pseudorobillardaceae
Pseudorobillarda phragmitis Water (river), foam (rivulet, stream) Austria, India, Poland Czeczuga etal. (1990), Voglmayr (1996),
Rajashekhar and Kaveriappa (2003)
Pseudorobillarda sojae Submerged decaying wood, decaying Licuala longicalycata
(peat swamp) Thailand Plaingam etal. (2005), Pinruan etal.
(2014)
Trichothyriaceae
Lichenopeltella nigroannulata Carex nigra, C. rostrata, Equisetum fluviatile, Eriophorum
angustifolium, Juncus filiformis (lake) Austria Magnes and Hafellner (1991)
Dothideomycetes genera incertae sedis
Alascospora evergladensis Submerged petiole of Nymphaea odorata USA Raja etal. (2010c)

Fungal Diversity
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Table 3 (continued)
Taxon Substrate Distribution References
Ascominuta lignicola Submerged decaying wood, submerged decaying Licuala
longicalycata and Nenga pumila (peat swamp) China, Thailand Ranghoo and Hyde (2000), Pinruan etal.
(2007), Hu etal. (2010b), Pinruan etal.
(2014)
Ascominuta ovalispora Submerged decaying wood (stream) Thailand Hu etal. (2010a)
Bactrodesmium aquaticum Submerged decaying wood (river, reservoir) India Borse (2019)
Bactrodesmium indicum Submerged beech test block (cooling tower) Rao (1983)
Bactrodesmium spilomeum Submerged decaying wood (river) UK Hyde and Goh (1999a)
Clavariopsis aquatica Submerged decaying wood (Alnus glutinosa, Acer pseu-
doplatanus, Corylus avellana, Tilia cordata, Salix spp.,
Populus spp., Betula glandulosa, and Alnus tenuifolia)
(stream), submerged decaying leaves of Coffea arabica
and Hevea brasiliensis (stream), foam (stream, rivulet,
river, spring, lake), water (river, stream), scum (stream),
sediment (dam), stemflow and throughfall of tree canopies
Austria, Canada, Czech Republic,
France, India, Ireland, Japan, Poland,
Portugal, Spain, UK, USA, Venezuela
Ingold (1942), Ingold (1943a), Ranzoni
(1953), Hudson and Ingold (1960),
Baxter (1960), Ingold (1960), Crane
(1968), Iqbal and Webster (1973), Bär-
locher (1987), Sridhar and Kaveriappa
(1988), Czeczuga etal. (1990), Gessner
etal. (1993), Sridhar and Bärlocher
(1993), Voglmayr (1996), Harrington
(1997), Marvanová (2000), Gulis and
Suberkropp (2003), Belliveau and
Bärlocher (2005), Hosoya and Tanaka
(2007), Smits etal. (2007), Sridhar
and Karamchand (2009), Sudheep and
Sridhar (2012), Menéndez etal. (2012),
Raposeiro etal. (2018), Pietryczuk etal.
(2018)
Clavariopsis azlanii Submerged decaying leaves, foam (stream) India, Malaysia, Venezuela Nawawi (1987a), Rajashekhar and Kave-
riappa (2003), Smits etal. (2007)
Clavariopsis brachycladia Submerged leaf of Acer Japan Tubaki (1957)
Clavariopsis bulbosa Foam (stream) India Rajashekhar and Kaveriappa (2003)
Gibbera myrtilli Submerged decaying leaves of Vaccinium macrocarpon, V.
oxycoccus, Ledum groenlandicum (lake) USA Fallah and Shearer (2001)
Lucidascocarpa pulchella Submerged decaying wood (river) Costa Rica, Ecuador, Peru Ferrer etal. (2008), Shearer etal. (2015)
Monodictys abuensis Submerged decaying wood (stream) China Ho etal. (2001)
Monodictys castaneae Submerged decaying wood (river) USA Shearer (1972)
Monodictys cruciseptata Submerged decaying wood USA Schoknecht and Crane (1983)
Monodictys fluctuata Submerged decaying wood Thailand Hu etal. (2010b)
Monodictys levis Submerged decaying wood and bamboo (river) Philippines Cai etal. (2003a)
Monodictys melanocephaloides Submerged decaying wood (reservoir) China, Thailand Goh and Hyde (1999), Hu etal. (2010b)
Monodictys pandani Submerged decaying wood (lake, river) Australia, China Hyde and Goh (1998a)
Monodictys paradoxa Submerged decaying wood Thailand Hu etal. (2010b)
Monodictys peruviana Submerged decaying wood (stream) China Ho etal. (2001)

Fungal Diversity
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Table 3 (continued)
Taxon Substrate Distribution References
Monodictys putredinis Submerged decaying wood (stream), water-filled treeholes Brazil, China, Hungary, South Africa Hyde etal. (1998a), Tsui etal. (2000,
2001d), Gönczöl and Révay (2003)
Monodictys sessilis Submerged decaying wood (river, stream) China Tsui etal. (2000, 2001d)
Monodictys striata Submerged decaying wood (stream) China Tsui etal. (2000)
Monodictys trichocladiopsis Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Ocala scalariformis Submerged decorticated wood (lake) USA Raja etal. (2009a)
Oncopodiella trigonella Water-filled treeholes Hungary Gönczöl and Révay (2003)
Quintaria aquatica Submerged decaying wood (river) Australia Hyde and Goh (1999b)
Quintaria microspora Submerged wood of Fraxinus excelsior (stream) France Zhang etal. (2008a)
Wettsteinina moniliformis Submerged decaying leaf sheath (lake) Denmark Van Ryckegem and Aptroot (2001)
Wettsteinina niesslii Submerged decaying wood and Phragmites australis culms
(lake, river) UK, USA Ingold (1955), Apinis etal. (1972a), Tali-
goola etal. (1972), Shearer (1972)
information about the availability of sequence data of the
listed fungal species.
Some 260 freshwater Leotiomycetes are listed in Table6,
but this figure is underestimated. Based on the results in
Table6, Helotiales has the largest freshwater fungal mem-
bers with 187 species across 12 families including Helotiales
genera incertae sedis. Few taxa under Lauriomycetales (3
species), Leotiales (5 species), Rhytismatales (4 species),
and Thelebolales (2 species) are reported in freshwater habi-
tats. Forty-four species are members of Leotiomycetes fami-
lies incertae sedis while 15 species are under Leotiomycetes
genera incertae sedis. Based on the current revisions of the
Leotiomycetes (Wijayawardene etal. 2020), many species
of freshwater Leotiomycetes have been updated and we refer
to the currently accepted names.
Freshwater Leotiomycetes listed in the Table6 includes
aero-aquatic species, Ingoldian fungi, meiosporic ascomy-
cetes (discomycetes) and hyphomycetes. Helicocentralis
hyalina found in Thailand, Pseudaegerita conifera and P.
websteri both from Spain, were documented as new species
of aero-aquatic fungi within Leotiomycetes and mostly found
on submerged lignocellulose materials (Abdullah etal. 2005;
Sri-indrasutdhi etal. 2015). Aero-aquatic fungi are equipped
with aerially release propagules and air trapping elements
which are distinctive to freshwater hyphomycetes.
Freshwater hypomycetous Leotiomycetes consist of semi-
terrestrial aquatic and Ingoldian fungi which are important
as decomposers with slight difference in substrate spe-
cialization and sporulation. Dactylaria, a semi-terrestrial
hyphomycetes can sporulate underwater and play a role as
decomposers of plant materials (Shearer etal. 2007; Bär-
locher etal. 2008; Tsui etal. 2016). Ingoldian fungi were
mostly isolated from floating foam or associated with leaf
substrates and often found as asexual morphs with unique
conidial shapes i.e. branched forms (Varicosporium) (Camp-
bell etal. 2009), tetraradiate (Alatospora, Articulaspora,
Tetracladium, Tricladium) and sigmoid (Flagellospora)
(Ranzoni 1956). Ingoldian fungi are chiefly leaf decompos-
ers in freshwater ecosystems (Wong etal. 1998a) by produc-
ing exoenzymes that increases leaf litters’ palatability for
invertebrate diets (Bärlocher and Kendrick 1974; Gessner
etal. 2007).
Members of freshwater Leotiomycetes, especially disco-
mycetes, are considered responsible for degrading woody
substrates in water bodies, even though their roles are not as
significant as Sordariomycetes and Dothideomycetes (Wong
etal. 1998a; Hyde etal. 2016a). Wood, plant materials com-
posed of lignin, cellulose, and hemicellulose, are difficult to
break down even in freshwater ecosystems. Hence, coloniza-
tion of freshwater ascomycetes including discomycetes are
capable to secrete enzymes for ligno-cellulose degradation
on woody substrates and play a prominent role in nutrient
cycling and energy transfer between trophic elements in

Fungal Diversity
1 3
freshwater ecosystems (Simonis etal. 2008; Gessner etal.
2010).
Freshwater Leotiomycetes biodiversity and prevalence
can be affected by substrate availability and pollution. A
study has been undertaken on the effect of substrate prefer-
ences on freshwater Leotiomycetes. For nutrition reasons,
Alastospora acuminata and Tetrachaetum elegans were
found mostly on Acacia and Eucalyptus leaves, respectively
(Thomas etal. 1992). Hyphomycetes fungi, Dactylaria spp.
were the only freshwater Leotiomycetes less affected by
organic pollution (Luo etal. 2004). However, other studies
reported that diversity of hyphomycetous and ascomycet-
ous fungi were greatly reduced in polluted environments
(Raviraja etal. 1998; Tsui etal. 2001e). The combination of
macroclimate and microclimate may also affect freshwater
fungal prevalence in general, including Leotiomycetes spe-
cies in certain regions either in temperate or tropical zones
(Webster and Descals 1981). While similar climate in dif-
ferent continents resulted parallel comparison in species
assemblages of freshwater ascomycetes (Raja etal. 2009b),
microclimate such as temperature, pH, and altitude may be
related to Ingoldian fungal prevalence in particular areas
(Chauvet 1991).
Orbiliomycetes
Orbiliaceae was previously assigned to Helotiales but it
was raised to a class rank based on morpho-phylogenetic
analysis. Wijayawardene etal. (2020) accepted twelve gen-
era in Orbiliaceae while Microdochiella (M. fusarioides)
and Mycoceros (M. antennatissimus) are members of
Orbiliales genus incertae sedis and Orbiliomycetes genus
incertae sedis, respectively. Baral etal. (2018) provided a
list of generic names of Orbiliomycetes while Baral etal.
(2020) provided a comprehensive taxonomic account of the
class and its geographical distribution. Orbiliomycetes are
collected in various habitats (hygric, semi-aquatic, xeric).
Nineteen species, in ten genera, (Arthrobotrys, Hyalor-
bilia, Dactylella, Dicranidion, Helicoon, Monacrosporium,
Orbilia, Orbiliella, Trinacrium, Vermispora), are listed in
Table7.
Pezizomycetes
Pezizomycetes thrive mostly in terrestrial habitats, such
as burnt land, deserts, forests, sand, steppe, and on dif-
ferent substrates (dung, leaves, soil, wood) (Hansen and
Pfister 2006; Pfister etal. 2008; Ekanayaka etal. 2018).
However, Pezizomycetes are rarely reported from fresh-
water habitats. Jones etal. (2014a) recorded eight species
from freshwater. Currently, ten species are included in the
Freshwater Ascomycetes Database (http:// fungi. life. illin
ois. edu/). We reviewed the number of freshwater fungi in
this class (Table8), and there are 13 species from four
families. All known freshwater Pezizomycetes are found
on woody substrates. Within these aquatic species, almost
half of them from Pachyella and Psilopezia. Species of
both genera are usually from wet habitats, however not
all species were found and exclusively described on the
water-soaked wood (Pfister 1972; Pfister 1973a, b). Sac-
cobolus beckii is usually reported on animal dung (Doveri
2014; Melo etal. 2014). One interesting account reports
them from water cooling tower (Udaiyan 1989). In addi-
tion, another unknown Peziza species has been reported
Fig. 1 Major families of Dothideomycetes taxa from freshwater habitats

Fungal Diversity
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Fig. 2 Freshwater Dothideomycetes. a, b Aliquandostipite
khaoyaiensis (MFLU 18-1573), c, d Ascagilis submersa (MFLU
18-1527), c Ascoma on host with associated appendages (arrow),
e, f Caryospora aquatica (MFLU 18-1202), g, h Parabambusicola
aquatica (MFLU 18-1524), i, j Longipedicellata aptrootii (MFLU
18-1000), k, l Caryospora submersa (MFLU 18-1539), m, n Heli-
cotruncatum palmigenum (MFLU 15-2715), o, p Shrungabeeja
aquatica (MFLU 18-0978), q, r Pleopunctum pseudoellipsoideum
(HKAS 102147), s, t Pseudoxylomyces elegans (MFLU 17-1712).
Scale bar: b, d, f, h, l, r = 20; j, n = 10; p = 30; t = 50

Fungal Diversity
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Fig. 3 Freshwater Eurotiomycetes. a–f Pseudobactrodesmium spp. g–h Minimelanolocus sp. i–k Thysanorea thailandensis. l,m Veronaea bot-
ryosa. n–p Aculeata aquatica. Scale bar: a–d, g, n = 200μm, e, f = 20μm, h, m = 10μm, i = 100μm, j, k = 5μm, o, p = 15μm

Fungal Diversity
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Table 4 Classification of freshwater Eurotiomycetes with information on the type of habitats, substrates, and geographical distribution of fungal
taxa
Taxon Substrate Distribution References
EUROTIOMYCETES
Chaetothyriomycetidae
Chaetothyriales
Cyphellophoraceae
Anthopsis microspora Scum (stream) Japan Ando and Tubaki (1985)
Cyphellophora oxyspora Water (swimming pool) Germany Feng etal. (2014)
Cyphellophora catalaunica Sediments (river) Spain Crous etal. (2013)
Kumbhamaya jalapriya Submerged roots of Hopea ponga (stream) India Nair and Bhat (2002)
Herpotrichiellaceae
Aculeata aquatica Submerged decaying wood (river) Thailand Dong etal. (2018)
Exophiala cancerae Water Germany de Hoog etal. (2011)
Exophiala castellani Drinking water Germany de Hoog etal. (2011)
Exophiala dermatitidis Groundwater, water (river) Poland, Slovenia Novak Babič etal.
(2016), Pietryczuk
etal. (2018)
Exophiala equina Drinking water Netherlands Ávila etal. (2005)
Exophiala lecanii-corni Tapwater Slovenia Novak Babič etal.
(2016)
Exophiala mesophila Waterlines, tapwater USA, Slovenia Porteous etal. (2003),
Novak Babič etal.
(2016)
Exophiala oligosperma Tapwater Slovenia Novak Babič etal.
(2016)
Exophiala phaeomuriformis Tapwater Slovenia Novak Babič etal.
(2016)
Exophiala pisciphila Swimming pool Germany McGinnis and Ajello
(1974)
Exophiala psychrophila From Atlantic salmon smolt – Pedersen and Langvad
(1989)
Exophiala salmonis Cut-throat trout (Oncorhynchus clarkii), fingerling
lake trout (Salvelinus namaycush)Alberta, Canada Carmichael (1967)
Exophiala angulospora Drinking well water Japan Iwatsu etal. (1991)
Exophiala lacus Lake Netherlands de Hoog etal. (2011)
Exophiala opportunistica Drinking water Germany de Hoog etal. (2011)
Exophiala xenobiotica Tapwater Slovenia Novak Babič etal.
(2016)
Minimelanolocus aquatilis Submerged decaying wood (stream) Brazil Fiuza etal. (2017c)
Minimelanolocus clavatus Submerged decaying wood (stream) China Wan etal. (2021)
Minimelanolocus nujiangensis Submerged decaying wood (river) China Wan etal. 2021)
Minimelanolocus submersus Submerged decaying wood (lake) China Hyde etal. (2016b)
Phialophora reptans Drinking water Germany Feng etal. (2014)
Phialophora aurantiaca Water UK Beyma (1940)
Phialophora benthica Sediment (lake) Canada Millar (1990)
Rhinocladiella amoena Submerged decaying twig Spain Madrid etal. (2016)
Rhinocladiella similis Groundwater, tapwater Slovenia Novak Babič etal.
(2016)
Thysanorea melanica Submerged decaying wood (stream) China Liu etal. (2015b)
Thysanorea obscura Submerged decaying wood (stream) China Liu etal. (2015b)
Thysanorea papuana Submerged decaying wood (stream) Thailand Wang etal. (2019)
Thysanorea thailandensis Submerged decaying wood (river) Thailand Dong etal. (2018)

Fungal Diversity
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Table 4 (continued)
Taxon Substrate Distribution References
Thysanorea aquatica Submerged decaying wood (river) Thailand Dong etal. (2018)
Thysanorea asiatica Submerged decaying wood (stream) China Liu etal. (2015b)
Thysanorea curvata Submerged decaying wood (stream) China Liu etal. (2015b)
Thysanorea nonramosa Submerged decaying wood (stream) Thailand Wang etal. (2019)
Thysanorea uniseptata Submerged decaying wood (stream) Thailand Hyde etal. (2019)
Thysanorea yunnanensis Submerged decaying wood (stream) China, Thailand Liu etal. (2015b)
Veronaea aquatica Submerged decaying wood (stream) China Chandrasiri etal. (2021)
Veronaea botryosa Submerged decaying wood (river) Thailand Dong etal. (2018)
Chaetothyriales genera incertae sedis
Uncispora sinensis Submerged decaying leaves (river) China Yang etal. (2011)
Uncispora wuzhishanensis Submerged decaying leaves (stream) China Liu etal. (2018b)
Verrucariales
Adelococcaceae
Sagediopsis lomnitzensis Stream and lake margin Europe Orange (2002)
Verrucariaceae
Agonimia allobata Stream Lithuania Krzewicka etal. (2017)
Agonimia tristicula Stream UK Motiejunaite (2003)
Clavascidium lacinulatum Stream UK Gilbert and Giavarini
(1997)
Catapyrenium lachneum Stream UK Gilbert and Giavarini
(1997)
Catapyrenium squamulosum River Watershed Canada Maloles etal. (2018)
Catillaria cf. chalybeia Stream Gwynedd Gilbert and Giavarini
(1997)
Dermatocarpon arnoldianum River and stream Europe Pentecost (1977)
Dermatocarpon intestiniforme Stream UK Gilbert and Giavarini
(1997), Shivarov etal.
(2018)
Dermatocarpon leptophyllodes Stream UK Gilbert and Giavarini
(1997)
Dermatocarpon luridum Stream Poland Krzewicka etal. (2017)
Dermatocarpon meiophyllizum Stream UK Gilbert and Giavarini
(1997)
Dermatocarpon miniatum River Bulgaria Shivarov etal. (2018)
Dermatocarpon rivulorum Submerged siliceous rock (stream, river) Bulgaria, Italy, UK Nascimbene etal.
(2007) Gilbert and
Giavarini (1997),
Shivarov and Lőkös
(2015)
Endocarpon adscendens Stream UK Gilbert and Giavarini
(1997)
Hydropunctaria orae Stream – Orange (2013), Thüs
etal. (2014)
Hydropunctaria rheitrophila Stream, submerged carbonic rock (river, stream) Australia, Italy,
Poland Nascimbene etal.
(2007), Krzewicka
etal. (2017) Gilbert
and Giavarini (1997),
McCarthy (2002),
Matura and Krzewicka
(2015)
Hydropunctaria scabra Siliceous rock (stream) Bulgaria Shivarov etal. (2017)
Muellerella pygmaea Stream and lake margin Europe Orange (2002)
Phaeospora lemaneae Parasitic on Lemanea sp. UK Ingold (1955)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Placopyrenium cinereoatratum Siliceous rocks (brook, lake, river, stream) Bulgaria, Finland,
Norway, Sweden,
UK
Orange (2009)
Placopyrenium formosum Rock (river) Finland, Iceland, UK Orange (2009)
Polyblastia ardesiaca – Italy Nascimbene and Nimis
(2006)
Psoroglaena stigonemoides Twig (waterfall) Canada Björk etal. (2009)
Sporodictyon cruentum River and Streams Alpine Australia, Italy,
UK Gilbert and Giavarini
(1997), McCarthy
(2002), Nascimbene
and Nimis (2006),
Hafellner (2010)
Staurothele acarosporoides Nature reserves Vietnam Gueidan etal. (2014)
Staurothele arenaria Nature reserves Vietnam Gueidan etal. (2014)
Staurothele chlorospora Nature reserves Vietnam Gueidan etal. (2014)
Staurothele clopimoides River Italy, USA Thomson (1991), Nas-
cimbene and Nimis
(2006)
Staurothele fauriei Nature reserves Vietnam Gueidan etal. (2014)
Staurothele fissa Lakes and Streams Europe Lindsay (1856), Gilbert
and Giavarini (1997),
Nascimbene and
Nimis (2006), Shiv-
arov (2017)
Staurothele fuscocuprea – Italy Nascimbene and Nimis
(2006)
Staurothele honghensis Nature reserves Vietnam Gueidan etal. (2014)
Staurothele kwapiensis Nature reserves Vietnam Gueidan etal. (2014)
Staurothele muliensis Nature reserves Vietnam Gueidan etal. (2014)
Staurothele ochroplaca Nature reserves Vietnam Gueidan etal. (2014)
Staurothele pachystroma Nature reserves Vietnam Gueidan etal. (2014)
Staurothele paraguayensis Nature reserves Vietnam Gueidan etal. (2014)
Staurothele sinensis Nature reserves Vietnam Gueidan etal. (2014)
Staurothele succedens Stream Italy, UK Gilbert and Giavarini
(1997), Nascimbene
and Nimis (2006)
Staurothele yunnanica Nature reserves Vietnam Gueidan etal. (2014)
Thelidium aethioboloides Splash water zone (creek) Germany Thüs and Nascimbene
(2008)
Thelidium aquaticum Stream Poland Krzewicka etal. (2017)
Thelidium carbonaceum Nature reserves Australia McCarthy and Kantvilas
(2020)
Thelidium circumspersellum Limestone (rivulet) Poland, Romania Thüs and Nascimbene
(2008), Matura etal.
(2017)
Thelidium epomphalum Creek and River Europe Thüs and Nascimbene
(2008)
Thelidium fontigenum Waterfall, amphibious on limestone, lime-containing
sandstone, silt-impregnated wood, and on mica
schist
Austria, Germany,
Switzerland Thüs and Nascimbene
(2008)
Thelidium klementii Calcareous rocks Germany Thüs and Nascimbene
(2008)
Thelidium lahmianum Inundated granite boulders Poland Thüs and Nascimbene
(2008)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Thelidium methorium Rocks (brook, creek, river) Austria, Czech Repub-
lic, Estonia, Italy,
Norway, Slovakia,
Sweden, Switzerland
Thüs and Nascimbene
(2008)
Thelidium minutulum splash water zone, limestone or
basic siliceous rocks (creeks, rivers) Austria, Czech
Republic, Germany Thüs and Nascimbene
(2008)
Thelidium nigricans Siliceous rocks (creek's splash water zone) Switzerland Thüs and Nascimbene
(2008)
Thelidium papulare Rocks (creek) Austria, Germany Thüs and Nascimbene
(2008)
Thelidium pertusatii Rocks (river) Italy Thüs and Nascimbene
(2008)
Thelidium pluvium Sandstone, schist (creek, river) Austria, Germany Thüs and Nascimbene
(2008)
Thelidium rehmii Sandstone rocks (creek) Germany Thüs and Nascimbene
(2008)
Thelidium rivulicolum Carbonaceous rock (creek, stream) France, Germany Thüs and Nascimbene
(2008)
Thelidium spadanum – Europe Thüs and Nascimbene
(2008)
Thelidium sublacteum – Europe Thüs and Nascimbene
(2008)
Thelidium submethorium Rocks (creek, river) Italy, Germany Thüs and Nascimbene
(2008)
Thelidium suzaeanum Limestone rock (river) Slovakia Thüs and Nascimbene
(2008)
Thelidium zahlbruckneri Stream Poland Krzewicka etal. (2017)
Thelidium zwackhii Stream Italy, Germany,
Poland Thüs and Nascimbene
(2008), Krzewicka
etal. (2017)
Verrucaria aethiobola Stream Italy, UK Pentecost (1977), Nas-
cimbene and Nimis
(2006)
Verrucaria alpicola Pebble (river) Bulgaria Shivarov etal. (2017)
Verrucaria amnica Mountain stream New Zealand McCarthy and Johnson
(1997)
Verrucaria aquatilis Stream, submerged carbonatic and siliceous rock
(stream) Italy, Poland Nascimbene etal.
(2007), Krzewicka
etal. (2017)
Verrucaria austroalpina Submerged rocks (stream) Australia Krzewicka etal. (2017)
Verrucaria basaltica Streams and lakes Poland Krzewicka (2012)
Verrucaria calkinsiana River Watershed Canada Maloles etal. (2018)
Verrucaria capitulata Submerged rocks (stream) Japan Harada (2012)
Verrucaria cernaensis Stream Poland Krzewicka etal. (2017)
Verrucaria consociata Stream UK Orange (2015)
Verrucaria conturmatula Stream and lake margin Europe Orange (2002)
Verrucaria cootapatambensis Sandstone boulder in snow-melt creek Australia McCarthy (2002)
Verrucaria craterigera Friable rocks by stream in a gorge Japan Harada (2012)
Verrucaria devensis Stones (river) Germany, UK Orange (2014)
Verrucaria dolosa Stream Poland, UK Gilbert and Giavarini
(1997), Matura and
Krzewicka (2015)
Verrucaria elaeina Stream Poland Matura and Krzewicka
(2015)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Verrucaria elaeomelaena Submerged carbonatic rock (stream) Italy Nascimbene etal.
(2007)
Verrucaria funckii Stream, submerged siliceous rock (stream) Italy, Poland Nascimbene etal.
(2007), Krzewicka
etal. (2017)
Verrucaria glaucina Stream UK Gilbert and Giavarini
(1997)
Verrucaria gongshanensis Non-calcareous rocks in a stream China Harada and Wang
(2008)
Verrucaria honghensis Riverside rocks China Harada and Wang
(2008)
Verrucaria humida Wet stones in bed of streamlet in Picea abiesforest Germany Krzewicka etal. (2017)
Verrucaria hydrela Inundated and submerged siltstone UK Orange (2013)
Verrucaria hydrophila Shaded stone by stream in woodland UK Orange (2013)
Verrucaria igii River Alpine Australia McCarthy (2002)
Verrucaria inconstans Aquatic limestone Australia McCarthy (1995)
Verrucaria latebrosa Submerged siliceous rock (stream) Germany, Italy, Slova-
kia, Switzerland Nascimbene etal.
(2007), Thüs and
Nascimbene (2008)
Verrucaria luchunensis Riverside rocks China Harada and Wang
(2008)
Verrucaria maas-geesterani Lake Switzerland Servít (1954)
Verrucaria madida Sloping siliceous rocks, shallowly submerged in
stream France Harada and Wang
(2008)
Verrucaria margacea Freshwater Australian McCarthy (1995)
Verrucaria morosa River Alpine Australia McCarthy (2002)
Verrucaria mundula Aquatic slate Australia McCarthy (1995)
Verrucaria nodosa Unshaded rock in stream UK Orange (2013)
Verrucaria nujiangensis Submerged stones in stream China Harada and Wang
(2008)
Verrucaria pachyderma Stream Australia, Italy,
Europe Nascimbene and Nimis
(2006), Krzewicka
(2012)
Verrucaria phaeoderma Aquatic basalt Australia McCarthy (1995)
Verrucaria placida Stone in shaded streamlet in woodland Norway, Germany,
UK Orange (2013)
Verrucaria praetermissa Freshwater Australian McCarthy (1995)
Verrucaria puncticulata Inundated and submerged siltstone Australia McCarthy (1995)
Verrucaria rosula Siliceous rocks and stones (reservoir, stream) France, UK Orange (2013)
Verrucaria ruttneri River Alpine Australia McCarthy (2002)
Verrucaria rhizicola Bark of roots ofAlnus glutinosairregularly sub-
mersed in stream France Lumbsch etal. (2011)
Verrucaria sublobulata Stream Poland Krzewicka etal. (2017)
Verrucaria submauroides Stream Poland Krzewicka etal. (2017)
Verrucaria submersella Stream Poland Krzewicka etal. (2017)
Verrucaria takagoensis Non-calcareous rocks submerged in stream Japan Harada (2001)
Verrucaria tasmanica Aquatic dolerite Australia McCarthy (1995)
Verrucaria tholocarpa Aquatic limestone Australia McCarthy (1995)
Verrucaria yoshimurae Rock (gorge) Japan Harada (2011)
Willeya australis Nature reserves Vietnam Gueidan etal. (2014)
Willeya diffractella Nature reserves Vietnam Gueidan etal. (2014)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Willeya fusca Nature reserves Vietnam Gueidan etal. (2014)
Willeya iwatsukii Nature reserves, river Japan, Vietnam Harada (1992), Gueidan
etal. (2014)
Willeya japonica Nature reserves Vietnam Gueidan etal. (2014)
Willeya laevigata Nature reserves Vietnam Gueidan etal. (2014)
Willeya malayensis Nature reserves Vietnam Gueidan etal. (2014)
Willeya microlepis Nature reserves Vietnam Gueidan etal. (2014)
Willeya pallidipora Nature reserves Vietnam Gueidan etal. (2014)
Willeya protrudens Nature reserves Vietnam Gueidan etal. (2014)
Willeya rimosa Nature reserves Vietnam Gueidan etal. (2014)
Chaetothyriomycetidae family incertae sedis
Rhynchostomataceae
Rhynchostomapyriforme On rotten wood in river Dominica Smith (1901)
Eurotiomycetidae
Arachnomycetales
Arachnomycetaceae
Arachnomyces pilosus River sediment Spain Gené etal. (1994)
Eurotiales
Aspergillaceae
Aspergillus aculeatus Freshwater UK Kinsey etal. (1998)
Aspergillus affinis Submerged riparian decomposing leaves Italy Davolos etal. (2012)
Aspergillus candidus River Egypt Abdel Hameed etal.
(2008)
Aspergillus clavatus Freshwater UK Kinsey etal. (1998)
Aspergillus flavus Freshwater Korea Heo etal. (2019)
Aspergillus fumigatus Water (river) Poland Pietryczuk etal. (2018)
Aspergillus israelensis Isolated from evaporation pond Israel Piontelli etal. (2019)
Aspergillus japonicus Freshwater Korea Heo etal. (2019)
Aspergillus niger Water (river) Korea, Poland Pietryczuk etal. (2018),
Heo etal. (2019)
Aspergillus ochraceus River Egypt Abdel Hameed etal.
(2008)
Aspergillus parasiticus River Egypt Abdel Hameed etal.
(2008)
Aspergillus parvulus Freshwater UK Kinsey etal. (1998)
Aspergillus sydowii River Egypt Abdel Hameed etal.
(2008)
Aspergillus tamarii River Egypt Abdel Hameed etal.
(2008)
Aspergillus terreus Lake India Bandh etal. (2012)
Aspergillus tubingensis Freshwater Korea Heo etal. (2019)
Aspergillus versicolor Lake India Bandh etal. (2012)
Aspergillus wentii Lake India Bandh etal. (2012)
Monascus pallens Sediment Iraq Cannon etal. (1995)
Monascus sanguineus Sediment Iraq Cannon etal. (1995)
Penicillium acidum Plant debris in pond Korea Wanasinghe etal.
(2018)
Penicillium aquaticum Plant debris in pond Korea Wanasinghe etal.
(2018)
Penicillium aurantiogriseum Freshwater UK Kinsey etal. (1998)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Penicillium brasilianum Freshwater Korea Heo etal. (2019)
Penicillium brevicompactum Lake Turkey Asan etal. (2003)
Penicillium capsulatum River Egypt Abdel Hameed etal.
(2008)
Penicillium camemberti Lake India Bandh etal. (2012)
Penicillium chrysogenum Lake India Bandh etal. (2012)
Penicillium citrinum River Egypt, UK Kinsey etal. (1998),
Abdel Hameed etal.
(2008)
Penicillium citreonigrum River Egyptian Abdel Hameed etal.
(2008)
Penicillium commune Lake India, UK Kinsey etal. (1998),
Bandh etal. (2012)
Penicillium corylophilum Freshwater UK Kinsey etal. (1998)
Penicillium crustosum Freshwater Korea Heo etal. (2019)
Penicillium dimorphosporum Lake India Bandh etal. (2012)
Penicillium echinulatum Freshwater UK Kinsey etal. (1998)
Penicillium echinulonalgiovense Freshwater Korea Pangging etal. (2021a)
Penicillium expansum Freshwater Korea Heo etal. (2019)
Penicillium funiculosus Water (lake, river) India, Poland Bandh etal. (2012),
Pietryczuk etal.
(2018)
Penicillium glabrum Lake Turkey, UK Kinsey etal. (1998),
Asan etal. (2003)
Penicillium globosum Freshwater Korea Pangging etal. (2021a)
Penicillium griseofulvum Freshwater UK Kinsey etal. (1998)
Penicillium griseoroseum Lake Turkey Asan etal. (2003)
Penicillium guanacastense Stream soil Korea Mun etal. (2020)
Penicillium hirsutum Freshwater UK Kinsey etal. (1998)
Penicillium janczewskii Freshwater UK Kinsey etal. (1998)
Penicillium lilacinum Lake India Bandh etal. (2012)
Penicillium miczynskii Lake Turkey Asan etal. (2003)
Penicillium ochrosalmoneum Freshwater UK Kinsey etal. (1998)
Penicillium onobense Freshwater Korea Pangging etal. (2021a)
Penicillium olivicolor Lake India Bandh etal. (2012)
Penicillium oxalicum Freshwater Korea Heo etal. (2019)
Penicillium pinophilum Freshwater UK Kinsey etal. (1998)
Penicillium piscarium Freshwater Korea Heo etal. (2019)
Penicillium piscium Internal organs of freshwater Osteichthyes – Reichenbach-Klinke
(1956)
Penicillium puberulum Lake Turkey Asan etal. (2003)
Penicillium purpureogenum Freshwater UK Kinsey etal. (1998)
Penicillium raistrickii Freshwater UK Kinsey etal. (1998)
Penicillium saturniforme River soil Korea Mun etal. (2020)
Penicillium scabrosum Stream soil, water Korea Mun etal. (2020),
Nguyen etal. (2020b)
Penicillium simplicissimum Freshwater UK Kinsey etal. (1998)
Penicillium solitum Freshwater UK Kinsey etal. (1998)
Penicillium spinulosum Freshwater UK Kinsey etal. (1998)
Penicillium variabile River Egypt Abdel Hameed etal.
(2008)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Penicillium verrucosum Lake Turkey Asan etal. (2003)
Penicillium waksmanii Freshwater UK Kinsey etal. (1998)
Penicillium wandoens Freshwater Korea Hyde etal. (2019)
Penicillium yunnanense Freshwater Korea Pangging etal. (2021a)
Thermoascaceae
Paecilomyces lilacinus Freshwater UK Kinsey etal. (1998)
Trichocomaceae
Talaromyces flavus Submerged decaying leaf litter (swamp), sand (creek) Australia, USA Pitt (1979), Shearer and
Crane (1986)
Talaromyces gwangjuensis Freshwater (reservoir) Korea Nguyen etal. (2021)
Talaromyces koreanus Freshwater (reservoir) Korea Nguyen etal. (2021)
Talaromyces liani Freshwater Korea Pangging etal. (2019)
Talaromyces pinophilus Freshwater Korea, UK Kinsey etal. (1998),
Heo etal. (2019)
Talaromyces siamensis Freshwater Korea Lim etal. (2021)
Talaromyces teleomorphus Freshwater (reservoir) Korea Nguyen etal. (2021)
Talaromyces versatilis Freshwater Korea Heo etal. (2019)
Onygenales
Arthrodermataceae
Arthroderma insingulare Water (river) Poland Pietryczuk etal. (2018)
Microsporum gypseum Water (river) Poland Pietryczuk etal. (2018)
Trichophyton mentagrophytes Water Poland Czeczuga (1996)
Trichophyton violaceum Water (river) Poland Pietryczuk etal. (2018)
Onygenaceae
Aphanoascus keratinophilus Water (river) Poland Pietryczuk etal. (2018)
Chrysosporium fluviale Sediment (river) Spain Vidal etal. (2000)
Chrysosporium minutisporosum Sediment (river) Spain Vidal etal. (2002)
Chrysosporium pseudomerdarium Sediment Korea Goh etal. (2021a)
Chrysosporium sanyaense Sediment Korea Goh etal. (2021a)
Chrysosporium submersum Sediment (river) Spain Vidal etal. (2002)
Chrysosporium undulatum Sediment (river) Spain Vidal etal. (1999)
Mycocaliciomycetidae
Mycocaliciales
Mycocaliciaceae
Chaenothecopsis debilis River watershed Canada Maloles etal. (2018)
Chaenothecopsis haematopus Rainwater from building roofs Poland Czeczuga and Orłowska
(1997)
Chaenothecopsis pusiola River watershed Canada Maloles etal. (2018)
Chaenothecopsis savonica River watershed Canada Maloles etal. (2018)
Phaeocalicium curtisii River watershed Canada Maloles etal. (2018)
Phaeocalicium polyporaeum River watershed Canada Maloles etal. (2018)
Phaeocalicium populneum River watershed Canada Maloles etal. (2018)
Sphinctrina anglica River watershed Canada Maloles etal. (2018)
Stenocybe pullatula River watershed Canada Maloles etal. (2018)
Sclerococcomycetidae
Sclerococcales
Dactylosporaceae
Cylindroconidiis aquaticus Freshwater Thailand Yu etal. (2018)
Pseudobactrodesmium aquaticum Submerged decaying wood (stream) China Dong etal. (2020a)
Pseudobactrodesmium chiangmaiensis Submerged decaying wood (stream) Thailand Dong etal. (2020a)

Fungal Diversity
1 3
Table 4 (continued)
Taxon Substrate Distribution References
Pseudobactrodesmium longisporum Submerged decaying wood (stream) Australia, Thailand Vijaykrishna and Hyde
(2006), Dong etal.
(2020a)
Pseudobactrodesmium stilboideum Submerged decaying wood (stream, river) Brazil, Philippines,
Thailand Cai etal. (2003a), Boon-
mee etal. (2021)
from a water-cooling tower (Eaton and Jones 1971a). Hu
etal. (2012c) proposed a new genus Aquapeziza and placed
in Pezizaceae based on the morphology and phylogeny of
Aquapeziza globispora on submerged wood. The opercu-
late asci, which is main feature of Pezizomycetes, are obvi-
ous in all freshwater Pezizomycetes except A. globispora
with no distinct ascus (Hu etal. 2012c). Peziza oliviae is
the latest report associated with freshwater habitat within
Pezizomycetes (Frank 2013).
Sordariomycetes
The Sordariomycetes is one of the largest classes of Asco-
mycota that comprises a highly diverse range of fungi mainly
characterized by perithecial ascomata and inoperculate uni-
tunicate asci (Maharachchikumbura etal. 2016; Hyde etal.
2020d). In freshwater habitats, Sordariomycetes is one of the
major and important fungal groups with a vital role in eco-
systems. Members of this class are known to produce bioac-
tive compounds (Poch etal. 1992; Jones etal. 2014a). The
latest systematic study of freshwater Sordariomycetes was
carried out by Luo etal. (2019) with taxonomic accounts for
each genus. Furthermore, all taxa recorded from freshwater
habitats are outlined with important ecological (freshwater
distribution) and taxonomic (morphology, holotype, avail-
able molecular data) notes of each species (Luo etal. 2019;
Calabon etal. 2020b).
Luo etal. (2019) included six subclasses, 30 orders, 56
families and 160 genera of Sordariomycetes thriving in
freshwater habitats, and representative taxa are illustrated
in Fig.5. From the review of published works on fresh-
water Sordariomycetes, a total 823 freshwater species were
reported from freshwater habitats, and distributed in 40
orders, 85 families, and 298 genera (Table9, Fig.6). Among
the 40 orders, Chaetosphaeriales has the highest number of
taxa with 149 freshwater species recorded.
The use of DNA sequence data coupled with phylo-
genetic analyses has significantly improved our under-
standing of the phylogeny and evolution of freshwater
ascomycetes (Vijaykrishna etal. 2006). Most of the new
taxa of freshwater fungi published in the past decade are
supported with molecular data (Pinruan etal. 2010; Liu
and Cai 2013; Su etal. 2016a; Yang etal. 2018d; Calabon
etal. 2021a). However, there are still taxa of freshwa-
ter fungi with no sequence data (Monteiro etal. 2017a,
2017b). The taxonomic placements of many freshwater
Sordariomycetes are currently uncertain and suffers from
lack of living cultures and type sequences. Therefore, to
better understand the phylogeny of freshwater Sordario-
mycetes in a natural classification system, further investi-
gations are needed. Future studies should put high priority
to obtain living cultures.
Freshwater lichens
Lichens are one of the most successful lifestyles among
fungi (Honegger 2009). Lichens are recently redefined
as “a self-sustaining ecosystem formed by the interaction
of an exhabitant fungus and an extracellular arrangement
of one or more photosynthetic partners and an indeter-
minate number of other microscopic organisms” (Hawk-
sworth and Grube 2020). The lichen forming fungi are
mainly terrestrial but can occur also in aquatic and semi-
aquatic habitats in both freshwater and marine environ-
ments (Kulikova etal. 2008). Santesson (1939) and Apt-
root and Seaward (2003) stated that freshwater lichens
are amphibious organisms as most of them are submerged
for particular period. The mycobiont is ascomycetes in
most freshwater lichens while the photobiont can be
commonly coccoid green alga or a cyanobacterium (Nash
1996; Honegger 1997; Nascimbene and Nimis 2006).
Verrucariales and Lichinales are frequently recorded
freshwater species from predominantly submerged zone
(Thüs etal. 2014). The species are mostly found in the
surface of water since the photobiont needs direct sun-
light (Nash 1996; Honegger 1997). However, their verti-
cal distribution is highly influenced by transparency of
the water as some species found in the deeper lakes and
rivers which has high water clarity (e.g., Hydropunc-
taria rheithrophila) (Thüs etal. 2014). Species have been
recorded on rocks, pebbles and wood and provide favora-
ble host surface for many lichenicolous fungi (Aptroot
and Thüs 2011; Thüs etal. 2014).
Freshwater lichens have worldwide distribution, but
this group are poorly studied (Motiejunaite 2003; Matura
and Krzewicka 2015). The diversity of freshwater lichens

Fungal Diversity
1 3
Table 5 Classification of freshwater Laboulbeniomycetes with information on the type of habitats, hosts, and geographical distribution of fungal
taxa
Taxon Substrate (Family of host-insects) Distribution Reference
ASCOMYCOTA
LABOULBENIOMYCETES
Laboulbeniales
Ceratomycetaceae
Autoicomyces acuminatus Hydrophilidae USA Thaxter (1908)
Autoicomyces aquatilis Hydrochidae, Dytiscidae Cuba, Europe, USA Majewski (2008)
Autoicomyces bicornis Hydrophilidae Argentina, South Korea Lee and Kim (1990)
Autoicomyces chinensis Hydrophilidae China Shen and Ye (2006)
Autoicomyces contortus Hydrophilidae USA Tavares (1985)
Autoicomyces costaricensis Hydrophilidae USA Tavares (1985)
Autoicomyces crassus Hydrophilidae Europe Santamaria (2003)
Autoicomyces falcatus Hydrophilidae Brazil, Trinidad, USA Goldmann and Weir (2018)
Autoicomyces falcifer Hydrophilidae Asia, Europe, Sierra Leone Rossi etal. (2019)
Autoicomyces fragilis Hydrophilidae Italy Spegazzini (1915)
Autoicomyces furcatus Hydrophilidae Cameroon, USA Thaxter (1931)
Autoicomyces helocharalis Hydrophilidae Asia, Cameroon Thaxter (1931)
Autoicomyces humilis Hydrophilidae Asia, Italy, USA Thaxter (1931)
Autoicomyces incurvatus Hydrophilidae Philippines Thaxter (1931)
Autoicomyces latens Hydrophilidae China, Philippines Sugiyama and Shazawa
(1977)
Autoicomyces longipes Hydrophilidae Philippines Thaxter (1931)
Autoicomyces marginalis Hydrophilidae Argentina Tavares (1985)
Autoicomyces marginicola Hydrophilidae Asia, Spain Santamaria (2003)
Autoicomyces minusculus Hydrophilidae South and North America Tavares (1985)
Autoicomyces occidentalis Hydrophilidae USA Tavares (1985)
Autoicomyces occultatus Hydrophilidae USA Thaxter (1931)
Autoicomyces ornithocephalus Hydrophilidae USA Thaxter (1908)
Autoicomyces recurvatus Hydrophilidae USA Goldmann and Weir (2018)
Autoicomyces semiopacus Hydrophilidae Trinidad Tavares (1985)
Autoicomyces siamensis Hydrophilidae Thailand Sugiyama and Phanichapol
(1984)
Autoicomyces trinitatis Hydrophilidae Trinidad Thaxter (1931)
Ceratomyces ansatus Hydrophilidae South and North America Weir and Rossi (2001)
Ceratomyces brasiliensis Hydrophilidae Brazil Thaxter (1908)
Ceratomyces californicus Hydrophilidae USA Thaxter (1908)
Ceratomyces camptosporus Hydrophilidae USA Thaxter (1896)
Ceratomyces cladophorus Hydrophilidae Ecuador, USA Bernardi etal. (2014)
Ceratomyces confusus Hydrophilidae USA Thaxter (1896)
Ceratomyces curvatus Hydrophilidae Venezuela Thaxter (1908)
Ceratomyces filiformis Hydrophilidae Central, North, South America Thaxter (1931)
Ceratomyces floridanus Hydrophilidae USA Thaxter (1908)
Ceratomyces hyalinus Hydrochidae China Shen etal. (2009)
Ceratomyces intermedius Hydrophilidae Argentina Thaxter (1931)
Ceratomyces longicornis Hydrophilidae USA Thaxter (1931)
Ceratomyces mexicanus Hydrophilidae Mexico Thaxter (1908)
Ceratomyces mirabilis Hydrophilidae Central and South America, USA Goldmann and Weir (2018)
Ceratomyces platensis Hydrophilidae Argentina Thaxter (1931)
Ceratomyces procerus Hydrophilidae Brazil Thaxter (1908)
Ceratomyces pyrenaeus Hydrophilidae Spain Santamaria (1999)

Fungal Diversity
1 3
Table 5 (continued)
Taxon Substrate (Family of host-insects) Distribution Reference
Ceratomyces rhizophorus Hydrophilidae Argentina Thaxter (1931)
Ceratomyces spiniger Hydrophilidae Costa Rica, Mexico Thaxter (1908)
Ceratomyces spiralis Hydrophilidae Spain, Poland Majewski (2008)
Ceratomyces sternolophi Hydrophilidae Indonesia Thaxter (1931)
Ceratomyces ventriosus Hydrophilidae Argentina Thaxter (1931)
Drepanomyces malayanus Hydrophilidae Asia (Bameul 1993)
Eusynaptomyces africanus Hydrophilidae East Africa Thaxter (1931)
Eusynaptomyces benjaminii Hydrophilidae Germany, Poland Majewski (2008)
Eusynaptomyces borealis Hydrophilidae USA Thaxter (1931)
Eusynaptomyces cornutus Hydrophilidae Spain Santamaria (2006)
Eusynaptomyces enochri Hydrophilidae Europe Majewski (2008)
Eusynaptomyces hydrobii Hydrophilidae Poland Majewski (2008)
Helodiomyces elegans Dryopidae Algeria, Europe De Kesel and Haelewaters
(2014)
Phurmomyces obtusus Hydrophilidae China, Indonesia Shen and Ye (2006)
Plectomyces gracilis Hydrophilidae Philippines, China Shen and Ye (2006)
Rhynchophoromyces amphiopsis Hydrophilidae Cameroon Thaxter (1931)
Rhynchophoromyces anacaenae Hydrophilidae Europe Rossi etal. (2019)
Rhynchophoromyces denticulatus Hydrophilidae Asia, Guam Shen and Ye (2006)
Rhynchophoromyces elephanthnus Hydrophilidae USA Shen and Ye (2006)
Rhynchophoromyces helophorii Hydrophilidae Spain Santamaria (1999)
Rhynchophoromyces minor Hydrophilidae Argentina, Philippines, USA Goldmann and Weir (2018)
Rhynchophoromyces rostratus Hydrophilidae Asia, North, Central and South
America, Europe Rossi etal. (2016)
Rhynchophoromyces sperchopsis Hydrophilidae USA Thaxter (1931)
Rhynchophoromyces unguicola Hydrophilidae China, Indonesia Shen and Ye (2006)
Synaptomyces argentinus Hydrophilidae Argentina Thaxter (1931)
Thaumasiomyces bipedalis Hydrophilidae Indonesia Thaxter (1931)
Thaumasiomyces scabellularius Hydrophilidae Cameroon Thaxter (1931)
Thaumasiomyces scaurus Hydrophilidae Cameroon Thaxter (1931)
Thripomyces italicus Hydraenidae Italy, Spain Santamaria (1993)
Thripomyces tessinensis Hydraenidae Poland Majewski (1994)
Laboulbeniaceae
Blasticomyces fastigiatus Hydrophilidae China, Indonesia Shen and Ye (2006)
Cantharomyces aquaticus Hydrochidae USA Thaxter (1931)
Cantharomyces bruchii Dryopidae Argentina Spegazzini (1917)
Cantharomyces denigratus Dryopidae Europe Thaxter (1931)
Cantharomyces flagellatus Dryopidae Trinidad Thaxter (1931)
Cantharomyces italicus Dryopidae Europe De Kesel and Haelewaters
(2014)
Cantharomyces permasculus Dryopidae Argentina Thaxter (1931)
Cantharomyces platensis Dryopidae Argentina Thaxter (1931)
Capillistichus tenellus Hydrophilidae Spain Santamaria (2004)
Chaetarthriomyces coelostomalis Hydrophilidae Cameroon, China, Malaysia Shen and Ye (2006)
Chaetarthriomyces crassiappendi-
catus Hydrophilidae Europe Haelewaters etal. (2014)
Chaetarthriomyces flexatus Hydrophilidae Indonesia Thaxter (1931)
Chitonomyces aculeifer Dytiscidae, Haliplidae Argentina, China, Europe De Kesel and Haelewaters
(2012)
Chitonomyces acuminatus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces aethiopicus Gyrinidae Africa Thaxter (1926)

Fungal Diversity
1 3
Table 5 (continued)
Taxon Substrate (Family of host-insects) Distribution Reference
Chitonomyces affinis Dytiscidae USA Goldmann and Weir (2012)
Chitonomyces africanus Dytiscidae Cameroon Thaxter (1926)
Chitonomyces amoyensis Dytiscidae China Thaxter (1926)
Chitonomyces appendiculatus Dytiscidae Central and South America, USA Goldmann and Weir (2012)
Chitonomyces arcuatus Dytiscidae Cameroon Thaxter (1926)
Chitonomyces argentinensis Noteridae Argentina Spegazzini (1917)
Chitonomyces aripensis Dytiscidae Trinidad Thaxter 1924)
Chitonomyces atricornis Gyrinidae Cameroon Thaxter (1926)
Chitonomyces aurantiacus Dytiscidae Brazil, USA Rossi and Bergonzo (2008)
Chitonomyces auriculatus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces australasiae Dytiscidae China, Fiji Shen and Ye (2006)
Chitonomyces bakeri Dytiscidae Asia, Greece, Spain Shen and Ye (2006)
Chitonomyces benjaminii Dytiscidae Sierra Leone Rossi (1994)
Chitonomyces bicolor Dytiscidae Grenada, Trinidad Thaxter (1924)
Chitonomyces bidessarius Dytiscidae Asia, USA Majewski (2008)
Chitonomyces bilabiatus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces borealis Dytiscidae USA Thaxter (1924)
Chitonomyces bullardii Haliplidae Spain, USA Santamaria (2003)
Chitonomyces cerviculatus Dytiscidae Grenada, Trinidad Thaxter (1924)
Chitonomyces chinensis Dytiscidae Asia, Sierra Leone Rossi and Leonardi (2018)
Chitonomyces conditus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces copelati Dytiscidae Cameroon Thaxter (1926)
Chitonomyces corniculatus Dytiscidae China Shen and Ye (2006)
Chitonomyces cornucopioides Gyrinidae Cameroon Thaxter (1926)
Chitonomyces dextricola Gyrinidae Cameroon Thaxter (1926)
Chitonomyces elegans Dytiscidae Cameroon, Indonesia Sugiyama and Nagasawa
(1986)
Chitonomyces elongatus Dytiscidae Italy, Spain Santamaria (2003)
Chitonomyces ensifer Dytiscidae Italy, Poland, Spain Majewski (2008)
Chitonomyces excavatus Dytiscidae China Shen and Ye (2006)
Chitonomyces floridanus Haliplidae USA Thaxter (1924)
Chitonomyces furcatus Dytiscidae China Shen and Ye (2006)
Chitonomyces gracilipes Dytiscidae China Shen and Ye (2006)
Chitonomyces grenadae Dytiscidae Grenada Thaxter (1924)
Chitonomyces helicoferus Dytiscidae Trinidad Thaxter (1924)
Chitonomyces hyalinus Dytiscidae Central and South America, USA Goldmann and Weir (2012)
Chitonomyces hydrocanthi Dytiscidae Sierra Leone Rossi (1990)
Chitonomyces hydropori Dytiscidae Europe, Philippines, USA Majewski (2008)
Chitonomyces incisus Dytiscidae Fiji Thaxter (1924)
Chitonomyces indentatus Dytiscidae Philippines, China Shen and Ye (2006)
Chitonomyces inflatus Dytiscidae Philippines Thaxter (1926)
Chitonomyces intermedius Dytiscidae Asia Shen and Ye (2006)
Chitonomyces interpositus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces introversus Dytiscidae Grenada, USA Thaxter (1924)
Chitonomyces iriomotensis Dytiscidae Asia, Spain Rossi etal. (2016)
Chitonomyces italicus Dytiscidae Europe De Kesel and Haelewaters
(2012)
Chitonomyces japanensis Dytiscidae China, Japan, Thailand Sugiyama and Phanichapol
(1984)

Fungal Diversity
1 3
Table 5 (continued)
Taxon Substrate (Family of host-insects) Distribution Reference
Chitonomyces javanicus Dytiscidae Asia Sugiyama and Phanichapol
(1984)
Chitonomyces laboulbenioides Haliplidae China Shen and Ye (2006)
Chitonomyces latens Gyrinidae Cameroon Thaxter (1926)
Chitonomyces ligulatus Haliplidae China Thaxter (1926)
Chitonomyces longirostratus Dytiscidae Trinidad, Japan Thaxter (1924)
Chitonomyces manubriolatus Dytiscidae Grenada, Thailand, Trinidad Sugiyama and Phanichapol
(1984)
Chitonomyces melanurus Dytiscidae Africa, Asia, Europe Try etal. (2017)
Chitonomyces obtusus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces occultus Haliplidae USA Thaxter (1908)
Chitonomyces oedipus Dytiscidae Grenada, Trinidad Thaxter (1924)
Chitonomyces ordinatus Dytiscidae Asia Try etal. (2017)
Chitonomyces orectogyri Gyrinidae Africa Thaxter (1908)
Chitonomyces orientalis Dytiscidae China, Japan, South Korea Shen and Ye (2006)
Chitonomyces paradoxus Dytiscidae Asia, Central America, Europe,
Sierra Leone, USA Goldmann and Weir (2012)
Chitonomyces pilicola Gyrinidae Cameroon Thaxter (1926)
Chitonomyces prolongatus Dytiscidae Brazil, USA Rossi and Bergonzo (2008)
Chitonomyces psittacopsis Dytiscidae Asia, North and Central America,
Sierra Leone Rossi and Leonardi (2018)
Chitonomyces pullus Haliplidae China Thaxter (1926)
Chitonomyces recurvatus Dytiscidae China Thaxter (1926)
Chitonomyces reynoldsii Dytiscidae Venezuela Thaxter (1926)
Chitonomyces rhynchostoma Dytiscidae Trinidad, USA Goldmann and Weir (2012)
Chitonomyces rugosus Dytiscidae Fiji, Thailand Sugiyama and Phanichapol
(1984)
Chitonomyces seticola Dytiscidae Grenada Thaxter (1924)
Chitonomyces sigmoideus Gyrinidae Cameroon Thaxter (1926)
Chitonomyces simplex Dytiscidae Brazil, Central America, Sierra
Leone, USA Goldmann etal. (2013)
Chitonomyces sinistricola Gyrinidae Cameroon Thaxter (1926)
Chitonomyces spinosus Dytiscidae Asia, Spain Shen and Ye (2006)
Chitonomyces striatus Dytiscidae Grenada Thaxter (1924)
Chitonomyces strictus Dytiscidae China Thaxter (1926)
Chitonomyces subterminalis Gyrinidae Cameroon Thaxter (1926)
Chitonomyces tarsicola Haliplidae China Thaxter (1926)
Chitonomyces thaxteri Dytiscidae Thailand, Trinidad Sugiyama and Phanichapol
(1984)
Chitonomyces tricornis Dytiscidae Cameroon Thaxter (1926)
Chitonomyces tylopus Dytiscidae Spain Santamaria (2001)
Chitonomyces uncinulatus Dytiscidae Trinidad Thaxter (1924)
Chitonomyces venezuelae Dytiscidae Venezuela Thaxter (1926)
Chitonomyces yunnanensis Dytiscidae China Shen and Ye (2006)
Chitonomyces zonatus Dytiscidae Asia Shen and Ye (2006)
Coreomyces acuminatus Corixidae Chile Thaxter (1918)
Coreomyces arcuatus Corixidae Europe, Jamaica Majewski (1994)
Coreomyces argentinensis Corixidae Argentina Spegazzini (1917)
Coreomyces australis Corixidae Argentina Spegazzini (1917)
Coreomyces bicolor Corixidae Jamaica Thaxter (1931)
Coreomyces chinensis Corixidae China Thaxter (1931)

Fungal Diversity
1 3
Table 5 (continued)
Taxon Substrate (Family of host-insects) Distribution Reference
Coreomyces confusus Corixidae Sweden, Denmark Sundberg etal. (2021)
Coreomyces corixae Corixidae Asia, Europe, North and South
America, Sweden, Turkey Sundberg etal. (2021)
Coreomyces curvatus Corixidae China, Italy, Jamaica, USA Shen and Ye (2006)
Coreomyces dextrorsus Corixidae Denmark, Sweden Sundberg etal. (2021)
Coreomyces elongatus Corixidae Europe Majewski (1994)
Coreomyces fastigiatus Corixidae Chile Thaxter (1931)
Coreomyces javanicus Corixidae Indonesia Lee and Na (2009)
Coreomyces macropus Corixidae Europe, Jamaica, Sweden Sundberg etal. (2021)
Coreomyces micronectae Corixidae Asia Shen and Ye (2006)
Coreomyces minor Corixidae China, Philippines Shen and Ye (2006)
Coreomyces orientalis Corixidae China, Indonesia Shen and Ye (2006)
Coreomyces ramosus Corixidae USA Thaxter (1931)
Coreomyces recurvatus Corixidae Asia Lee etal. (2008)
Coreomyces sigmoideus Corixidae Jamaica Thaxter (1931)
Coreomyces sp. Corixidae South Africa Goldmann and Weir (2018)
Coreomyces subdivisus Corixidae Chile Thaxter (1931)
Coreomyces thaxteri Corixidae Ethiopia Thaxter (1931)
Hydraeomyces halipli Haliplidae Asia, Europe, North and South
America, Rossi and Christian (2020)
Hydrophilomyces aduncus Hydraenidae Spain Santamaria (2006)
Hydrophilomyces arcuatus Hydraenidae Finland Huldén (1983)
Hydrophilomyces atroseptatus Hydraenidae Poland, Spain Santamaria (2003)
Hydrophilomyces coneglianensis Hydrophilidae Europe Castaldo etal. (2004)
Hydrophilomyces deflexus Hydraenidae Spain Santamaria etal. (2020)
Hydrophilomyces digitatus Hydraenidae Europe Santamaria (2003)
Hydrophilomyces gracilis Hydrophilidae Belgium, Greece, Poland Majewski (1994)
Hydrophilomyces hamatus Hydrophilidae Europe, Sierra Leone Majewski (1994)
Hydrophilomyces hydraenae Hydraenidae USA Boonmee etal. (2021)
Hydrophilomyces limnebii Hydraenidae Europe Majewski (1994)
Hydrophilomyces lumbricoides Hydrophilidae Argentina Tavares (1985)
Hydrophilomyces major Hydrophilidae Argentina Thaxter (1931)
Hydrophilomyces pusillus Hydrophilidae Poland Majewski (2008)
Hydrophilomyces reflexus Hydrophilidae Argentina, USA Thaxter (1908)
Hydrophilomyces rhynchophorus Hydrophilidae Central and South America, USA Goldmann and Weir (2018)
Hydrophilomyces rhytidopus Hydrophilidae Argentina, Trinidad Thaxter (1931)
Hydrophilomyces riberae Hydraenidae Spain Santamaria etal. (2020)
Laboulbenia afrogyri Gyrinidae Africa Rossi (1986)
Laboulbenia anomala Gyrinidae Africa Thaxter (1908)
Laboulbenia aquatica Gyrinidae Bolivia, Venezuela Weir and Rossi (2001)
Laboulbenia bicornis Gyrinidae Africa, Asia Balazuc (1982)
Laboulbenia birmanica Gyrinidae Myanmar Spegazzini (1915)
Laboulbenia borealis Gyrinidae Asia, North and Central America,
Madagascar Shen and Ye (2006)
Laboulbenia coarctata Gyrinidae India Thaxter (1908)
Laboulbenia constricta Gyrinidae Africa Rossi (1986)
Laboulbenia crassipes Gyrinidae Myanmar Spegazzini (1915)
Laboulbenia cubensis Gyrinidae Cuba Thaxter (1908)
Laboulbenia dactylophora Gyrinidae Africa Rossi (1986)

Fungal Diversity
1 3
is influenced by the duration of submergence, dynamics
of stream currents, type and stability of the substrate, light
availability, chemical and physical parameters of the water
(Matura and Krzewicka 2015). The type of substrate, such as
bedrock and size of the rocks, is the most important factor in
determining their distribution. Many aquatic lichens became
endangered or rare in some regions of Europe due to high
demands in the water clarity and the sensitivity of lichens
to pollution (Krzewicka etal. 2017). Furthermore, the water
current and scouring effect of sediments drifting over the
thallus can cause physical stress to lichen (Thüs etal. 2015).
The studies on freshwater lichens started during early
lichenological history (Rehman 1879; Boberski 1886;
Olech 1972, 1973; Kulikova etal. 2008; Matura and
Table 5 (continued)
Taxon Substrate (Family of host-insects) Distribution Reference
Laboulbenia denticulata Gyrinidae Australia Thaxter (1908)
Laboulbenia desgodii Gyrinidae Africa Rossi (1986)
Laboulbenia dineutis Gyrinidae Asia, Africa, Papua New Guinea Balazuc (1982)
Laboulbenia drepanalis Gyrinidae Panama, “Amazon” Thaxter (1908)
Laboulbenia fallax Gyrinidae Brazil, Panama, Trinidad Haelewaters etal. (2017)
Laboulbenia fennica Gyrinidae Europe Rossi and Christian (2020)
Laboulbenia franciscoloi Gyrinidae Sierra Leone Rossi (1986)
Laboulbenia funeralis Gyrinidae North, Central, and South
America Haelewaters etal. (2017)
Laboulbenia guerinii Gyrinidae North, Central, and South
America Thaxter (1908)
Laboulbenia gyrinicola Gyrinidae Algeria, Europe, Morocco Rossi and Christian (2020)
Laboulbenia gyrinidarum Gyrinidae Mexico, North America Thaxter (1908)
Laboulbenia heterocheila Gyrinidae “Timor” Thaxter (1908)
Laboulbenia leathsi Gyrinidae Chile Spegazzini (1917)
Laboulbenia leptopus Gyrinidae Argentina Spegazzini (1917)
Laboulbenia oberthuri Gyrinidae Madagascar Thaxter (1896)
Laboulbenia orechtochilii Gyrinidae Asia Thaxter (1908)
Laboulbenia orechtochilicola Gyrinidae Myanmar Spegazzini (1915)
Laboulbenia pachystoma Gyrinidae Argentina, Brazil Rossi and Bergonzo (2008)
Laboulbenia rotundata Gyrinidae Indonesia Thaxter (1908)
Laboulbenia satanas Gyrinidae Philippines Balazuc (1973)
Laboulbenia songhaiensis Gyrinidae Sierra Leone Rossi (1986)
Laboulbenia strangulata Gyrinidae Asia Try etal. (2017)
Laboulbenia valida Gyrinidae Indonesia Spegazzini (1915)
Limnaiomyces amphiopsis Hydrophilidae Cameroon Thaxter (1926)
Limnaiomyces hydrocharis Hydrophilidae USA Thaxter (1908)
Limnaiomyces tropisterni Hydrophilidae Mexico Thaxter (1908)
Rickia admirabilis Hydrophilidae Cameroon, Sierra Leone Rossi (1982)
Rickia coelostomalis Hydrophilidae Indonesia, Japan, China Shen and Ye (2006)
Scepastocarpus peritheciiformis Hydrochidae Spain Santamaria (2004)
Zodiomyces modae Hydrophilidae Japan Majewski and Sugiyama
(1989)
Zodiomyces rhizophorus Hydrophilidae Trinidad Rossi etal. (2016)
Zodiomyces subseriatus Hydrophilidae Asia, Poland Santamaria (2004)
Zodiomyces vermiformis Hydrophilidae Ecuador Das etal. (2018)
Zodiomyces vorticellarius Hydrophilidae Africa, Asia, Central and South
America, Europe, USA Goldmann and Weir (2018)
Pyxidiophorales
Pyxidiophoraceae
Pyxidiophora spinulorostrata Submerged decaying Quercus (river) UK Webster and Hawksworth
(1986)

Fungal Diversity
1 3
Fig. 4 Freshwater Laboulbeniales. a Whirligig beetle (Gyrinus caspius) bearing several thalli of Laboulbenia gyrinicola (arrow). b Laboulbe-
nia gyrinicola. c Ceratomyces cladophorus. d Chitonomyces paradoxus. Scale bars: b–d = 50µm
Krzewicka 2015; Krzewicka etal. 2017). Studies of fresh-
water lichens in British Isles has a huge contribution to
the taxonomic knowledge of this group (Gilbert 1996).
Thüs etal. (2014) estimated a total number of freshwater
lichens to be 270 but this is an underestimate, and the
number could be higher (Sonina and Androsova 2021).
The taxonomic studies of freshwater lichens are in the
urgent need of revision due to lack of molecular support
and morphological data (Nascimbene etal. 2007). Most
of the studies on freshwater lichens were solely based
on floristics surveys (Pentecost 1977; Keller and Schei-
degger 1994; Gilbert 1996; Gilbert and Giavarini 1997;
Molitor and Diederich 1997). Ecological studies have
been done also in aquatic lichens: effect of substrates in
aquatic lichens (Santesson 1939); influence of mineral
composition of water to the occurrence of freshwater
lichens and their thalli development (Pentecost 1977);
pollution indicator of water level fluctuations, water acid-
ity, water quality (Gilbert and Giavarini 1997; Thüs etal.
2014; Matura etal. 2017). Thüs etal. (2014) revised the
freshwater lichens and lichen-like organisms. The pre-
sent review provides the updated checklist of lichens and
lichenicolous fungi in freshwater habitats (Table10).
Lichens and lichenicolous taxa under Arthoniomycetes,
Dothideomycetes, Eurotiomycetes, and Sordariomycetes
included in this section are already listed for each taxo-
nomic class above.
Ascomycota genera incertae sedis
The list of freshwater fungal taxa in the Ascomycota with
uncertain taxonomic placements are provided in Table11. A
total of 252 species under 141 genera are recorded.
Classication offreshwater Basidiomycota
When compared to Ascomycota, few Basidiomycota have
been recorded from freshwater habitat. Jones and Choeyklin
(2008) and Jones etal. (2014a, b) listed freshwater Basidi-
omycota with 29 and 115 species, respectively. Asexual
morphs, with elaborate conidial forms, are mainly reported
and they are taxonomically diverse including both yeasts
and filamentous fungi. A special section on freshwater
yeast discusses both ascomycetous and basidiomycetous
yeasts. Freshwater Basidiomycota have been isolated from
variety of substrates: submerged decaying wood (Bandoni
and Marvanová 1989; Hyde and Goh 1998a; Mongkol etal.
2009; Yamaguchi etal. 2009; Ordynets 2009; Frank etal.
2010; Kohout etal. 2012; Chuaseeharonnachai etal. 2013),
leaf litter (Cunningham 1888; Engler etal. 1900; Fragoso
1917; Lavrov 1937; Thirumalachar and Mundkur 1951;
Vánky 1999; Verma and Jha 1999; Piatek etal. 2008; Lotz-
Winter etal. 2011), foam (Marvanová and Stalpers 1987;
Marvanová and Bärlocher 1988, 2000; Kirschner 2004), and

Fungal Diversity
1 3
Table 6 Classification of freshwater Leotiomycetes with information on the type of habitats, substrates, and geographical distribution of fungal
taxa
Taxon Substrate Distribution References
ASCOMYCOTA
LEOTIOMYCETES
Leotiomycetidae
Leotiomycetes families incertae sedis
Calloriaceae
Cistella graminicola Carex rostrata, Equisetum fluviatile Austria Magnes and Hafellner (1991)
Dactylaria africana Submerged decaying wood (river) Philippines Cai etal. (2003a)
Dactylaria appendiculata Submerged decaying wood (river) Brazil Cazau etal. (1990)
Dactylaria aquatica Water India Nair and Bhat (2016)
Dactylaria aspirensis Grass China Wong and Hyde (2001)
Dactylaria belliana Submerged leaves Brazil da Silva and Gusmão (2015)
Dactylaria biguttulata Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Dactylaria botulispora Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Dactylaria candidula Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Dactylaria fecundissima Submerged decaying wood China Tsui (1999)
Dactylaria flammulicornuta Submerged decaying Eleiodoxa conferta (peat
swamp) Thailand Pinnoi etal. (2006)
Dactylaria fusifera Submerged decaying leaves of Calophyllum brasil-
iense (stream) Brazil Fiuza etal. (2019)
Dactylaria fusiformis Submerged decaying balsa wood (river) and leaves
(stream) Brazil, USA Shearer and Crane (1971), Bar-
bosa etal. (2013b)
Dactylaria hemibeltranioidea Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Dactylaria hoogii Submerged decaying wood (lake) China Cai etal. (2002b), Luo etal.
(2004)
Dactylaria irregularis Submerged decaying wood (stream) China Ho (1998)
Dactylaria longidentata Grass, submerged bamboo (lake), wood (river) China, Philippines Luo etal. (2004)
Dactylaria lunata Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Dactylaria malaysianum Submerged wood China Tsui etal. (2000)
Dactylaria mucronulata Submerged wood China Tsui (1999)
Dactylaria naviculiformis Submerged decaying leaves of Calophyllum brasil-
iense (stream) Brazil Fiuza etal. (2019)
Dactylaria obscuriseptata Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Dactylaria obtriangularia Water-filled treeholes Hungary Gönczöl and Révay (2003)
Dactylaria plovercovensis Submerged decaying wood (lake, reservoir) China Goh and Hyde (1999)
Dactylaria saccardoana Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Dactylaria splendida Grass, submerged decaying wood (lake) China Luo etal. (2004)
Dactylaria triseptata Grass, submerged decaying wood (lake) Brunei, China Ho etal. (2001), Luo etal. (2004)
Dactylaria uliginicola Submerged petiole of Eleiodoxa conferta (peat
swamp) Thailand (Pinnoi etal. (2006), Pinruan etal.
(2007)
Diplonaevia emergens Juncus filiformis,
Eriophorum angustifolium Austria Magnes and Hafellner (1991)
Diplonaevia seriata Carex rostrata Austria Magnes and Hafellner (1991)
Mycoarthris corallina Foam (stream), freshwater UK, Korea Baschien etal. (2013), Lim etal.
(2021)
Rodwayella citrinula Phragmites australis stems and leaves, wood UK, Soviet Union Dudka (1963), Apinis etal.
(1972b), Taligoola etal. (1972),
Han etal. (2014)
Roseodiscus equisetinus Equisetum fluviatile Austria Magnes and Hafellner (1991)
Stamnaria persoonii Equisetum fluviatile Austria Magnes and Hafellner (1991),
Haelewaters etal. (2018)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Cenangiaceae
Moellerodiscus advenulus Submerged decaying Picea mariana needles (lake) USA Fallah and Shearer (2001)
Hamatocanthoscyphaceae
Chalara aotearoae Submerged wood China Tsui (1999)
Chalara siamense Submerged decaying Eleiodoxa conferta (peat
swamp) Thailand Pinnoi etal. (2006)
Chalara ungeri Grass China Wong and Hyde (2001)
Pezizellaceae
Calycina citrina Submerged decaying wood (swamp) USA Shearer and Crane (1986), Haele-
waters etal. (2018)
Calycina parilis (Chalara state) Submerged decaying wood Scotland Friggens etal. (2017)
Chalara alabamensis Submerged decaying petioles (stream) Brazil Barbosa etal. (2013b)
Rutstroemiaceae
Lambertella tubulosa Submerged decaying twigs of Acer pseudoplatanus,
Fagus sp. (stream, pond), water-filled treeholes Hungary, Ireland,
UK Abdullah and Webster (1980),
Abdullah and Webster (1981),
Gönczöl and Révay (2003), Vu
etal. (2019)
Sclerotiniaceae
Monilia aquatilis Water Germany Petrak (1930)
Leotiomycetes genera incertae sedis
Aquadiscula appendiculata Acer rubrum leaves USA Shearer and Crane (1985), Shearer
and Crane (1986)
Coleosperma lacustre Submerged decaying stalks of Schoenoplectus
lacustris (lake) UK Ingold (1954)
Coronellaria caricinella Carex nigra,
Carex echinata Austria Magnes and Hafellner (1991)
Flagellospora curvula Foam (stream, river, rivulet), leaves, submerged
wood (Alnus glutinosa, Corylus avellana, Fraxinus
excelsior, Fagus sylvatica, Tilia cordata), sub-
merged leaves of Alnus glutinosa, Coffea arabica
and Hevea brasiliensis (stream, river), water (river,
stream), scum (stream), sediment (dam, stream),
stemflow and throughfall of tree canopies
Austria, Brazil.
Canada, China,
France, Ghana,
India, Ire-
land, Jamaica,
Nigeria, Poland,
Spain, South
Africa, Sweden,
Switzerland,
Uganda, UK,
USA, Venezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Fenton (1950),
Ranzoni (1953), Nilsson (1958),
Ingold (1958b), Dixon (1959),
Nilsson (1960), Suzuki and
Nimura (1960), Hudson and
Ingold (1960)ngold (1960,
Ingold (1960), Greathead
(1961), Alasoadura (1968a),
Iqbal and Webster (1973),
Bärlocher (1987), Sridhar and
Kaveriappa (1988), Yu etal.
(1988), Gessner etal. (1993),
Voglmayr (1996), Harrington
(1997), Chan etal. (2000),
Marvanová (2000), Gulis and
Suberkropp (2003), Smits etal.
(2007), Schoenlein-Crusius etal.
(2009), Sridhar and Karamchand
(2009), Sudheep and Sridhar
(2012), Menéndez etal. (2012),
Baschien etal. (2013), Cornut
etal. (2014), Pietryczuk etal.
(2018)
Flagellospora fusarioides Submerged leaf of Crataegus
monogyna, foam (stream) Austria, UK Marvanová (2000), Baschien etal.
(2013)
Flagellospora leucorhynchos Foam (stream) Czech Republic Baschien etal. (2013)
Flagellospora minuta Submerged decaying leaves Pakistan Iqbal and Bhatty (1980)
Flagellospora saccata Foam (stream) Canada, India Rajashekhar and Kaveriappa
(2003), Baschien etal. (2013)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Flagellospora stricta Submerged decaying leaves, water from melting ice
(river) Poland, Sweden Nilsson (1962a), Czeczuga and
Orłowska (1999)
Hysteronaevia advena Eriophorum angustifolium Austria Magnes and Hafellner (1991)
Hysteronaevia olivacea Eriphorum vaginatum, Carex rostrata, C. pauper-
cula, C. Nigra, Juncus filiforme Austria Magnes and Hafellner (1991)
Hysteropezizella diminuens Carex rostrata, C. paupercula Austria Magnes and Hafellner (1991)
Hysteropezizella fuscella Carex nigra, C. rostrata Austria Magnes and Hafellner (1991)
Hysteropezizella phragmitina Carex rostrata Austria Magnes and Hafellner (1991)
Helotiales
Discinellaceae
Cladochasiella divergens Culture contaminant Czech Republic Baschien etal. (2013)
Fontanospora alternibrachiata Submerged decaying leaves (river), water (stream) France, USA Dyko (1978), Gulis and Suberk-
ropp (2003), Cornut etal. (2014)
Fontanospora eccentrica Foam (river, stream), submerged decaying leaves of
Quercus (spring), water (stream) Canada, Cuba,
Ireland, Japan,
Portugal, USA
Petersen (1962), Marvanová and
Marvan (1969), Dyko (1978),
Harrington (1997), Gulis and
Suberkropp (2003), Hosoya and
Tanaka (2007), Baschien etal.
(2013), Duarte etal. (2015),
Raposeiro etal. (2018)
Fontanospora fusiramosa Foam (stream) Czech Republic Baschien etal. (2013)
Gyoerffyella craginiformis Liriodendron tulipifera, decaying leaves Netherland, USA Petersen (1962), Baschien etal.
(2013)
Gyoerffyella entomobryoides Rosa sp. Netherland Baschien etal. (2013)
Gyoerffyella gemellipara Submerged decaying leaves (river), foam (stream,
rivulet) Austria, Czech
Republic Marvanová (1975), Voglmayr
(1996), Marvanová (2000),
Baschien etal. (2013)
Gyoerffyellamyrmecophagi-
formis River, foam (stream), melted ice (river) Austria, Poland Czeczuga and Orłowska (1999),
Marvanová (2000)
Gyoerffyella rotula Foam (stream, river, rivulet) Austria, Czech
Republic,
Ireland, New
Zealand, UK
Voglmayr (1996), Harrington
(1997), Cooper (2005), Baschien
etal. (2013)
Gyoerffyella speciosa Foam (stream) Canada, Japan Miura (1971), Ingold (1974),
Bärlocher (1987)
Gyoerffyella tatrica Foam (tidal ditch) UK Marvanová etal. (1967)
Gyoerffyella tricapillata Decaying leaf Punus and Rosa sp. in a pond, rainwa-
ter from building roofs Poland, UK Ingold (1964), Czeczuga and
Orłowska (1997), Baschien etal.
(2013)
Lemonniera alabamensis Submerged wood of Betula nigra (stream), sub-
merged decaying Alnus glutinosa leaves (stream),
decaying submerged leaf litter of Polygonum
nepalensis and Nerium sp., foam (stream)
Brazil, China,
France, India,
Portugal, Spain,
USA
Sinclair and Morgan-Jones
(1979a), Gessner etal. (1993),
Ho (1998), Menéndez etal.
(2012), Fiuza and Gusmão
(2013a), Sati (2014), Duarte
etal. (2015)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Lemonniera aquatica Water (pond, river, stream), scum (stream), spring
on flower petals of Malus sylvestris, Prunus spp,
Pyrus communis, foam (stream, rivelet), sub-
merged wood (Alnus glutinosa, Acer pseudoplata-
nus, Corylus avellana, Fraxinus excelsior, Fagus
sylvatica, Tilia cordata, Ulmus sp.), submerged
decaying leaves of Salix spp., Populus spp., Betula
glandulosa, and Alnus tenuifolia, (stream), foam
(lake, spring, stream, river), stemflow and through-
fall of tree canopies
Australia, Austria,
Brazil, Canada,
China, Czech
Republic,
Estonia, France,
Germany, India,
Ireland, Japan,
Norway, Poland,
Portugal, Roma-
nia, Russia,
South Africa,
Spain, Sweden,
Switzerland,
UK, Ukraine,
USA
Ingold (1942), Ingold (1943a),
Ranzoni (1953), Ingold (1960),
Ingold (1967), Crane (1968),
Descals etal. (1977), Bärlocher
(1987), Czeczuga etal. (1990),
Gessner etal. (1993), Sridhar
and Bärlocher (1993), Voglmayr
(1996), Harrington (1997),
Czeczuga and Orłowska (1999),
Marvanová (2000), Baschien
etal. (2008), Schoenlein-Crusius
etal. (2009), Sridhar and
Karamchand (2009), Menéndez
etal. (2012), Sati (2014), Cornut
etal. (2014), Duarte etal.
(2015), Raposeiro etal. (2018),
Pietryczuk etal. (2018), Vu etal.
(2019)
Lemonniera centrosphaera Water (pond, river), foam (stream, river, rivulet),
submerged leaf litter of Alnus glutinosa (stream),
scum (stream)
Austria, Canada,
China, Czech
Republic,
France, Ireland,
Japan, Poland,
UK, Ukraine
Miura (1971, 1972a), Descals
etal. (1977), Bärlocher (1987),
Gessner etal. (1993), Voglmayr
(1996), Harrington (1997),
Czeczuga and Orłowska (1999),
Pietryczuk etal. (2018), Baudy
etal. (2019)
Lemonniera cornuta Submerged decaying leaves and twigs of Alnus sp.,
Salix sp. and Quercus kellogi, submerged decaying
leaves and twigs of dicots (river, stream), foam
(stream, spring, river, lake), water (river. stream),
stemflow and throughfall of tree canopies
China, France,
India, Japan,
Spain, UK,
Ukraine, USA
Ranzoni (1953), Petersen (1963b),
Miura (1967), Crane (1968),
Iqbal (1972a), Iqbal and Webster
(1973), Dudka (1974), Des-
cals etal. (1977), Zhu and Yu
(1992), Campbell etal. (2006),
Sridhar and Karamchand (2009),
Menéndez etal. (2012), Cornut
etal. (2014)
Lemonniera filiformis Pond, foam (stream), water (stream), throughfall Austria, Canada,
China, India,
Germany,
Poland, Russia,
Ukraine, USA
Petersen (1963b), Arnold (1967),
Arnold (1968), Iqbal (1972a),
Dudka (1974), Bärlocher (1987),
Czeczuga and Orłowska (1999),
Marvanová (2000), Campbell
etal. (2006), Ghate and Sridhar
(2015)
Lemonniera pseudofloscula Submerged decaying leaves (river) water (stream),
foam (river), throughfall of palm species Brazil, India, USA Ingold (1958a), Descals etal.
(1977), Gulis and Suberkropp
(2003), Campbell etal. (2006),
Fiuza and Gusmão (2013a),
Ghate and Sridhar (2015)
Lemonniera terrestris Foam (pond, stream, river, rivulet), submerged
leaves (pond, stream, river), leaf baits (stream),
scum (stream), submerged decaying leaves of Salix
spp., Populus spp., Betula glandulosa, Alnus glu-
tinosa, and Alnus tenuifolia (stream), water (river,
stream), submerged decaying woods, stemflow and
throughfall of tree canopies
Australia, Aus-
tria, Canada,
China, Cuba,
Czech Repub-
lic, France,
Germany,
Hungary, India,
Ireland, Japan,
Poland, Russia,
Slovakia, South
Africa, Spain,
Sweden, UK,
Ukraine, USA
Ingold (1958a), Ingold (1958b),
Baxter (1960), Ingold (1960),
Marvanová and Marvan (1969),
Iqbal and Webster (1973),
Bärlocher (1987), Gessner
etal. (1993), Voglmayr (1996),
Czeczuga and Orłowska (1999),
Gulis and Suberkropp (2003),
Baschien etal. (2008), Sridhar
and Karamchand (2009), Baudy
etal. (2019)
Lemonniera yulongensis Submerged decaying leaves (river) China Yu etal. (2019)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Margaritispora aquatica Pond, foam (stream, rivulet), submerged decaying
leaves of Salix spp., Populus spp., Betula glandu-
losa, and Alnus tenuifolia, A. glutinosa (stream)
Austria, Brazil,
Canada, Korea,
Poland, UK,
USA
Ingold (1942), Ingold (1943a),
Baxter (1960), Bärlocher (1987),
Voglmayr (1996), Czeczuga and
Orłowska (1999), Marvanová
(2000), Schoenlein-Crusius
etal. (2009), Mun etal. (2016),
Baudy etal. (2019)
Margaritispora monticola Rainwater in forest, foam (stream), bedrock, decay-
ing angiosperm leaves (brook, stream, river),
submerged leaf litter of Alnus glutinosa (stream),
water (stream)
Australia, Aus-
tria, Canada,
Czech Republic,
France, New
Zealand, Por-
tugal, Slovakia,
UK, USA
Dyko (1978), Almer and Segedin
(1985), Gessner etal. (1993),
Marvanová (2000), Belliveau
and Bärlocher (2005), Duarte
etal. (2012), Cornut etal. (2014)
Pezoloma rhodocarpum Plant debris Malawi Fisher and Spooner (1987)
Tetrachaetumelegans Water (river, stream), foam (rivulet, river, stream,
spring, lake), submerged decaying leaves of Alnus
glutinosa, A. tenuifolia, Salix spp., Populus spp.,
Betula glandulosa (stream)
Australia,
Austria, Brazil,
Canada, France,
Germany, India,
Japan, Poland,
Portugal, Spain,
UK, USA
Ingold (1942), Ingold (1943a),
Ranzoni (1953), Baxter (1960),
Petersen (1963a), Crane (1968),
Iqbal and Webster (1973),
Marvanová and Descals (1985),
Bärlocher (1987), Gessner
etal. (1993), Voglmayr (1996),
Czeczuga and Orłowska (1999),
Marvanová (2000), Gulis and
Suberkropp (2003), Rajashek-
har and Kaveriappa (2003),
Belliveau and Bärlocher (2005),
Hosoya and Tanaka (2007),
Schoenlein-Crusius etal. (2009),
Menéndez etal. (2012), Cornut
etal. (2014), Funck etal. (2015),
Raposeiro etal. (2018), Vu etal.
(2019)
Gelatinodiscaceae
Dimorphospora foliicola Castanea leaf, stream, submerged decaying leaves
of Alnus glutinosa (stream, river), water-filled
treeholes
France, Germany,
Hungary, Japan,
Portugal, UK
Iqbal and Webster (1973), Gessner
etal. (1993), Gönczöl and Révay
(2003), Belliveau and Bärlocher
(2005), Baschien etal. (2013)
Fluminispora ovalis Submerged decaying leaves of Alnus glutinosa UK Ingold (1958a)
Helotiaceae
Articulospora atra Picea abies twiglet (stream), foam (river) Czech Republic,
UK Descals and Webster (1982),
Baschien etal. (2013)
Articulospora moniliformis Submerged Alnus and Salix New Zealand,
USA Ranzoni (1953), Petersen (1962),
Almer and Segedin (1985)
Articulospora proliferata Submerged leaves ofPhragmites communis (river) Poland, Spain Czeczuga and Orłowska (1999),
Baudy etal. (2019)
Articulospora tetracladia Foam, foam/water (stream) Canada, Korea,
France, Por-
tugal, USA,
Venezuela
Gulis and Suberkropp (2003),
Belliveau and Bärlocher (2005),
Smits etal. (2007), Mun etal.
(2016), Raposeiro etal. (2018)
Cudoniella indica Foam (river) South Africa Webster etal. (1995), Vu etal.
(2019)
Cudoniella clavus Ash, alder, oak twigs UK Willoughby and Archer (1973)
Dimorphospora foliicola Water (stream) USA Gulis and Suberkropp (2003)
Helicocentralis hyaline Submerged decaying wood (peat swamp) Thailand Sri-indrasutdhi etal. (2015)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Helicodendron triglitziense Submerged leaves of Salix (reservoir), water-filled
treeholes Germany, Hun-
gary, Ireland,
Netherlands,
USA
Linder (1929), van Beverwijk
(1953), Glen-Bott (1955),
Abdullah and Webster (1980),
Gönczöl and Révay (2003)
Helicodendron westerdijkiae Water-filled treeholes Hungary Gönczöl and Révay (2003)
Crocicreas megalosporum Carex rostrata Austria Magnes and Hafellner (1991)
Hymenoscyphus africanus Alder twigs UK Descals etal. (1984)
Hymenoscyphus imberbis Submerged decaying twig (stream) Slovakia Marvanová and Descals (1996)
Hymenoscyphus foliicola Acer pseudoplatanus twigs UK Abdullah etal. (1981)
Hymenoscyphus malawiensis Plant debris Malawi Fisher and Spooner (1987)
Hymenoscyphus paradoxus Willow twigs Switzerland Fisher and Webster (1983)
Hymenoscyphus phyllophilus Skeletal leaves Soviet Union Dudka (1963)
Hymenoscyphus scutula Submerged decaying stems of Scirpus atrovirens
(lake) USA Fallah and Shearer (2001)
Hymenoscyphus splendens Beech cupules UK Abdullah etal. (1981)
Hymenoscyphus tetracladius Beech and willow leaves (stream, lake), submerged
wood (Alnus glutinosa, Corylus avellana, Tilia
cordata, Ulmus sp.) (stream), foam (stream,
rivulet, river), submerged decaying leaves of Alnus
glutinosa, A. tenuifolia, Salix spp., Populus spp.,
Betula glandulosa (stream, river), water (river),
scum (stream)
Austria, Brazil,
Canada, France,
Ireland, India,
Japan, Nigeria,
Spain, Switzer-
land, UK, USA
Ingold (1942), Ingold (1943a),
Ingold (1949), Ranzoni (1953),
Hudson and Ingold (1960),
Baxter (1960), Ingold (1960),
Petersen (1962), Alasoadura
(1968a), Iqbal and Webster
(1973), Abdullah and Webster
(1980), Abdullah etal. (1981),
Bärlocher (1987), Gessner etal.
(1993), Sridhar and Bärlocher
(1993), Harrington (1997),
Marvanová (2000), Rajashek-
har and Kaveriappa (2003),
Menéndez etal. (2012), Fiuza
and Gusmão (2013a)
Hyaloscyphaceae
Arbusculina fragmentans Spume (river), flower petal by spring, pond, and
river Slovakia, Slove-
nia, Poland Marvanová (1988), Czeczuga
etal. (2007), Baschien etal.
(2013)
Arbusculina irregularis Roof top rainwater, submerged debris (creek), foam
(stream) India, Poland,
USA (Petersen (1963a), Sridhar and
Kaveriappa (1992), Czeczuga
and Orłowska (1997), Czeczuga
and Orłowska (1999)
Brocchiosphaera brocchiata Submerged decaying Licuala longicalycata (peat
swamp), submerged decaying wood (e.g., Diptero-
carpus alatus and Xylia dolabriformis test blocks)
(lake, stream, river, reservoir) and bamboo (river)
Brazil, Brunei,
China, Japan,
Malaysia, Philip-
pines, Thailand
(Tubaki (1975), Goh and Hyde
(1999), Ho etal. (2001), Cai
etal. (2003a), Luo etal. (2004),
Pinruan etal. (2007), Chua-
seeharonnachai etal. (2013),
Barbosa etal. (2013b)
Brocchiosphaera microspora Submerged decaying woods (stream), submerged
leaves of Calophyllum brasiliense (stream) Brazil, Thailand Chuaseeharonnachai etal. (2013)
Clathrosphaerinazalewskii On decaying leaves ofBetula in water Netherland van Beverwijk (1951b), Abdullah
and Webster (1980), Vu etal.
(2019)
Clavidisculum karstenii Submerged Umbelliferae stem (stream) UK Ekanayaka etal. (2019)
Clavidisculum graminicola Carex rostrata, Equisetum fluviatile Austria Magnes and Hafellner (1991)
Dematioscypha dematiicola Submerged decaying Eleiodoxa conferta (peat
swamp) Thailand Pinnoi etal. (2006)
Hyaloscypha albohyalina Twigs, rainwater from building roofs, Decaying
wood of Nothofagus fusca (stream, waterfall) Poland, New
Zealand, UK Abdullah and Webster (1983),
Czeczuga and Orłowska (1997),
Vu etal. (2019)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Hyaloscypha spinulosa Submerged decaying leaves of Alnus and Betula
(brook, ponds, rainwater-filled bombcrater, tree-
holes), decaying leaves of Nothofagus
Argentina, Hun-
gary, Ireland,
Japan, Nether-
lands, Switzer-
land, UK
van Beverwijk (1951a, b), Bot-
tomley (1954), Tubaki (1958),
Gamundi etal. (1977), Gönczöl
and Révay (2003)
Hyaloscypha zalewskii Catkins of Alnus, spine of Castanea sativa (stream),
decaying stems of Endymion non-scriptus (river),
Quercus cupule, Alnus glutinosa strobilus (river),
submerged roots, rainwater from building roofs
Ireland, Nether-
land, Poland,
UK
Descals and Webster (1976),
Czeczuga and Orłowska (1997)
Pseudaegerita caffra Dead twig (stream) Japan Matsushima (1996)
Pseudaegerita conifera Submerged needles ofPinus halepensis ( river) Spain Abdullah etal. (2005)
Pseudaegeritacorticalis River, submerged decaying wood (stream) Poland, Thailand Czeczuga and Orłowska (1999),
Chuaseeharonnachai etal.
(2013), Vu etal. (2019)
Pseudagaerita foliicola Oak leaf (lake) UK Field and Webster (1985)
Pseudaegerita ossiformis Decaying twig of Quercus (river) Italy Abdullah etal. (1997)
Pseudaegerita viridis River, water-filled treeholes, submerged decaying
wood (stream) Brazil, Hungary,
Poland Czeczuga and Orłowska (1999),
Gönczöl and Révay (2003),
Bruzone etal. (2015), Fiuza
etal. (2017b)
Pseudaegerita websteri Submerged decaying wood, decaying leaves of Calo-
phyllum brasiliense (stream) Brazil, Spain Abdullah etal. (2005), Fiuza etal.
(2019)
Hydrocinaceae
Clathrosporiumintricatum Submerged decaying angiosperm and unidentified
leaves (stream) Malaysia, Thai-
land Nawawi and Kuthubutheen
(1987b), Voglmayr (2004),
Chuaseeharonnachai etal.
(2013)
Clathrosporium retortum From leaf litter submerged in fresh water Brazil Moro etal. (2015a)
Hydrocina chaetocladia Submerged alder and oak twigs (stream) UK Shearer and Webster (1991),
Webster etal. (1991), Vu etal.
(2019)
Varicosporium aquaticum Water (river) Rusia Anonymus (1950)
Varicosporiumdelicatum Submerged of rotting leaves ofAcer pseudoplatanus,
foam (rivulet, river, stream), water (river) Austria, Canada,
Ireland, Poland,
Portugal
Bärlocher (1987), Voglmayr
(1996), Harrington (1997),
Czeczuga and Orłowska (1999),
Duarte etal. (2015), Pietryczuk
etal. (2018)
Varicosporiumelodeae Foam (rivulet, stream, spring, river, lake), sub-
merged decaying leaves of Alnus glutinosa
(stream), submerged decaying leaves of broad-
leaved trees, shrubs, and angiosperms, stemflow
and throughfall of tree canopies, water-filled
treeholes, water (river, stream)
Austria, Bra-
zil, Canada,
Czech Repub-
lic, France,
Hungary, India,
Ireland, Japan,
New Zealand,
Poland, Slova-
kia, Sweden,
UK, USA
Ingold (1942), Ingold (1943a),
Fenton (1950), Tubaki (1957),
Nilsson (1958), Willen (1958),
Petersen (1963b), Crane (1968),
Bärlocher (1987), Gessner etal.
(1993), Sridhar and Bärlocher
(1993), Voglmayr (1996), Har-
rington (1997), Czeczuga and
Orłowska (1999), Marvanová
(2000), Gönczöl and Révay
(2003), Baschien etal. (2008),
Schoenlein-Crusius etal. (2009),
Sridhar and Karamchand (2009),
Menéndez etal. (2012), Cornut
etal. (2014), Pietryczuk etal.
(2018), Vu etal. (2019)
Varicosporium giganteum Foam (stream, spring, river, lake), submerged decay-
ing leaves (stream) Canada, China,
Ireland, New
Zealand, Thai-
land, USA
Crane (1968), Bärlocher (1987),
Zhu and Yu (1992), Sridhar and
Bärlocher (1993), Harrington
(1997), Sakayaroj etal. (2005),
Campbell etal. (2009)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Varicosporium helicosporum Foam (river) Malaysia Nawawi (1974)
Varicosporium macrosporum Foam (river) Malaysia, Thai-
land Nawawi (1974), Sakayaroj etal.
(2005)
Varicosporium scoparium Foam (river) Spain Baschien etal. (2013)
Varicosporium tricladiiforme Submerged leaves of Salix atrocinerea, foam
(stream) Austria, Spain Roldan etal. (1992), Marvanová
(2000)
Varicosporium trimosum Foam (stream) Canada, Czech
Republic Marvanová and Bärlocher (2001),
Baschien etal. (2013)
Lachnaceae
Lachnum calycioides Eriophorum vaginatum
Juncus filiformis Austria Magnes and Hafellner (1991)
Lachnum controversum Stems of Phragmites australis UK, Germany Shearer (1993a)
Lachnum cf. virgineum wood USA Lamore and Goos (1978)
Lachnum imbecille Eriophorum angustifolium Austria Magnes and Hafellner (1991),
Kosonen etal. (2020)
Lachnum nudipes Stems of Phragmites australis UK Taligoola etal. (1972), Cantrell
and Hanlin (1997)
Loramycetaceae
Loramyces juncicola Dead stalks of Scirpus, Eleocharis, Equisetum,
Juncus UK, USA Ingold (1955), Digby and Goos
(1987), Vu etal. (2019)
Loramyces macrosporus Internodes of Equisetum fluviatile UK Ingold and Chapman (1952),
Baschien etal. (2013)
Obtectodiscus aquaticus Carex rostrata Austria, Switzer-
land (Muller etal. (1979), Magnes
and Hafellner (1991), Vu etal.
(2019)
Mitrulaceae
Mitrula alba Submerged leaves UK Cannon etal. (1987)
Mitrula paludosa Submerged leaves, wood UK Dennis (1978), Vu etal. (2019)
Mollisiaceae
Belonopsis guestphalicum Submerged decaying Phragmites stem, Schoenoplec-
tus lacustris stalks, Typha angustifolia leaf bases
(lake)
UK Ingold (1954)
Mollisiacasaresiae Submerged birch dowel UK Webster etal. (1993)
Mollisia cinerea Wood, Typha latifolia Soviet Union Dudka (1963), Tanney and Seifert
(2020)
Mollisia gigantea Picea twigs Finland Fisher and Webster (1983)
Mollisia hydrophila Phragmites Czechoslovakia,
Denmark, UK,
Finland, France,
Germany, Poland,
Scotland, Swe-
den,
Switzerland
Nannfeldt (1985), Tanney and
Seifert (2020)
Mollisia mediella Phragmites,
Schoenoplectus lacustris UK, Finland,
France, Ger-
many,
Ireland, Sweden,
Switzerland
Nannfeldt (1985)
Mollisia melaleuca Wood Soviet Union Dudka (1963)
Mollisia pilosa Carex, Schoenoplectus lacustris UK, Soviet Union Dudka (1963), Dennis (1978), Vu
etal. (2019)
Mollisia pulla Submerged decaying Carex, Eleocharis, Juncus,
Phragmites, Phalaris (lake) UK Dennis (1978), Vu etal. (2019)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Mollisia retincola Phragmites, Carex,Typha UK, Finland,
France, Ger-
many, Ireland,
Sweden,
Switzerland
Nannfeldt (1985)
Mollisia uda Wood UK Shearer (1993a)
Nimbomollisia eriophori Carex rostrata, Eriophorum
vaginatum, Juncus filiformis Austria Magnes and Hafellner (1991)
Nimbomollisia melatephroides Carex rostrata Austria Magnes and Hafellner (1991)
Niptera bispora Carex rostrata Austria Magnes and Hafellner (1991)
Niptera guestphalica Schoenoplectus lacustris, Typha leaf bases UK Ingold (1954)
Niptera lacustris Schoenoplectus lacustris, Scirpus, Phragmites UK, Scandinavia Shearer (1993a)
Niptera melanophae Schoenoplectus lacustris Soviet Union Dudka (1963)
Niptera tephromelas Typha Italy Nannfeldt (1985)
Phialocephala fluminis Submerged decaying balsa wood (river) USA Shearer etal. (1976)
Phialocephala xalapensis Submerged decaying wood (stream) Brunei, China Ho etal. (2001)
Scutomollisia morvernensis Carex rostrata, Equisetum
fluviatile, Eriophorum
angustifolium, Glyceria fluitans, Juncus filiformis
Austria Magnes and Hafellner (1991)
Scutomollisia punctum Carex rostrata Austria Magnes and Hafellner (1991)
Tapesia kneiffii Phragmites stalks UK Ingold (1954)
Trichobelonium kneiffii Phragmites, Carex, Typha Czech Republic,
Finland, France,
Germany,
Ireland, Sweden,
Switzerland, UK
Nannfeldt (1985)
Trimmatostroma betulinum Leaves China Au etal. (1992), Vu etal. (2019)
Variocladium giganteum Submerged decaying leaves of Crataegus monogyna
and Poaceae, foam (stream) Canada, UK Bärlocher (1987), Baschien etal.
(2013)
Variocladium rangiferinum Foam (river) UK Descals and Webster (1982)
Ploettnerulaceae
Belonium excelsior Submerged decaying Phragmites Finland UK Ingold (1954, 1955)
Cadophora novi-eboraci Freshwater Korea Lim etal. (2021)
Collembolispora aristata Foam (stream) Czech Republic Crous etal. (2012)
Collembolispora barbata Foam (stream) Portugal Marvanová etal. (2003)
Pyrenopeziza arundinacea Phragmites australis leaves UK Apinis etal. (1972b)
Pyrenopeziza huetteri Carex rostrata Austria Magnes and Hafellner (1991)
Pyrenopeziza perminuta Carex rostrata Austria Magnes and Hafellner (1991)
Volucrispora aurantiaca Foam (rivulet, stream) Austria, Canada,
UK Haskins (1958), Ingold (1959a),
Voglmayr (1996), Marvanová
(2000)
Volucrispora graminea Foam (river, rivulet, stream), water (river), stemflow
and throughfall of tree canopies and palm species Austria, Canada,
India, Ireland,
Poland, Portu-
gal, UK
Ingold etal. (1968b), Iqbal and
Webster (1973), Bärlocher
(1987), Voglmayr (1996),
Harrington (1997), Marvanová
(2000), Marvanová and
Bärlocher (2001), Sridhar and
Karamchand (2009), Baschien
etal. (2013), Ghate and Sridhar
(2015), Pietryczuk etal. (2018)
Tricladiaceae

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Geniculospora grandis Decomposing plant litter, foam (stream), water
(river), submerged decaying leaves (stream) Australia, Aus-
tria, Canada,
Poland, Por-
tugal, Spain,
Thailand
Czeczuga etal. (1990), Srid-
har and Bärlocher (1993),
Marvanová (2000), Marvanová
and Hywel-Jones (2000),
Marvanová etal. (2003), Bel-
liveau and Bärlocher (2005),
Duarte etal. (2012)
Geniculospora inflata Foam (river, stream), submerged decaying leaves
of Alnus glutinosa (stream), water (river, stream),
Submerged decaying leaves of Calophyllum brasil-
iense (stream)
Austria, Brazil,
France, Ireland,
Portugal, Spain,
UK, USA
Ingold (1943a), Anastasiou
(1964), Iqbal and Webster
(1973), Czeczuga etal. (1990),
Gessner etal. (1993), Harrington
(1997), Marvanová (2000), Gulis
and Suberkropp (2003), Menén-
dez etal. (2012), Raposeiro etal.
(2018), Fiuza etal. (2019)
Graddonia coracina Wood UK Dennis (1978), Friggens etal.
(2017)
Mycofalcella calcarata Submerged branch of Quercus, water (stream) France, Portugal,
UK, USA Gulis and Suberkropp (2003),
Baschien etal. (2013), Cornut
etal. (2014), Raposeiro etal.
(2018)
Mycofalcella iqbalii Submerged decaying culm of Saccharum munja
(canal) Pakistan Firdaus-E-Bareen and Braun
(2007)
Trichopezizella hystricula Carex rostrata Austria Magnes and Hafellner (1991)
Tricladium aciculum Foam (river, stream), submerged decaying leaves Malaysia, Thai-
land Nawawi (1985c), Sakayaroj etal.
(2005), Hosoya etal. (2006)
Tricladium alaskense Stream USA Baschien etal. (2013)
Tricladiumangulatum Pond water, foam (stream, spring, river, lake),
submerged decaying leaves of Alnus glutinosa, A.
tenuifolia, Salix spp., Populus spp., Betula glandu-
losa (stream), water (river), scum (stream)
Austria, Canada,
Czech Repub-
lic, Germany,
Poland, Swit-
zerland, UK,
USA, Venezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Ranzoni (1953),
Baxter (1960), Ingold (1960),
Petersen (1962), Crane (1968),
Bärlocher (1987), Czeczuga
etal. (1990), Czeczuga and
Orłowska (1999), Marvanová
(2000), Belliveau and Bärlocher
(2005), Baschien etal. (2006),
Smits etal. (2007), Baschien
etal. (2008), Duarte etal.
(2015), Pietryczuk etal. (2018),
Baudy etal. (2019)
Tricladium anomalum On submerged decaying leaves ofTypha latifolia,
river, foam and scum Poland, UK, USA Anastasiou (1964), Ranzoni
(1979), Czeczuga and Orłowska
(1999)
Tricladium attenuatum Foam (rivulet, stream) Austria, Canada,
Czech Republic,
Portugal
Bärlocher (1987), Voglmayr
(1996), Baschien etal. (2013),
Duarte etal. (2015)
Tricladium biappendiculatum Foam (rivulet, stream), submerged decaying leaves
(stream) Austria, Canada,
Czech Republic,
UK
Marvanová and Descals (1987),
Sridhar and Bärlocher (1993),
Voglmayr (1996), Marvanová
(2000), Campbell etal. (2009),
Baschien etal. (2013)
Tricladium brunneum Foam (river) India, Japan,
Malaysia Nawawi (1974), Sridhar and
Kaveriappa (1992), Hosoya and
Tanaka (2007)
Tricladium castaneicola Foam (stream, river), water-filled treeholes Czech Republic,
Hungary, Ireland Harrington (1997), Gönczöl and
Révay (2003), Baschien etal.
(2013)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Tricladium caudatum Foam (rivulet, stream) Austria, Canada,
Czech Republic Voglmayr (1996), Marvanová
(2000), Marvanová and Bär-
locher (2001), Baschien etal.
(2013)
Tricladium chaetocladium Water (stream, river), submerged decaying leaves
(e.g., Alnus glutinosa, Alnus sp. (stream, river),
foam (stream)
Australia,
Canada, France,
Malaysia, Portu-
gal, Spain, USA
Nawawi (1985b), Bärlocher
(1987), Gessner etal. (1993),
Gulis and Suberkropp (2003),
Belliveau and Bärlocher (2005),
Duarte etal. (2012), Menéndez
etal. (2012), Baschien etal.
(2013), Cornut etal. (2014),
Funck etal. (2015), Raposeiro
etal. (2018)
Tricladium curvisporum Foam (rivulet, river, stream) Austria, Brazil,
Ireland, UK Descals and Webster (1983),
Voglmayr (1996), Harrington
(1997), Marvanová (2000),
Baschien etal. (2013), Fiuza
etal. (2015a)
Tricladium fallax Foam (river, stream) Brazil, Czech
Republic Marvanová (1984), Fiuza and
Gusmão (2013a)
Tricladium fuscum Rotten leaves and foam (stream) Malaysia Nawawi (1985c)
Tricladium gracile Pond, submerged decaying leaves, water (river),
foam (rivulet, stream, spring, river, lake) Austria, Canada,
Ireland, Poland,
UK, USA
Ingold (1943a), Ranzoni (1953),
Petersen (1962), Crane (1968),
Iqbal and Webster (1973), Bär-
locher (1987), Voglmayr (1996),
Harrington (1997)
Tricladium indicum Foam, on submerged rotten leaves of Pinus rox-
burghii China, India,
Malaysia, USA Ho etal. (2001), Hyde (2007),
Baschien etal. (2013)
Tricladium intermedium On submerged of rotting twigs, submerged decaying
angiosperm leaves, foam (river, stream, spring,
lake)
USA Petersen (1962), Crane (1968)
Tricladium kelleri Stream USA Baschien etal. (2013)
Tricladium marylandicum Foam (pond, lake, stream, spring, river), submerged
decaying angiosperm leaves Poland, USA Crane (1968), Czeczuga and
Orłowska (1999)
Tricladium minutum Juncus culms, foam (stream) Canada, UK Bärlocher (1987), Baschien etal.
(2013)
Tricladium obesum Foam (stream) Czech Republic Marvanová (2004), Baschien etal.
(2013)
Tricladium patulum Foam (rivulet, river, stream), submerged leaf litter of
Alnus glutinosa (stream) Austria, Canada,
Czech Republic,
France, Ireland,
Poland
Bärlocher (1987), Gessner etal.
(1993), Voglmayr (1996),
Harrington (1997), Marvanová
(2000), Belliveau and Bärlocher
(2005), Baschien etal. (2013),
Pietryczuk etal. (2018)
Tricladium procerum Submerged dead stem of Juncus, rainwater from
building roofs Poland, Slovenia Marvanová (1988), Czeczuga and
Orłowska (1997), Baschien etal.
(2013)
Tricladiumrobustum In mountain stream, on rotten leaf ofFagus syl-
vatica, foam (stream) Austria, Czech
Republic, Spain Marvanová (1984), Descals
(1987), Marvanová (2000)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Tricladium splendens Foam (rivulet, stream, river, spring, lake), sub-
merged decaying leaves of Alnus glutinosa, A. ten-
uifolia, Salix spp., Populus spp., Betula glandulosa
(stream), foam (stream), stemflow and throughfall
of tree canopies, water (stream)
Austria, Brazil,
Canada, Czech
Republic,
France, India,
Ireland, Japan,
Poland, Slova-
kia, Switzerland,
UK, USA,
Venezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Ranzoni (1953),
Baxter (1960), Petersen
(1962), Crane (1968), Eaton
and Jones (1971b), Bärlocher
(1987), Gessner etal. (1993),
Voglmayr (1996), Harrington
(1997), Marvanová (2000),
Belliveau and Bärlocher (2005),
Baschien etal. (2006), Hosoya
and Tanaka (2007), Smits etal.
(2007), Baschien etal. (2008),
Schoenlein-Crusius etal. (2009),
Sridhar and Karamchand (2009),
Baschien etal. (2013), Cornut
etal. (2014), Pietryczuk etal.
(2018)
Tricladium terrestre Foam (stream) Ireland, Portugal Baschien etal. (2013)
Tricladium varicosporioides Twigs, submerged decaying wood (e.g., Dipterocar-
pus alatus and Xylia dolabriformis test blokcs) Japan, Thailand Tubaki (1966), Sivichai etal.
(2002), Johnston etal. (2019)
Tricladium varium Submerged decaying wood of Pinus sylvestris and
Fagus sylvatica (water-cooling towers) UK Jones and Stewart (1972)
Vibrisseaceae
Gorgoniceps boltonii Submerged decaying Equisetum fluviatile
Stems (lake) Austria, UK Ingold (1954), Magnes and Hafell-
ner (1991)
Leucovibrissea obconica Alder twigs Scandinavia Schumacher (1976)
Vibrissea decolorans Twigs Sweden, Norway Schumacher (1976)
Vibrissea dura Eucalyptus logs Australia Beaton and Weste (1976)
Vibrissea filisporia Submerged alder, willow and Corylus twigs (stream) UK, Hungary,
France, Norway,
Sweden
Graddon (1965), Schumacher
(1976), Révay and Gönczöl
(1990), Han etal. (2014),
Vibrissea flavovirens Submerged decaying wood and leaves (stream, river) UK Graddon (1965), Descals and Sut-
ton (1976), Zheng and Zhuang
(2017)
Vibrissea guernisacii Twigs, water (river, lake) Poland, UK, USA Ingold (1954), Lamore and Goos
(1978), Czeczuga etal. (1990)
Vibrissea leptospora Prunus, Rosa, Ribes, Salix Sweden, UK, Graddon (1965)
Vibrissea norvegica Phragmites australis Norway Sánchez (1969)
Vibrissea sporogyra Submerged decaying stalks of Equisetum, Carex,
Eleocharis (lake) UK Ingold (1954)
Vibrissea truncorum Maple, alder, birch
willow, juniper branches Scandinavia Kjuller (1960), Tanney and Seifert
(2020)
Vibrissea vibrisseoides Wood Scandinavia Kjuller (1960), Tanney and Seifert
(2020)
Helotiales genera incertae sedis
Aquapoterium pinicola Submerged Pinus needles (swamp) USA Raja etal. (2008b)
Chlorovibrissea bicolor Wood Australia Beaton and Weste (1977)
Chlorovibrissea melanochlora Eucalyptus logs Australia Beaton and Weste (1976)
Chlorovibrissea tasmanica Eucalyptus logs Australia Beaton and Weste (1976)
Filosporella annelidica Foam (river, stream), water, submerged decaying
leaves of Platanus occidentalis and Acer sacchari-
num (river)
Austria, Spain,
UK, USA Shearer and Crane (1976),
Marvanová (2000), Baschien
etal. (2013)
Filosporella aquatica Ssubmerged decaying leaves of Pandanus helicopus Malaysia Nawawi (1976a)
Filosporella exilis Equisetum fluviatile Belarus Baschien etal. (2013)
Filosporella fistucella Submerged roots of Alnus glutinosa UK Baschien etal. (2013)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Filosporella pinguis Foam (stream) Canada Marvanová and Bärlocher (1998c)
Filosporella versimorpha Submerged roots of Alnus glutinosa UK Baschien etal. (2013)
Tetracladium apiense Plant debris (stream), foam (stream) South Africa,
Spain Roldán etal. (1989), Baschien
etal. (2013)
Tetracladium breve Leaf Frangula cf. alnus, foam (stream), water (river) Austria, Brazil,
Poland, Spain Roldán etal. (1989), Marvanová
(2000), Baschien etal. (2013),
Pietryczuk etal. (2018)
Tetracladium furcatum Leaves, foam (river, stream), water (river, stream) Austria, China,
Poland, Portu-
gal, UK
Descals and Webster (1983), Zhu
and Yu (1992), Marvanová
(2000), Fiuza and Gusmão
(2013a), Baschien etal. (2013),
Raposeiro etal. (2018), Pietryc-
zuk etal. (2018)
Tetracladium globosum Freshwater Korea Lim etal. (2021)
Tetracladiummarchalianum Water (pond, river, stream), submerged decaying
wood (Alnus glutinosa, Acer pseudoplatanus,
Corylus avellana, Fraxinus excelsior, Fagus
sylvatica, Tilia cordata, Ulmus sp., submerged
decaying leaves of Salix spp., Populus spp., Betula
glandulosa, and Alnus tenuifolia, (stream), foam
(stream, river, rivulet), sediment (dam, stream)
Austria, Canada,
Cuba, Czech
Repub-
lic, France,
Germany,
India, Jamaica,
Nigeria, Poland,
Portugal, Spain,
Switzerland,
UK, USA, Ven-
ezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Ranzoni (1953),
Hudson and Ingold (1960),
Baxter (1960), Ingold (1960),
Marvanová and Marvan (1969),
Iqbal and Webster (1973),
Bärlocher (1987), Roldán etal.
(1989), Czeczuga etal. (1990),
Gessner etal. (1993), Sridhar
and Bärlocher (1993), Voglmayr
(1996), Czeczuga and Orłowska
(1999), Marvanová (2000),
Smits etal. (2007), Baschien
etal. (2008), Anderson and
Shearer (2011), Sudheep and
Sridhar (2012), Menéndez etal.
(2012), Cornut etal. (2014),
Raposeiro etal. (2018), Pietryc-
zuk etal. (2018)
Tetracladium maxilliforme Water (pond, river), submerged decaying wood
(Alnus glutinosa) in stream, foam (stream, rivulet),
foam (stream)
Austria, Canada,
France, New
Zealand,
Poland, USA,
Venezuela
Petersen (1962), Bärlocher (1987),
Gessner etal. (1993), Voglmayr
(1996), Czeczuga and Orłowska
(1999), Marvanová (2000),
Cooper (2005), Smits etal.
(2007), Pietryczuk etal. (2018),
Baudy etal. (2019)
Tetracladium nainitalense Root endophyte riparian plants, foam (stream) Brazil, India Sati etal. (2009), Fiuza and
Gusmão (2013a)
Tetracladium palmatum River spume, foam (stream, rivulets) Austria, Portugal,
Spain Roldán etal. (1989), Voglmayr
(1996), Marvanová (2000),
Marvanová etal. (2003),
Baschien etal. (2013)
Tetracladium setigerum Water (pond, river), foam (stream), submerged
leaves of Salix atrocinerea (river) and Alnus gluti-
nosa leaves (stream), sediment (stream)
Austria, Canada,
France, India,
Jamaica, Malay-
sia, Poland,
Portugal, Spain,
Switzerland,
Thailand, UK,
USA, Venezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Ranzoni (1953),
Petersen (1962), Nawawi
(1985b), Bärlocher (1987),
Roldán etal. (1989), Gessner
etal. (1993), Czeczuga and
Orłowska (1999), Marvanová
(2000), Smits etal. (2007),
Schoenlein-Crusius etal. (2009),
Sudheep and Sridhar (2012),
Raposeiro etal. (2018), Vu etal.
(2019)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Lauriomycetales
Lauriomycetaceae
Lauriomyces heliocephalus Submerged decaying leaf Brazil da Silva and Gusmão (2015)
Lauriomyces pulcher Submerged decaying wood (stream) Hong Kong Ho (1998)
Lauriomyces sakaeratensis Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Leotiales
Leotiales genera incertae sedis
Alatospora acuminata Unpolluted water (stream), foam (stream, river, rivu-
let), submerged decaying wood and leaves (Alnus
glutinosa, Acer pseudoplatanus, Corylus avellana,
Fraxinus excelsior, Fagus sylvatica, Picea abies,
Tilia cordata, Ulmus sp.), submerged leaves (Cof-
fea arabica and Hevea brasiliensis) (stream), water
(river), scum (stream), sediment (dam), stemflow
and throughfall of tree canopies and palm species,
water-filled treeholes
Austria, Brazil,
Canada, Czech
Republic,
France, Hungary,
India, Ireland,
Jamaica, Japan,
Netherlands,
New Zealand,
Poland, Portu-
gal, Rhodesia,
Uganda, South
Africa, Spain,
Sweden, Swit-
zerland, UK,
USA, Venezuela
Ingold (1942), Ingold (1943a),
Ingold (1949), Fenton (1950),
Ranzoni (1953), Tubaki (1957),
Umphlett (1957), Nilsson
(1958), Ingold (1958b), Nilsson
(1960), Hudson and Ingold
(1960), Ingold (1960), Great-
head (1961), Anastasiou (1964),
Iqbal and Webster (1973),
Almer and Segedin (1985),
Marvanová and Descals (1985),
Sridhar (1987), Bärlocher
(1987), Gessner etal. (1993),
Sridhar and Bärlocher (1993),
Voglmayr (1996), Czeczuga
(1996), Harrington (1997),
Marvanová (2000), Gönczöl and
Révay (2003), Belliveau and
Bärlocher (2005), Smits etal.
(2007), Baschien etal. (2008),
Sridhar and Karamchand (2009),
Sudheep and Sridhar (2012),
Menéndez etal. (2012), Fiuza
and Gusmão (2013a), Cornut
etal. (2014), Ghate and Sridhar
(2015), Duarte etal. (2015),
Funck etal. (2015), Raposeiro
etal. (2018), Pietryczuk etal.
(2018), Vu etal. (2019)
Alatospora crassipes Submerged decaying leaves ofFagus sylvatica in
(river) Czech Republic Marvanová (1977)
Alatospora flagellata Fagus sylvatica leaf (stream), foam (stream, rivulet),
submerged leaf litter of Alnus glutinosa (stream),
water (river)
Austria, Canada,
Czech Republic,
France, Hun-
gary, Poland
Marvanová and Descals (1985),
Bärlocher (1987), Gessner
etal. (1993), Voglmayr (1996),
Marvanová (2000), Baschien
etal. (2013), Pietryczuk etal.
(2018)
Alatospora pulchella Leaves (stream), foam (stream, rivulet), submerged
decaying leaves of Alnus (stream), water (stream),
stemflow and throughfall
Australia,
Austria, China,
Czech Republic,
France, India,
Italy, Nether-
lands, Portugal,
Spain, UK, USA
Marvanová and Descals (1985),
Au etal. (1992), Gessner
etal. (1993), Voglmayr 1996),
Marvanová (2000), Gulis and
Suberkropp (2003), Menéndez
etal. (2012), Baschien etal.
(2013), Ghate and Sridhar
(2015), Duarte etal. (2015)
Rhytismatales
Rhytismataceae
Hypoderma alpinum Carex nigra Austria Magnes and Hafellner (1991)
Hypohelion scirpinum Submerged decaying stalks of Carex nigra UK Ingold (1954)
Lophodermium licualae Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Lophodermium macci Water China Song and Wu (2010)

Fungal Diversity
1 3
Table 6 (continued)
Taxon Substrate Distribution References
Thelebolales
Thelebolaceae
Geomyces sp. Water distribution system Polland Grabińska-Łoniewska etal. (2007)
Thelebolales genera incertae sedis
Gorgomyces honrubiae Foam (stream) Spain Baschien etal. (2013)
Miniancora allisoniensis Foam (stream) Canada Baschien etal. (2013)
water (van Uden and Ahearn 1963; Sláviková etal. 1992;
Gadanho etal. 2006; de García etal. 2007; Russo etal. 2008;
Libkind etal. 2009; Branda etal. 2010; Brandão etal. 2011;
Fell etal. 2011; Medeiros etal. 2012). Most of the taxa of
freshwater Basidiomycota in Table12 were observed from
plant substates. Fungi from foam records 19 species, eight
(Leptosporomyces crucelliger, Taeniospora nasifera, Fibulo-
taeniella canadensis, Nodulospora inconstans, Helicogloea
angustispora, Anguillomyces acadiensis, A. acadiensis,
Arcispora bisagittaria) of which were reported exclusively
from foam. There are notions on fungi observed from foam
as spores originating from the neighboring terrestrial envi-
ronment and just trapped in air bubbles, however studies
showed that the vast majority of these spores originated from
aquatic habitats (Nilsson 1964; Ingold 1975). Thus, they are
no different from ascomycetous spores that are also recov-
ered from foam samples. Foam serves as a natural trap for
conidia and gives a fairly accurate indication of the fungal
flora of stream or river. For instance, Nilsson (1964) con-
cluded that the foam and scum samples give us an overview
of the species from the stream system in the sampling area.
In many cases, the host substrate is determined in subsequent
studies, e.g., Mrakia aquatica was initially isolated from
foam samples (Jones and Slooff 1966), but later collected
on Equisetum culms (Webster unpublished).
In this section, we update the list of Jones etal. (2014b)
with a total of 218 species (in 100 genera) (Tables12 and
13). Eleven classes comprise freshwater Basidiomycota and
distributed in 26 orders and 43 families.
Freshwater yeasts
Few freshwater yeasts are known when compared with
the extensive number of hyphomycetes and ascomycetes
documented in this study. This can be attributed to the
lack of studies undertaken (Fell etal. 2011; Libkind etal.
2014). Yeasts are eukaryotic microorganisms that belong
to the Ascomycota and Basidiomycota and require differ-
ent techniques to filamentous fungi for their study. Early
studies in their identification required not only isolation
and morphological observation but also chemical reactions
such as utilization and fermentation of different substrates.
The development of molecular techniques has revolution-
ized their study and enabled their higher order classification
(Kurtzman and Robnett 1998) with extensive revisions of the
classification of Basidiomycota yeasts by Liu etal. (2015c,
d) and Wang etal. (2015a, b, c). Kurtzman and Fell (2006)
estimated there were circa 1500 yeast species but the figure
is nearer to 2000 to 2200.
Freshwater yeasts have been recovered from a variety of
environments: drinking water (Yamaguchi etal. 2007), surface
water (Biedunkiewicz and Baranowska 2011), bottled and tap
water (Yamaguchi etal. 2007), wastewater and polluted rivers
(Spencer etal. 1970; Woollett and Hedrick 1970; Pires etal.
2017; Assress etal. 2019), rivers (van Uden and Ahearn 1963;
Sláviková and Vadkertiová 1997a, b), trapped in foam (Jones
and Slooff 1966), freshwater estuaries (Fell etal. 2011; Lib-
kind etal. 2017), glacial melt water (Libkind etal. 2003, 2009,
2014), fish ponds (Sláviková and Vadkertiová 1995a), artificial
lakes (Sláviková etal. 1992) and lakes (Kwasniewska 1988;
Brandão etal. 2010, 2011, 2017; Biedunkiewicz etal. 2013).
Yeasts have also been isolated from freshwater plants (Mra-
kia aquatica from Equisetum culms, Vishniacozyma victoriae
from submerged roots), while a wide range of taxa occur in
the water tanks of bromeliads: Bromelia karatas, Encholirium
sp., Paepalanthus bromelioides and Vriesea minarum (Morais
etal. 2020). Others occur on the surfaces of surface and mouth
cavity of two Chelonian (turtles) species, Podocnemis expansa
and P. unifilis (Morais etal. 2010).
Yeast distribution in lakes has yielded a wide range of
taxa (Biedunkiewicz etal. 2013) as well as those found in the
freshwater zone of the Everglades (Fell etal. 2011). Many of
these studies yielded novel species: Candida aquatica (cur-
rent name = Mrakia aquatica), Cryptococcus agrionensis,
Sporobolomyces patagonicus, Rhodotorula himalayensis
(= Phenoliferia himalayensis) and Rhodotorula meli (= Sak-
aguchia meli). With recent revisions of the basidiomycetous
yeasts, many of the early names are now obsolete and, in this
paper, we refer to the currently accepted names.
Many taxa listed in Table13 were only isolated once
in the various studies, for example, the 317 isolates made
by Medeiros etal. (2012) from lakes and rivers in Brazil,

Fungal Diversity
1 3
Table 7 Classification of freshwater Orbiliomycetes with information on the type of habitats, substrates, and geographical distribution of fungal
taxa
Taxon Substrate Distribution References
ASCOMYCOTA
ORBILIOMYCETES
Orbiliales
Orbiliaceae
Arthrobotrys dactyloides Submerged twig Brazil da Silva and Gusmão (2015)
Arthrobotrys oligosporus Water (river), submerged decaying
leaves (stream), decaying Licuala
longicalycata (peat swamp)
Brazil, Poland, Thailand Czeczuga etal. (1990), Barbosa etal.
(2013b) Pinruan etal. (2014)
Hyalorbilia oviparasitica Foam (stream) India Sridhar and Kaveriappa (1992),
Rajashekhar and Kaveriappa (2003)
Dactylella microaquatica Submerged decaying of unidenti-
fied leaves and Shiia sieboldii,
submerged decaying wood (river),
water (stream)
Cuba, Japan, Philippines, USA Petersen (1962), Marvanová and
Marvan (1969), Cai etal. (2003a),
Gulis and Suberkropp (2003) (Ran-
zoni (1979), Cai etal. (2003a)
Dactylella submersa Submerged decaying leaves of
Quercus, foam (stream), stemflow
and throughfall of palm species
Cuba, India, Ireland, Poland, UK Ingold (1943a), Ingold (1944),
Marvanová and Marvan (1969),
Czeczuga (1996), Rajashekhar
and Kaveriappa (2003), Ghate and
Sridhar (2015)
Dactylella yunnanensis Submerged leaf Brazil da Silva and Gusmão (2015)
Dicranidion fissile Rainwater from Quercus myrsinae-
folia leaves Japan Ando and Tubaki (1984a)
Dicranidion gracile Stemflow and throughfall of tree
canopies India Sridhar and Karamchand (2009)
Helicoon myosuroides Submerged decaying leaves of Calo-
phyllum brasiliense (stream) Brazil Fiuza etal. (2019), Fiuza etal.
(2017b)
Helicoon pluriseptatum Rainwater from building roofs,
leaves in water, submerged decay-
ing Quercus leaves (lake)
Ireland, Netherlands, Poland van Beverwijk (1954), Abdullah and
Webster (1980), Czeczuga and
Orłowska (1997)
Monacrosporium tentaculatum Foam (stream, river), submerged
leaves India, Thailand, USA Anastasiou (1964), Sridhar and
Kaveriappa (1992), Sakayaroj etal.
(2005)
Orbilia luteorubella Submerged decaying wood (river) USA Lamore and Goos (1978)
Orbilia ellipsospora Submerged decaying twigs and
leaves (stream), Calophyllum
brasiliense
Brazil Barbosa etal. (2013b) Fiuza etal.
(2019)
Orbiliella armeniaca Twigs USA Shearer and Von Bodman (1983)
Trinacrium incurvum Foam (river), stemflow and through-
fall of palm species Brazil, India Fiuza and Gusmão (2013a), Ghate
and Sridhar (2015)
Trinacrium indicum Submerged decaying leaves of Cof-
fea arabica (stream) India Soosamma etal. (2001)
Trinacrium robustum Stemflow and throughfall of tree
canopies and palm species India Sridhar and Karamchand (2009),
Ghate and Sridhar (2015)
Trinacrium subtile Rainwater from building roofs,
leaves in water, stemflow and
throughfall of tree canopies and
palm species
India, Netherlands, Poland Czeczuga and Orłowska (1997), Srid-
har and Karamchand (2009)
Vermispora cauveriana Foam (stream), submerged decaying
leaves of Ficus religiosa (river) India Rajashekhar and Kaveriappa (2003),
Rajashekhar etal. (1991)

Fungal Diversity
1 3
yielded 21 genera and 48 species but few were isolated from
each of the 18 sites. The most numerous genera are Can-
dida (31 species) and Rhodotorula (9 species), and with
fewer species Papiliotrema (6) and Naganishia (5). Com-
mon species included Cryptococcus luteolous (= Hannaella
luteola) and Rhodotorula glutinis in the dry season, while
Aureobasidium pullulans was common in the rainy season.
Brandão etal. (2011) noted that Candia krusei (= Teuno-
myces kruisii), and C. guilliermondii (= Meyerozyma guil-
liermondii) were the most common in three freshwater lakes
in Brazil. These are faecal coliforms, probably present in
the water samples as the result of contamination by warm-
blooded animals (Brandão etal. 2010). Other taxa occurred
only once and can be regarded as rare.
While most are regular taxa in freshwater environments:
Mrakia aquatica, M. frigida, Kluyveromyces aestuarii, oth-
ers have been described also from mangrove habitats (Kwo-
niella mangrovensis) (Statzell-Tallman etal. 2008), or from
glacial melt water, e.g. Cryptococcus agrionensis (= Gof-
feauzyma agrionensis), Cystofilobasidium lacus-mascardii,
and Sakaguchia meli (de García etal. 2007; Libkind etal.
2014). Yeasts such as Candida albicans, Candida parapsilo-
sis, Candida krusei (= Teunomyces kruisii), Candida guilli-
ermondii (= Meyerozyma guilliermondii), Candida glabrata
and Candida tropicalis, are opportunistic pathogens often
associated with human activity. Fell etal. (2011) however,
did not encounter serious human pathogens (such as in water
Candida tropicalis) in samples from the freshwater zone of
the Everglades.
A number of factors determine the occurrence of yeasts
in freshwater habitats: temperature, exposure/tolerance to
UV light, and nutrient availability. Libkind etal. (2006) con-
cluded that in Patagonian freshwater yeasts, there had an
apparent relationship between the ability to produce PPCs
(photoprotective compounds), their tolerance to UV radia-
tion (UVR) exposure and their success in colonizing habi-
tats highly exposed to UVR. Brandão etal. (2011) opined
those yeasts isolated from an oligotrophic lake in Patagonia
(Argentina) are metabolically active and could contribute to
organic matter recycling even under these harsh conditions.
Some 325 yeasts (162 Basidiomycota, 163 Ascomycota)
species are listed in Table13, but this figure is undoubtedly
an underestimate. These yeast taxa are not recorded in the
previous taxonomic groups above and we separately listed
in this section.
Ingoldian fungi
Ingoldian fungi, also known as “aquatic (Ingoldian) hypho-
mycetes”, were named after Prof. Cecil Terence Ingold
Table 8 Classification of freshwater Pezizomycetes with information on the type of habitats, substrates, and geographical distribution of fungal
taxa
Taxon Substrate Distribution References
PEZIZOMYCETES
Pezizales
Ascobolaceae
Saccobolus beckii Pine bark India Udaiyan (1989)
Thecotheus rivicola Twigs and wood of Prunus Czech Republic, USA Pfister (1973a)
Pezizaceae
Adelphella babingtonii Twigs France, Hungary, UK, USA Ingold (1954), Pfister (1973b), Pfister etal. (2008)
Aquapeziza globispora Wood China Hu etal. (2012c)
Pachyella adnata Wood USA, Japan Pfister (1973b)
Pachyella clypeata Wood USA, Japan Pfister (1973b)
Pachyella punctispora Wood USA Pfister (1973b)
Pachyella violaceonigra Wood Ukraine Dudka (1963)
Peziza oliviae Wood USA Frank (2013)
Peziza sp. Wood UK Eaton and Jones (1971a)
Pyronemataceae
Miladina lecithina Wood, foam (stream) Austria, UK, USA Pfister and Korf (1974), Descals and Webster
(1978), Marvanová (2000)
Scutellinia scutellata Wood Ukraine Dudka (1963)
Rhizinaceae
Psilopezia deligata Poplar log or unidentified wood USA, Argentina Pfister (1973a)
Psilopezia nummularia Wood USA Pfister (1973a)

Fungal Diversity
1 3
Fig. 5 Freshwater Sordariomycetes. a, b Sporidesmium sp. c,
d Aquapteridospora sp. e, f Canalisporium sp. g–i Cordana sp. j–l
Sporidesmium sp. m, n Sprochisma sp. o, p Phaeoisaria sp. q, r Dis-
toseptispora sp. s, t Xylaria sp. u, v Ceratosphaeria sp. w–y Peroneu-
typa sp. z, a1 Cosmospora aquatica. Scale bars: s, w, z = 300μm, a,
c, e, g, j, k, m, o, q = 100 μm, r = 40μm, f, l, v = 20μm, b, d, h, i, n,
p, t, x, y, a1 = 10μm

Fungal Diversity
1 3
Table 9 Classification of freshwater Sordariomycetes with information on the type of habitats, substrates, and geographical distribution of fun-
gal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
SORDARIOMYCETES
Diaporthomycetidae
Annulatascales
Annulatascaceae
Annulatascus apiculatus Submerged decaying wood (stream) Brazil Barbosa etal. (2008)
Annulatascus aquaticus Submerged decaying wood (stream) China Ho etal. (1999c)
Annulatascus chiangmaiensis Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Annulatascus fusiformis Submerged decaying wood (stream) Australia,
China, Phil-
ippines
Hyde and Wong (2000), Cai
etal. (2006), Vijaykrishna
etal. (2006)
Annulatascus hongkongensis Submerged decaying wood (reservoir) China Ho etal. (1999b)
Annulatascus joannae Submerged decaying wood (river) China Tsui etal. (2002)
Annulatascus lacteus Submerged decaying wood (river) China Tsui etal. (2002)
Annulatascus liputii Submerged decaying bamboo (river) Philippines Cai etal. (2002a)
Annulatascus menglensis Submerged decaying wood (stream) China Hu etal. (2012a)
Annulatascus nakhonensis Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Annulatascus palmietensis Submerged decaying wood (river) Brunei, South
Africa Hyde etal. (1998a), Fryar etal.
(2004), Hyde and Sarma
(2006)
Annulatascus saprophyticus Submerged decaying wood (stream) Thailand Luo etal. (2015)
Annulatascus songkhlaensis Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Annulatascus thailandensis Submerged decaying wood (stream) Thailand Hyde etal. (2020d)
Annulatascus tropicalis Submerged decaying wood (stream) China Tsui etal. (2002)
Annulatascus velatisporus Submerged decaying wood (stream, river, waterfall) and
bamboo (river), submerged decaying Licuala longicaly-
cata and Eleiodoxa conferta (peat swamp)
Australia,
Brazil, Bru-
nei, China,
Philippines,
India, Japan,
Malaysia,
South
Africa,
Thailand
Hyde (1992c), Hyde etal.
(1998a), Wong etal. (1999b),
Ho etal. (2001), Cai etal.
(2003a), Tsui etal. (2003c),
Pinnoi etal. (2006), Hyde
(2007) Pinruan etal. (2007),
Hu etal. (2010b), Sudheep
and Sridhar (2011), Barbosa
etal. (2013a), Dayarathne
etal. (2016)
Annulusmagnus triseptatus Submerged decaying wood (river, stream) Australia, Bru-
nei, Canada,
China,
Malaysia,
USA, Ven-
ezuela
Wong etal. (1999b), Ho etal.
(2001), Campbell and Shearer
(2004)
Aqualignicola hyalina Submerged decaying wood (river) China, India Ranghoo etal. (2001), Sudheep
and Sridhar (2011)
Aqualignicola vaginata Submerged decaying wood (stream) China Hu etal. (2012a)
Ascitendus aquaticus Submerged decaying wood (river) Australia Hyde etal. (2018)
Ascitendus austriacus Submerged decaying wood Australia,
Canada,
USA, Ven-
ezuela
Réblová and Winkac (2001),
Campbell and Shearer (2004)
Ayria appendiculata Submerged decaying wood USA Raja etal. (2009b)
Ayria nubispora Submerged decaying wood (lake, stream) Costa Rica,
USA Raja etal. (2009a)
Cataractispora appendiculata Submerged decaying wood (river, stream) Australia,
Brunei,
Seychelles
Hyde etal. (1999b)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Cataractispora aquatica Submerged decaying wood (river, stream, waterfall) Australia,
Brunei,
Malaysia,
Seychelles
Hyde etal. (1999b, Kurniawati
etal. (2010)
Cataractispora bipolaris Submerged decaying wood (river, stream) Australia,
Brunei,
Malaysia,
USA
Hyde (1992c), Ho etal. (2001),
Raja etal. (2009b)
Cataractispora receptaculorum Submerged decaying wood and bamboo (reservoir) China Ho etal. (2004)
Cataractispora viscosa Submerged decaying wood (stream) Australia,
China,
Malaysia,
Mauritius
Hyde etal. (1999b, Ho etal.
(2001)
Chaetorostrum quincemilense Submerged decaying wood (stream) Peru Zelski etal. (2011b)
Fusoidigranularius nilensis Submerged decaying wood (Phragmites australis) (river) Egypt Abdel-Wahab etal. (2011)
Longivarius aquatorbae Submerged decaying wood (Erythrophleum teysmannii)Thailand Boonyuen etal. (2012)
Longicollum biappendiculatum Submerged decaying wood (stream) Brazil, Costa
Rica, Peru,
USA
Zelski etal. (2011a)
Submersisphaeria aquatica Submerged decaying wood, submerged decaying Licuala
longicalycata (peat swamp) Australia,
France,
Thailand,
USA
Hyde (1996), Campbell etal.
(2003b), Pinruan etal. (2007),
Fournier etal. 2016)
Submersisphaeria palmae Submerged decaying Eleiodoxa conferta petiole and rachis,
Nenga pumila rachis, Licuala longecalycata trunk and
petiole, (peat swamp)
Thailand Pinnoi etal. (2004, 2006)
Vertexicola ascoliberata Submerged decaying wood (stream) Costa Rica Barbosa etal. (2013a)
Vertexicola caudata Submerged decaying wood China Ranghoo etal. (2000)
Aquapteridosporaceae
Aquapteridospora aquatica Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Aquapteridospora bambusi-
num Submerged decaying wood Thailand Bao etal. (2021a)
Aquapteridospora lignicola Submerged decaying wood (stream) Thailand Yang etal. (2015)
Aquapteridospora fusiformis Submerged decaying wood (stream) China Luo etal. (2019)
Atractosporales
Atractosporaceae
Atractospora aquatica Submerged decaying wood (stream) China Luo etal. (2019)
Atractospora decumbens Submerged decaying wood (stream) France Réblová etal. (2016a)
Atractospora ellipsoidea Submerged decaying wood Brunei,
China, USA Ho etal. (1999d), Fryar etal.
(2004), Campbell and Shearer
(2004), Dong etal. (2021a)
Atractospora reticulata Submerged decaying Sambucus nigra wood (stream) France Réblová etal. (2016a)
Atractospora thailandensis Submerged decaying wood (river) Thailand Zhang etal. (2017a)
Atractospora verruculosa Submerged decaying wood France Réblová etal. (2016a)
Rubellisphaeria abscondita Submerged decaying wood France Réblová etal. (2016a)
Jobellisiales
Jobellisiaceae
Jobellisia guangdongensis Submerged decaying wood (stream) China Liu etal. (2012), Chethana etal.
(2021)
Jobellisia luteola Submerged decaying wood USA Raja etal. (2009b)
Jobellisia viridifusca Submerged decaying wood (river) China Ranghoo etal. (2001)
Barbatosphaeriales
Barbatosphaeriaceae
Barbatosphaeria lignicola Submerged decaying wood (stream) China Luo etal. (2019)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Cancellidiales
Cancellidiaceae
Cancellidium applanatum Submerged decaying wood of Dipterocarpus alatus and
Xylia dolabriformis test blocks, unidentified wood
(stream), submerged decaying Licuala longicalycata and
Eleiodoxa conferta (peat swamp)
Brazil, Perú,
Thailand Sivichai etal. (2002), Pinnoi
etal. (2006), Pinruan etal.
(2007), Chuaseeharonnachai
etal. (2013), Barbosa etal.
(2013b), Zelski etal. (2014)
Cancellidium atrobrunneum Submerged decaying wood (stream) Thailand Dong etal. (2021a), Hyde etal.
(2021)
Cancellidium cinereum Submerged decaying wood (stream) China Hyde etal. (2021)
Cancellidium griseonigrum Submerged decaying wood (stream) Thailand Hyde etal. (2021)
Cancellidium thailandense Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Obliquiminima hyalina Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Ceratolentales
Bullimycetaceae
Bullimyces aurisporus Submerged decaying wood (stream) Costa Rica Ferrer etal. (2012)
Bullimyces communis Submerged decaying wood (reserve, stream) Costa Rica Ferrer etal. (2012)
Bullimyces costaricensis Submerged decaying wood (stream) Costa Rica Ferrer etal. (2012)
Conlariales
Conlariaceae
Conlarium aquaticum Submerged decaying wood (river, stream) China, Thai-
land Zhang etal. (2017a), Dong
etal. (2021a)
Conlarium dupliciascosporum Submerged decaying wood (stream) China Liu etal. (2012)
Conlarium subglobosum Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Riomyces rotundus Submerged decaying wood (stream) Costa Rica Ferrer etal. (2012)
Diaporthales
Diaporthaceae
Diaporthe aquatica Submerged decaying wood (ditch) China Hu etal. (2012b)
Diaporthe beckhausii Submerged decaying wood (stream) China Ho etal. (2001)
Diaporthe longicolla Freshwater Korea Nam etal. (2020)
Gnomoniaceae
Ambarignomonia petiolorum Submerged partially decomposed Acer rubrum leaves (lake) USA Fallah and Shearer (2001)
Gnomonia papuana Submerged leaves Papua New
Guinea Sivanesan and Shaw (1977)
Gnomoniella microspora Submerged decaying wood (stream) Malaysia Ho etal. (2001)
Gnomoniella rubicola Submerged decaying wood (stream) Brunei Ho etal. (2001)
Sydowiellaceae
Sillia ferruginea Submerged decaying wood UK Eaton and Jones (1971a, b)
Diaporthales genera incertae sedis
Phruensis brunneispora Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2004a, 2007)
Distoseptisporales
Distoseptisporaceae
Distoseptispora appendiculata Submerged decaying wood (stream) Thailand Luo etal. (2019)
Distoseptispora aquatica Submerged decaying wood (stream) China Su etal. (2016a)
Distoseptispora cangshanensis Submerged decaying wood (stream) China Luo etal. (2018b)
Distoseptispora euseptata Submerged decaying wood (river) China Li etal. (2021c)
Distoseptispora fasciculata Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Distoseptispora fluminicola Submerged decaying wood (stream) China Su etal. (2016a)
Distoseptispora guttulata Submerged decaying wood (stream) Thailand Yang etal. (2018d), Luo etal.
(2019)
Distoseptispora lignicola Submerged decaying wood (waterfall) Thailand Luo etal. (2019)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Distoseptispora martinii Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Distoseptispora multiseptata Submerged decaying wood (stream) Thailand Hyde etal. (2016b), Luo etal.
(2019)
Distoseptispora neorostrata Submerged decaying wood (stream) Thailand Luo etal. (2019)
Distoseptispora obclavata Submerged decaying wood (stream) Thailand Luo etal. (2019)
Distoseptispora obpyriformis Submerged decaying wood (river) China Luo etal. (2018b)
Distoseptispora phangngaen-
sis Submerged decaying wood (stream) Thailand Yang etal. (2018d)
Distoseptispora rayongensis Submerged decaying wood (stream) Thailand Hyde etal. (2020d)
Distoseptispora rostrata Submerged decaying wood (river) China Luo etal. (2018b)
Distoseptispora saprophytica Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Distoseptispora songkhlaensis Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Distoseptispora submersa Submerged decaying wood (river) China Luo etal. (2018b)
Distoseptispora suoluoensis Submerged decaying wood (river) China Yang etal. (2018d)
Distoseptispora tectonae Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Distoseptispora yunnanensis Submerged decaying wood (river) China Li etal. (2021c)
Magnaporthales
Ceratosphaeriaceae
Ceratosphaeria aquatica Submerged decaying wood (river) China Luo etal. (2019)
Ceratosphaeria lampadophora Submerged decaying wood (water-cooling tower) Australia Jones (1972), Hyde etal. (1997)
Ceratosphaeria phialidica Submerged decaying wood (river) USA Shearer (1989a)
Ceratosphaeria pusilla Submerged decaying wood (swamp) USA Shearer and Crane (1986)
Ceratosphaeria lignicola Submerged decaying wood (stream) Thailand Luo etal. (2019)
Magnaporthaceae
Aquafiliformis lignicola Submerged decaying wood (river, stream) China Luo etal. (2019)
Buergenerula biseptata Carex rostrata Austria Magnes and Hafellner (1991)
Clasterosporium flexum Foam (stream), submerged decaying Thuja plicata foliage
(pond) Japan, UK Abdullah (1980), Hosoya and
Tanaka (2007)
Gaeumannomyces graminis Carex rostrata Australia Magnes and Hafellner (1991)
Ophioceraceae
Ophioceras aquaticum Submerged decaying wood (reservoir) China Hu etal. (2012b)
Ophioceras arcuatisporum Submerged Typha sp. (lake) Canada,
China, USA Shearer etal. (1999), Luo etal.
(2004)
Ophioceras commune Submerged decaying wood (lake, stream, river, waterfall) China, Egypt,
Japan,
Malaysia,
Panama,
Thailand,
USA
Shearer etal. (1999), Tsui etal.
(2001d, 2003c), Luo etal.
(2004), Hyde (2007) Raja
etal. (2009b), Kurniawati
etal. (2010), Abdel-Aziz
(2016b), Chethana etal.
(2021)
Ophioceras dolichostomum Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (lake, river) Australia, Bru-
nei, China,
Japan,
Malaysia,
Philippines,
Seychelles,
Thailand,
USA
Conway and Barr (1977), Hyde
(1992b), Hyde and Goh
(1998c), Sivichai etal. (2002),
Cai etal. (2003a), Tsui etal.
(2003c), Luo etal. (2004)
Ophioceras fusiforme Submerged decorticated woody debris (stream) USA Shearer etal. (1999)
Ophioceras guttulatum Submerged decaying wood (river, stream) China Tsui etal. (2001g), Cai etal.
(2006)
Ophioceras hongkongense Submerged decaying wood (river) China Tsui etal. (2001h)
Ophioceras leptosporum Submerged plant stalks UK Iqbal (1972b)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Ophioceras submersum Submerged decaying wood (waterfall) Thailand Luo etal. (2019)
Ophioceras tenuisporum Submerged decaying twig (creek) Panama Shearer etal. (1999)
Ophioceras venezuelense Submerged decaying wood (stream) Brazil, Ven-
ezuela Shearer etal. (1999)
Pseudohalonectriaceae
Pseudohalonectria adversaria Submerged decaying wood (swamp, stream) China, Malay-
sia, South
Africa, UK
Shearer (1989b), Shearer
(1989a), Hyde etal. (1998a),
Hyde and Goh (1999a), Ho
etal. (2001), Luo etal. (2004)
Pseudohalonectria falcata Submerged decaying wood (river) USA Shearer (1989a)
Pseudohalonectria fuxianii Submerged decaying wood (lake) China Cai etal. (2002b)
Pseudohalonectria lignicola Submerged decaying wood and Acer saccharinum twig
(creek, lake, stream, waterfall) China, Japan,
Thailand,
USA
Minoura and Muroi (1978),
Shearer (1989b), Shearer
(1989a), Cai etal. (2002b),
Tsui etal. (2003c), Luo etal.
(2004), Kurniawati etal.
(2010)
Pseudohalonectria longiro-
strum Submerged decaying wood (river) Panama, Phil-
ippines Shearer (1989a), Cai etal.
(2003a)
Pseudohalonectria lutea Submerged decaying Quercus wood (stream) China, Chile,
USA Shearer (1989a), Fallah and
Shearer (2001), Cai etal.
(2002b)
Pyriculariaceae
Neocordana musae Submerged decaying wood (stream) China Ho etal. (2001)
Pyricularia fusispora Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Pyricularia rabaulensis Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Myrmecridiales
Myrmecridiaceae
Myrmecridium aquaticum Submerged decaying wood (river, stream) China Luo etal. (2019)
Myrmecridium fluviae Freshwater Korea Tibpromma etal. (2017)
Myrmecridium montseguri-
num Submerged wood of Fraxinus excelsior, Fagus sylvatica,
Hedera helix and Alnus glutinosa (stream) France Réblová etal. (2016a)
Myrmecridium schulzeri Water (stream), submerged decaying wood China, South
Korea Nguyen etal. (2018), Wijaya-
wardene etal. (2021)
Ophiostomatales
Ophiostomataceae
Subbaromyces aquaticus Submerged decaying wood India Chary and Ramarao (1974)
Parasympodiellales
Parasympodiellaceae
Parasympodiella laxa Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Phomatosporales
Phomatosporaceae
Phomatospora aquatica Submerged decaying wood (stream) Australia,
Japan Minoura and Muroi (1978),
Vijaykrishna and Hyde (2006)
Phomatospora berkeleyi Submerged decaying wood (Typha latifolia, Carex comosa,
Scirpus brevicaudatus), wood (lake, river, stream), decay-
ing Licuala longicalycata (peat swamp)
Australia,
China,
Philippines,
Thailand,
USA
Fallah and Shearer (2001),
Vijaykrishna and Hyde (2006)
Phomatospora helvetica Submerged decaying wood (stream) Norway Nordén etal. (2015)
Phomatospora luteotingens Submerged decaying wood (Alnus glutinosa, Fraxinus
excelsior)France, Spain Fournier and Lechat (2010)
Phomatospora muskellungensis Submerged decaying wood and stem of Typha latifolia
(lake) USA Fallah and Shearer (1998)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Phomatospora striatigera Submerged Carex gracili Australia Scheuer (1988)
Phomatospora triseptata Submerged decaying wood (swamp) USA Raja and Shearer (2008)
Rhamphoriales
Rhamphoriaceae
Rhamphoria pyriformis Submerged decaying wood (stream) China Ho etal. (2001)
Rhodoveronaea aquatica Submerged decaying wood (stream) China Luo etal. (2019)
Sporidesmiales
Sporidesmiaceae
Sporidesmium altum Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Sporidesmium appendicula-
tum Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Sporidesmium aquaticivagi-
natum Submerged decaying wood (stream) Thailand Hyde etal. (2016b)
Sporidesmium aturbinatum Submerged decaying wood China Bao etal. (2021a)
Sporidesmium brachypus Submerged decaying wood (river) China Luo etal. (2019)
Sporidesmium cangshanense Submerged decaying wood (stream) China Luo etal. (2019)
Sporidesmium chiangmaiense Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Sporidesmium dulongense Submerged decaying wood (river) China Hyde etal. (2020d)
Sporidesmium ensiforme Foam (tarn) UK Descals and Webster (1982)
Sporidesmium fluminicola Submerged decaying wood (stream) China Su etal. (2016a)
Sporidesmium gyrinomor-
phum Submerged decaying wood (stream) Thailand Yang etal. (2018d)
Sporidesmium lageniforme Submerged decaying wood (river) China Luo etal. (2019)
Sporidesmium lignicola Submerged decaying wood (river, stream) China Luo etal. (2019)
Sporidesmium macrurum Submerged decaying wood (stream) Brunei Ho etal. (2001)
Sporidesmium olivaceoco-
nidium Submerged decaying wood (stream) Thailand Hyde etal. (2016b)
Sporidesmium nujiangense Submerged decaying wood (river) China Bao etal. (2021a)
Sporidesmium paludosum Submerged decaying wood (lake) and bamboo (river) China, Philip-
pines Cai etal. (2003a), Luo etal.
(2004)
Sporidesmium ponapense Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Sporidesmium pyriformatum Submerged decaying wood (stream) Thailand Hyde etal. (2016b)
Sporidesmium submersum Submerged decaying wood (stream) China Su etal. (2016a)
Sporidesmium thailandense Submerged decaying wood (river) Thailand Zhang etal. (2017a, b), Yang
etal. (2018d)
Sporidesmium tropicale Foam (river), submerged decaying wood (stream) Brazil, China,
Thailand Sakayaroj etal. (2005), Barbosa
etal. (2013b), Yang etal.
(2018d), Bao etal. (2021a)
Tirisporellales
Tirisporellaceae
Thailandiomyces bisetulosus Submerged decaying wood, decaying Licuala longicalycata
(peat swamp) Thailand Pinruan etal. (2008, 2014)
Tirisporella beccariana Foam (stream) Malaysia Jones etal. (1996)
Togniniales
Togniniaceae
Erostella minutissima Submerged decaying wood (stream) China Ho etal. (2001)
Phaeoacremonium aquaticum Submerged decaying wood (stream) China Hu etal. (2012b)
Phaeoacremonium ovale Submerged decaying wood (stream) China Huang etal. (2018b)
Phaeoacremonium thailan-
dense Submerged decaying wood (river) Thailand Calabon etal. (2021a)
Trichosphaeriales
Trichosphaeriaceae

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Brachysporium obovatum Submerged decaying wood USA Raja etal. (2009b)
Brachysporium nigrum Submerged decaying wood USA Lamore and Goos (1978)
Koorchaloma bambusae Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Unisetosphaeria penguinoides Submerged petiole of Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2003a, 2006)
Xenospadicoidales
Xenospadicoidaceae
Neospadicoides aquatica Submerged decaying wood (stream) China Luo etal. (2019)
Neospadicoides lignicola Submerged decaying wood (stream) China Luo etal. (2019)
Neospadicoides thailandica Submerged decaying wood Thailand Bao etal. (2021a)
Neospadicoides yunnanensis Submerged decaying wood (river) China Luo etal. (2019)
Spadicoides americana Submerged decaying wood (stream) China Wongsawas etal. (2008)
Spadicoides atra Submerged decaying wood (stream) China Zhang etal. (2017a)
Spadicoides bambusicola Submerged decaying wood (stream) China Cai etal. (2006)
Spadicoides cordanoides Submerged decaying wood (stream) Australia Goh and Hyde (1996b)
Spadicoides hodgkissii Submerged decaying wood (stream) China Ho etal. (2002)
Spadicoides klotzschii Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Spadicoides minuta Submerged decaying Phyllostachys bambusoides (stream) China Cai etal. (2004a)
Spadicoides obovata Submerged decaying wood, decaying Licuala longicalycata
(peat swamp) China, Thai-
land Zhuang etal. (2001), Pinruan
etal. (2014)
Spadicoides stoveri Decaying Nenga pumila (peat swamp) Thailand Pinruan etal. (2014)
Spadicoides subsphaerica Submerged decaying wood (stream) Brazil Fiuza etal. (2017b)
Torrentispora aquatica Submerged decaying wood (stream) China Vijaykrishna etal. (2005)
Torrentispora biatriispora Submerged decaying wood (river, stream) Australia,
China, Costa
Rica, Sey-
chelles
Hyde (1995d), Hyde and Goh
(1998c), Tsui etal. (2002),
Barbosa etal. (2013a)
Torrentispora calembola Submerged decaying wood of Fagus sylvatica and Fraxinus
excelsior (lake) Czech Repub-
lic, France Réblová etal. (2018)
Torrentispora crassiparietis Submerged decaying wood (river, stream) Brazil, Brunei,
Costa Rica Fryar etal. (2004), Fryar and
Hyde (2004), Barbosa etal.
(2013a)
Torrentispora fibrosa Submerged decaying wood (lake, reservoir, stream) China, New
Zealand Hyde etal. (2000), Réblová
etal. (2018)
Torrentispora fusiformis Submerged decaying wood (river) Brunei Fryar and Hyde (2004)
Torrentispora pilosa Submerged decaying wood (stream) Costa Rica Barbosa etal. (2013a)
Diaporthomycetidae family incertae sedis
Acrodictyaceae
Acrodictys aquatica Submerged decaying wood (stream) Thailand Hyde etal. (2018), Luo etal.
(2019)
Acrodictys fluminicola Submerged decaying wood (stream) China Luo etal. (2019)
Acrodictys liputii Submerged decaying wood and bamboo (river) Philippines,
Thailand Cai etal. (2002c, 2003a), Bao
etal. (2021a)
Acrodictys porosiseptata Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Acrodictys sacchari Submerged decaying bamboo (river) Philippines Cai etal. (2003a)
Acrodictys similis Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Junewangiaceae
Dictyosporella aquatica Submerged decayed stem of Phragmites australis (river) Egypt Ariyawansa etal. (2015)
Dictyosporella chiangmaiensis Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Dictyosporella ellipsoidea Submerged decaying wood (stream) China Dong etal. (2021a)
Dictyosporella guizhouensis Submerged decaying wood (river) China Yuan etal. (2020)
Dictyosporella hydei Submerged decaying wood (stream) China Song etal. (2018a)
Dictyosporella thailandensis Submerged decaying wood (river) Thailand Zhang etal. (2017a)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Jennwenomyces navicularis Submerged decaying culm of Miscanthus floridulus (stream) China Goh and Kuo (2020)
Junewangia aquatica Submerged decaying wood China Song etal. (2018b)
Junewangia thailandica Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Sporidesmiella aquatica Submerged decaying wood (stream) China Luo etal. (2019)
Sporidesmiella aspera Submerged decaying angiosperm twigs (stream) Malaysia Kuthubutheen and Nawawi
(1993a)
Sporidesmiella ciliaspora Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Sporidesmiella claviformis Submerged decaying twigs (stream) Malaysia Kuthubutheen and Nawawi
(1993a)
Sporidesmiella cornuta Submerged decaying twigs (stream) Malaysia Kuthubutheen and Nawawi
(1993a)
Sporidesmiella incrassata Submerged decaying twigs (stream) Malaysia Kuthubutheen and Nawawi
(1993a)
Sporidesmiella hyalosperma Submerged decaying wood (river, stream, swamp) China, USA Crane (1972), Luo etal. (2019)
Sporidesmiella novae-zelan-
diae Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) China, Thai-
land Sivichai etal. (2002), Luo etal.
(2019)
Sporidesmiella obovoidia Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Sporidesmiella oraniopsidis Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Sporidesmiella parvissima Submerged decaying wood and leaves (stream) Brazil, Malay-
sia Kuthubutheen and Nawawi
(1993a), Barbosa etal.
(2013b)
Sporidesmiella vignalensis Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Papulosaceae
Brunneosporella aquatica Submerged decaying wood (reservoir) China Ranghoo etal. (2001)
Fluminicola aquatica Submerged decaying wood (river) Thailand Zhang etal. (2017a, b)
Fluminicola bipolaris Submerged decaying wood (river, stream, waterfall) China,
Philippines,
Thailand
Wong etal. (1999a), Cai etal.
(2003a), Luo etal. (2004),
Kurniawati etal. (2010)
Fluminicola saprophytica Submerged decaying wood (river) Thailand Zhang etal. (2017a, b), Luo
etal. (2019)
Fluminicola striata Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Wongia aquatica Submerged decaying wood China Luo etal. (2019)
Wongia fusiformis Submerged decaying wood Thailand Bao etal. (2021a)
Pseudoproboscisporaceae
Diluviicola capensis Submerged decaying wood (stream) Brunei, Thai-
land Hyde etal. (1998b), Dong etal.
(2021a)
Neodiluviicola aquatica Submerged decaying bamboo (river) Thailand Zhang etal. (2017a)
Pseudoproboscispora aquatica Submerged decaying wood Australia,
Philippines,
Japan
Wong and Hyde (1999), Tsui
etal. (2003c)
Pseudoproboscispora caudae-
suis Submerged decaying wood (Acer ruburm, Fraxinus excel-
sior) (lake) UK, USA Ingold (1951), Fallah and
Shearer (2001), Campbell
etal. (2003b)
Pseudoproboscispora thailan-
densis Submerged decaying bamboo (river) Thailand Zhang etal. (2017a)
Pseudostanjehughesiaceae
Pseudostanjehughesia aqui-
tropica Submerged decaying wood (stream) Thailand Yang etal. (2018d)
Pseudostanjehughesia ligni-
cola Submerged decaying wood (stream) China Luo etal. (2019)
Woswasiaceae
Cyanoannulus petersenii Submerged decaying wood (stream) USA Raja etal. (2003)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Diaporthomycetidae genera incertae sedis
Aquaticola hyalomura Submerged decaying wood (stream) China Ho etal. (1999d)
Aquaticola longicolla Submerged decaying wood (creek, river) Australia,
Brunei Tsui etal. (2003b), Fryar etal.
(2004)
Aquaticola minutiguttulata Submerged decaying wood China Tsui etal. (2003b)
Aquaticola triseptata Submerged decaying wood (river) China Tsui etal. (2003b)
Aquidictyomyces appendicu-
latus Submerged decaying wood (stream) Thailand Dong etal. (2021a)
Ceratostomella hyalocoronata Submerged decaying wood (stream) China Inderbitzin (2000)
Clohiesia corticola Submerged decaying wood (river) Australia,
Japan Hyde (1995d), Tsui etal.
(2003c)
Clohiesia curvispora Submerged decaying wood (stream) China Cai and Hyde (2007c)
Clohiesia lignicola Submerged decaying wood (river) China Tsui etal. (1998a)
Hyalorostratum brunneispo-
rum Submerged decaying wood (stream) USA Raja etal. (2010a)
Hypocreomycetidae
Coronophorales
Bertiaceae
Bertia convolutispora Submerged decaying wood (stream) Australia Hyde (1995e)
Chaetosphaerellaceae
Chaetosphaerella phaeostroma Submerged bark Brazil da Silva and Gusmão (2015)
Nitschkiaceae
Rhagadostoma lichenicola Lake USA Hafellner (2014)
Coronophorales genera incertae sedis
Papulaspora pulmonaria Submerged decaying leaves UK, Nether-
lands van Beverwijk (1954)
Papulaspora sepedonioides Submerged bamboo culms (stream) China Cai etal. (2006)
Glomerellales
Australiascaceae
Dischloridium gloeosporioides Submerged leaves Brazil da Silva and Gusmão (2015)
Dischloridium inaequiseptatum Submerged decaying wood (stream) Brazil, China Ho etal. (2001), Barbosa etal.
(2013b)
Glomerellaceae
Colletotrichum godetiae Submerged decaying leaves (stream) Korea Goh etal. (2020)
Plectosphaerellaceae
Acrostalagmus luteoalbus Water (pond, stream) Korea Nguyen etal. (2019a)
Lectera nordwiniana Freshwater Korea Lim etal. (2021)
Reticulascaceae
Cylindrotrichum aquaticum Submerged decaying wood (stream) China Hyde etal. (2016b), Luo etal.
(2019)
Cylindrotrichum clavatum Submerged decaying wood (Alnus glutinosa, Fraxinus sp.,
Platanus sp.) (river) China, France Maharachchikumbura etal.
(2018)
Cylindrotrichum gorii Submerged decaying wood (river) China Maharachchikumbura etal.
(2018)
Cylindrotrichum helisciforme Foam (stream) Canada, Slo-
vakia Marvanová (1979), Bärlocher
(1987)
Cylindrotrichum oligospermum Submerged decaying wood (river) UK Hyde and Goh (1999a)
Cylindrotrichum submersum Submerged decaying wood (stream) China Luo etal. (2019)
Kylindria aquatica Submerged decaying wood (stream) China Maharachchikumbura etal.
(2018)
Kylindria chinensis Submerged decaying wood (stream) China Maharachchikumbura etal.
(2018)
Hypocreales

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Bionectriaceae
Acremonium guillematii Freshwater Korea Lim etal. (2021)
Bionectria byssicola Submerged decaying wood (stream) Brunei, China,
Malaysia Ho etal. (2001)
Didymostilbe australiensis Submerged decaying wood (stream) Australia Hyde and Goh (1997)
Fusariella sarniensis Submerged decaying wood China Chen etal. (2011)
Ovicuculispora parmeliae River watershed Canada Maloles etal. (2018)
Hypocreaceae
Gliocladium cylindrosporum Submerged decaying wood (stream) China Ho etal. (2001)
Sibirina orthospora Submerged decaying wood (stream) Brunei, China Ho etal. (2001)
Trichoderma glaucum Submerged decaying wood China Au etal. (1992)
Trichoderma harzianum Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Trichoderma polysporum Submerged decaying wood (stream) China Ho etal. (2001)
Myrotheciomycetaceae
Emericellopsis terricola Submerged organic debris and freshwater muds of Lake
Sengari Ireland,
Japan Park (1972), Tubaki (1973)
Nectriaceae
Aquanectria devians Submerged decaying leaf (reservoir) Singapore Gordillo and Decock (2019)
Aquanectria jacinthicolor Submerged decaying wood (stream) China Huang etal. (2018a)
Aquanectria penicillioides Submerged decaying wood and leaves of broad-leaved trees,
shrubs and angiosperm, foam (lake, spring, stream, river),
sediment (dam), stemflow and throughfall of tree canopies
and palm species
Australia,
China,
Cuba,
India, Italy,
Ireland,
Jamaica,
Korea, New
Zealand,
Portugal,
South
Africa,
Spain, UK,
USA
Ingold (1943a), Ranzoni (1953,
1956), Hudson and Ingold
(1960), Greathead (1961),
Crane (1968), Yu etal.
(1988), Au etal. (1992), Har-
rington (1997), Sridhar and
Karamchand (2009), Duarte
etal. (2012), Sudheep and
Sridhar (2012), Ghate and
Sridhar (2015), Lombard etal.
(2015), Mun etal. (2016), Luo
etal. (2019)
Aquanectria submersa Submerged decaying leaves of Alnus glutinosa (stream),
foam (stream) Jamaica,
Portugal,
Venezuela
Hudson (1961), Marvanova
etal. (2003), Smits etal.
(2007)
Aquanectria tenuispora Submerged decaying leaf (reservoir) Singapore Gordillo and Decock (2019)
Bacillispora inflata Water (river) Poland Czeczuga etal. (1990)
Baipadisphaeria spathulos-
pora Submerged trunk of Licuala longicalycata (peat bog) Thailand Pinruan etal. (2010)
Bisifusarium dimerum Groundwater Slovenia Novak Babič etal. (2016)
Calonectria candelabrum Submerged leaf Brazil da Silva and Gusmão (2015)
Chaetopsina aquatica Submerged decaying stem of Phragmites australis (river) Egypt Bakhit and Abdel-Aziz (2021)
Chaetopsina beijingensis Submerged decaying wood China Luo etal. (2019)
Chaetopsina fulva Submerged decaying leaves and petioles (stream), sub-
merged decaying test blocks (Dipterocarpus alatus and
Xylia dolabriformis)
Brazil, Thai-
land Barbosa etal. (2013b)
Chaetopsina hongkongensis Submerged decaying wood China Ho etal. (2002)
Chaetopsina illini Submerged decayed wood of Taxodium distichum (swamp) USA Crane (1971)
Chaetopsina penicillata Submerged decaying Xylia dolabriformis test block (stream) Thailand Sivichai etal. (2002)
Chaetopsina polyblastia Submerged decaying Xylia dolabriformis test block (stream) Thailand Sivichai etal. (2002)
Chaetopsina splendida Submerged decaying petioles (stream) Brazil Barbosa etal. (2013b)
Chaetopsinectria chaetopsinae Submerged decaying Xylia dolabriformis test block (stream) Thailand Sivichai etal. (2002)
Cosmospora aquatica Submerged decaying wood (stream) China Luo etal. (2019)
Cosmosporella olivacea Submerged decaying wood (lake) China Huang etal. (2018a)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Cylindrocarpon aquaticum Water (river), foam (stream), submerged decaying plant
materials (stream) India, Poland,
USA Ranzoni (1979), Czeczuga
etal. (1990), Rajashekhar and
Kaveriappa (2003)
Cylindrocarpon destructans
var. destructans Submerged leaves (river) UK Iqbal and Webster (1973)
Cylindrocladiella infestans Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2019)
Cylindrocladiella parva Submerged leaves Brazil da Silva and Gusmão (2015)
Fusarium solani Submerged decaying wood and bamboo (river, stream) Philippines,
USA Shearer and Von Bodman
(1983), Cai etal. (2003a)
Fusicolla aquaeductuum Submerged decaying wood (lake), water (river), submerged
decaying leaves of angiosperm, foam (river, lake, spring,
stream)
Brazil, China,
Poland, USA Crane (1968), Czeczuga etal.
(1990), Schoenlein-Crusius
etal. (2009), Huang etal.
(2018a)
Gliocladiopsis aquaticus Submerged decaying wood (lake) Thailand Hyde etal. (2018)
Gliocladiopsis guangdon-
gensis Submerged decaying wood (stream) China Liu and Cai (2013)
Gliocladiopsis singaporiensis Submerged decaying leaf (stream) Singapore Gordillo and Decock (2019)
Gliocladiopsis tenuis Submerged petioles India da Silva and Gusmão (2015)
Heliscus tentaculus Submerged decaying leaves of Platanus occidentalis, uni-
dentified leaves (stream) Jamaica, USA Umphlett (1957), Hudson and
Ingold (1960)
Ilyonectria destructans Foam (stream) India Rajashekhar and Kaveriappa
(2003)
Mariannaea aquaticola Submerged decaying wood (stream) Thailand Cai etal. (2010)
Mariannaea catenulata Submerged decaying wood (stream) China, Thai-
land Hao etal. (2021)
Mariannaea chlamydospora Submerged decaying wood (stream) China Hu etal. (2017)
Mariannaea cinerea Submerged decaying wood (reservoir) China Hu etal. (2017)
Mariannaea elegans Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2019)
Mariannaea fusiformis Submerged decaying wood (reservoir, stream) China, Korea Hu etal. (2017), Pangging etal.
(2021b)
Mariannaea lignicola Submerged decaying wood (stream) China Hu etal. (2017)
Mariannaea samuelsii Submerged decaying wood (stream) China Luo etal. (2019)
Mariannaea submersa Submerged decaying wood (river) China Hao etal. (2021)
Mariannaea superimposita Submerged decaying wood (river) China Luo etal. (2019)
Nectria curta Submerged wood of Fraxinus excelsior UK Webster (1993)
Nectriella lacustris Carex rostrata Austria Magnes and Hafellner (1991)
Nectria lugdunensis Submerged decaying wood (river) UK Shearer and Webster (1991)
Nectria submersa Water Japan Kobayasi and Ookubo (1952)
Neocosmospora haemato-
cocca Submerged decaying wood (river, stream) China Tsui etal. (2001d), Tsui and
Hyde (2004)
Neonectria coccinea Submerged beech (stream) Hungary Révay and Gönczöl (1990)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Neonectria lugdunensis Submerged decaying wood (Acer pseudoplatanus, Fraxinus
excelsior), rainwater from building roofs, water (river,
stream), foam (stream, rivulet, spring, river, lake), sub-
merged decaying leaves of Alnus glutinosa (stream), water
(stream)
Austria, Can-
ada, Czech
Republic,
France, Ger-
many, Ire-
land, Japan,
Poland,
Portugal,
Rhodesia,
Slovakia,
Spain, Swe-
den, UK,
USA
Ingold (1942), Fenton (1950),
Ranzoni (1953), Tubaki
(1957), Ingold (1958b), Nils-
son (1960), Baxter (1960),
Crane (1968), Iqbal and
Webster (1973), Bärlocher
(1987), Gessner etal. (1993),
Sridhar and Bärlocher (1993),
Czeczuga and Orłowska
(1997), Harrington (1997),
Marvanová (2000), Gulis and
Suberkropp (2003), Baschien
etal. (2006, 2008), Cornut
etal. (2014), Raposeiro etal.
(2018), Pietryczuk etal.
(2018)
Ophionectria trichospora Submerged decaying wood (stream) Mexico Castañeda-Ruíz etal. (2007)
Paracremonium binnewi-
jzendii Submerged decaying wood (lake) China Luo etal. (2019)
Payosphaeria minuta Submerged decaying wood (stream) China Cai etal. (2006)
Thelonectria discophora Submerged oak twigs (river) UK Shearer and Webster (1991)
Thelonectria lucida Submerged twigs (stream) USA Shearer and Von Bodman
(1983)
Thelonectria discophora Submerged decaying wood (stream) UK Shearer and Webster (1991)
Varicosporella aquatica Submerged decaying wood wood (Populus sp. Buxus sem-
pervirens, Salix sp., Alnus glutinosa, Fraxinus sp.) France Lechat and Fournier (2015)
Volutella citrinella Freshwater Korea Pangging etal. (2021b)
Niessliaceae
Paraniesslia aquatica Submerged decaying wood (stream) China Cai and Hyde (2007c)
Paraniesslia tuberculata Submerged decaying wood (river) China Tsui etal. (2001f)
Stachybotryaceae
Koorchalomella salmonispora Submerged decayed stem of Phragmites australis Egypt Hyde etal. (2017b)
Stachybotrys chartarum Submerged decaying wood (lake, river, stream) China Luo etal. (2019)
Stachybotrys chlorohalonatus Submerged decaying wood (river) China Luo etal. (2019)
Stachybotrys nephrosporus Submerged leaf Brazil da Silva and Gusmão (2015)
Stachybotrys sansevieriae Water (river) South Korea Lee etal. (2019)
Stachybotrys sphaerosporus Submerged decaying petioles (stream) Brazil Barbosa etal. (2013b)
Virgatospora echinofibrosa Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Sarocladiaceae
Sarocladium implicatum Water (river) Poland Pietryczuk etal. (2018)
Hypocreales genera incertae sedis
Idriella lunata Submerged decaying leaves (stream) Korea Goh etal. (2020)
Illosporiopsis christiansenii River watershed Canada Maloles etal. (2018)
Illosporium carneum River watershed Canada Maloles etal. (2018)
Microascales
Gondwanamycetaceae
Knoxdaviesia undulatistipes Submerged decaying petiole of Eleiodoxa conferta (peat
swamp) Thailand Pinnoi etal. (2003b, 2006)
Halosphaeriaceae
Aniptodera aquadulcis Submerged decaying wood (e.g., Dipterocarpus alatus
and Xylia dolabriformis test blocks) (lake, stream, river,
waterfall)
China, Thai-
land Hsieh etal. (1995), Hyde etal.
(1999a), Luo etal. (2004),
Kurniawati etal. (2010)
Aniptodera aquibella Submerged decaying wood (stream) Thailand Li etal. (2016)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Aniptodera chesapeakensis Submerged decaying wood (lake, stream, reservoir) Australia, Bra-
zil, Brunei,
China, India,
Iraq, Japan,
Mauritius,
South
Africa, USA
Shearer and Miller (1977),
Minoura and Muroi (1978),
Shearer and Crane (1986),
Shearer (1989b), Hyde
etal. (1998a), Goh and
Hyde (1999), Hyde etal.
(1999a), Fryar etal. (2004),
Luo etal. (2004), Patil and
Borse (2012b), Barbosa etal.
(2013a), Al-Saadoon and Al-
Dossary (2014)
Aniptodera fusiformis Submerged decaying cottonwood twigs (Populus deltoides) Iraq, USA Shearer (1989b), Al-Saadoon
and Al-Dossary (2014)
Aniptodera inflatiascigera Submerged decaying wood (river, stream) Brunei, China,
Philippines Tsui etal. (1997), Hyde etal.
(1999a), Ho etal. (2001)
Aniptodera lignicola Submerged decaying wood (lake, stream) and bamboo
(river) Australia,
Brunei,
China, India,
Malaysia,
Philippines
Hyde etal. (1999a), Cai etal.
(2003a), Luo etal. (2004),
Patil and Borse (2012b)
Aniptodera limnetica Submerged decaying wood (creek) USA Shearer (1989b)
Aniptodera margarition Submerged decaying wood UK, USA Shearer (1989b)
Aniptodera mauritaniensis Submerged decaying wood (river, lake) China, Iraq,
Mauritius Hyde etal. (1999a), Luo etal.
(2004), Al-Saadoon and Al-
Dossary (2014)
Aniptodera megaloascocarpa Submerged decaying wood (lake) USA Raja and Shearer (2008)
Aniptodera megalospora Submerged decaying wood (stream) Brunei, China,
Malaysia Hyde etal. (1999a)
Aniptodera palmicola Submerged decaying wood (lake), submerged Raphia aus-
tralia rachis China, Iraq,
South Africa Hyde etal. (1999a), Luo etal.
(2004), Al-Saadoon and Al-
Dossary (2014)
Aniptosporopsis lignatilis Submerged decaying wood (river, stream, waterfall) Australia,
China,
Mauritius
Philippines,
Seychelles,
South Africa
Hyde (1992a), Hyde etal.
(1999a), Ho etal. (2001)
Ascosacculus aquaticus Submerged decaying wood (river) Australia Hyde etal. (1999a)
Ascosacculus heteroguttulatus Submerged decaying wood (creek, river, stream) Australia, Bru-
nei, China,
Mauritius,
Philippines,
South Africa
Hyde etal. (1999a)
Ascosacculus fusiformis Submerged decaying wood (stream) Thailand Luo etal. (2019)
Cirrenalia macrocephala Submerged decaying wood (stream) China Ho etal. (2001)
Cucurbitinus constrictus Submerged decaying wood (lake) China Liu etal. (2020a)
Cucurbitinus ibericus Submerged wood Spain Hernández-Restrepo etal.
(2017)
Clavatospora filiformis Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (stream), foam (stream),
submerged decaying Eleiodoxa conferta, Licuala longi-
calycata
Metroxylon sagu (peat swamp)
Brunei, India,
Malaysia,
Thailand
Nawawi (1973a), Marvanová
(1980), Nawawi (1985b),
Sridhar and Kaveriappa
(1992), Kuthubutheen etal.
(1992), Sivichai etal. (2002),
Rajashekhar and Kaveriappa
(2003), Hosoya etal. (2006),
Pinnoi etal. (2006), Kurni-
awati etal. (2010), Yang etal.
(2018c), Hosoya etal. (2019)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Clavatospora longibrachiata Foam (stream, rivulet), submerged decaying leaves of Alnus
glutinosa (stream), water (river, stream), scum (stream) Austria, Can-
ada, France,
Ireland,
Poland, Por-
tugal, South
Africa, Swe-
den, Spain,
UK, USA
Ingold (1942), Ingold (1943a),
Willen (1958), Ingold (1960),
Greathead (1961), Iqbal and
Webster (1973), Bärlocher
(1987), Gessner etal. (1993),
Sridhar and Bärlocher (1993),
Voglmayr (1996), Harrington
(1997), Marvanová (2000),
Gulis and Suberkropp (2003),
Menéndez etal. (2012), Cor-
nut etal. (2014), Raposeiro
etal. (2018), Pietryczuk etal.
(2018)
Clavatospora tentacula Submerged leaves (Coffea arabica, Hevea brasiliensis,
unidentified) (stream), stemflow and throughfall of tree
canopies, foam (stream)
Cuba, India,
Malaysia,
Thailand,
USA, Ven-
ezuela
Marvanová and Marvan (1969),
Ranzoni (1979), Sridhar
(1987), Sridhar and Kave-
riappa (1988), Sridhar and
Kaveriappa (1992), Sivichai
and Boonyene (2004), Hosoya
etal. (2006), Smits etal.
(2007)
Fluviatispora reticulata Submerged Livistona sp. and Eleiodoxa conferta (peat
swamp) Papua New
Guinea,
Thailand
Hyde (1994a), Pinnoi etal.
(2006)
Fluviatispora tunicata Submerged Livistona sp. (river) Papua New
Guinea Hyde (1994a)
Haligena elaterophora Submerged decaying wood (river) UK Eaton and Jones (1971b)
Halosphaeria viscosa Submerged decaying wood (stream) Philippines,
South Africa Hyde etal. (1999a)
Halosphaeriopsis medioseti-
gera Submerged decaying balsa wood (river) USA Shearer and Crane (1971)
Luttrellia estuarina Submerged decorticated wood (river, stream) Canada, USA Shearer (1978), Ferrer and
Shearer (2007)
Luttrellia guttulata Submerged decorticated wood (stream) Costa Rica,
Panama Ferrer and Shearer (2007)
Luttrellia halonata Submerged decorticated wood (river) Ecuador Ferrer and Shearer (2007)
Luttrellia parvulospora Submerged decorticated wood (river) USA, French
Guiana Ferrer and Shearer (2007)
Magnisphaera stevemossago Submerged decorticated wood (creek, pond, river) USA Campbell etal. (2003a)
Nais aquatica Submerged decaying wood (river, stream, waterfalls) Australia,
China, South
Africa
Hyde etal. (1999a)
Nais inornata Submerged decaying wood of Pinus sylvestris and Fagus
sylvatica (water-cooling towers), wood (lake) China, Japan,
USA, UK Jones and Stewart (1972),
Minoura and Muroi (1978),
Luo etal. (2004), Raja etal.
(2009b)
Natantispora lotica Submerged decaying wood Australia,
China, Mau-
ritius, Philip-
pines, South
Africa,
USA
Hyde etal. (1999a), Campbell
etal. (2003a)
Natantispora retorquens Submerged decaying wood (river, reservoir, stream) and
wood (river), submerged rachis of Raphia australia Australia,
China, Mau-
ritius, Philip-
pines, South
Africa,
USA
Goh and Hyde (1999), Hyde
etal. (1999a), Cai etal.
(2002b), Cai etal. (2003a),
Campbell etal. (2003a)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Okeanomyces cucullatus Submerged decaying wood (lake) China Luo etal. (2004)
Phaeonectriella appendiculata Submerged decaying wood (stream) Australia,
Philippines Hyde etal. (1999a)
Phaeonectriella lignicola Submerged decaying wood (lake, river, stream, water-
cooling towers) China, Mauri-
tius, UK Eaton and Jones (1970), Hyde
etal. (1999a), Luo etal.
(2004)
Sigmoidea aurantiaca Foam (rivulet), submerged decaying wood (river) Austria, UK Descals and Webster (1982),
Voglmayr (1996)
Sigmoidea contorta Submerged decaying leaves (stream) Thailand Marvanová and Hywel-Jones
(2000)
Sigmoidea praelonga Foam (stream) Austria Marvanová (1986)
Sigmoidea prolifera Submerged decaying wood and leaves (stream), foam (lake,
stream, river, spring) USA Petersen (1963b), Crane (1968)
Microascaceae
Scedosporium cereisporum Fluid (wastewater treatment plant) France Crous etal. (2016a)
Scedosporium minutisporum Sediment (river) Spain Gilgado etal. (2005)
Triadelphiaceae
Triadelphia fusiformis Submerged decaying wood (stream) Thailand Lu etal. (2018b)
Triadelphia heterospora Submerged decaying wood (river) USA Shearer and Crane (1971)
Triadelphia hexaformispora Submerged decaying wood Thailand Chuaseeharonnachai etal.
(2020)
Triadelphia hungarica Water-filled treeholes Hungary Gönczöl and Révay (2003)
Triadelphia morgoensis Submerged decaying wood Hungary Révay (1992)
Microascales genera incertae sedis
Sporendocladia bactrospora Submerged decaying wood (stream), Submerged decaying
leaves of Calophyllum brasiliense (stream) Brazil Barbosa etal. (2013b), Fiuza
etal. (2018)
Torpedosporales
Juncigenaceae
Juncigena fruticosae Submerged decayed stems of Suaeda fruticosa (lake) Egypt Abdel-Wahab etal. (2010)
Vermiculariopsiellales
Vermiculariopsiellaceae
Vermiculariopsiella cornuta Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Vermiculariopsiella cubensis Submerged decaying leaves and petioles (stream) Brazil Barbosa etal. (2013b)
Vermiculariopsiella falcata Submerged decaying leaves and twigs (stream) Malaysia Nawawi etal. (1990)
Vermiculariopsiella immersa Submerged decaying leaves and twigs (stream), Submerged
decaying leaves of Calophyllum brasiliense (stream) Brazil, Malay-
sia Nawawi etal. (1990), Barbosa
etal. (2013b), Fiuza etal.
(2018)
Vermiculariopsiella parvula Submerged decaying twigs (stream) Malaysia Nawawi etal. (1990)
Hypocreomycetidae genera incertae sedis
Campylospora brasiliensis Submerged decaying leaves (stream, river, waterfall) Brazil Moro etal. (2015b)
Campylospora chaetocladia Submerged decaying leaves of Acer sp., Coffea arabica,
Hevea brasiliensis (stream, river), decaying unidentified
leaves (spring), foam, scum (lake, spring, stream, river),
stemflow and throughfall of tree canopies
Brazil, Cuba,
Ghana,
India,
Jamaica,
Malaysia,
Nigeria,
Portugal,
Thailand,
USA, Ven-
ezuela
Ranzoni (1953), Dixon (1959),
Hudson and Ingold (1960),
Crane (1968), Alasoadura
(1968a), Nawawi (1985b),
Sridhar and Kaveriappa
(1988), Rajashekhar and
Kaveriappa (2003), Sakayaroj
etal. (2005), Hosoya etal.
(2006), Smits etal. (2007),
Sridhar and Karamchand
(2009), Fiuza and Gusmão
(2013b), Raposeiro etal.
(2018)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Campylospora filicladia Submerged decaying leaves of Coffea arabica, and Hevea
brasiliensis (stream), foam (river, stream) Brazil, India,
Venezuela Sridhar and Kaveriappa (1988),
Rajashekhar and Kaveriappa
(2003), Smits etal. (2007),
Fiuza and Gusmão (2013b)
Campylospora parvula Foam (water, river, rivulet. stream), stemflow and through-
fall of tree canopies Austria, Bra-
zil, Canada,
India, Ire-
land, Japan,
Venezuela
Bärlocher (1987), Voglmayr
(1996), Harrington (1997),
Hosoya and Tanaka (2007),
Smits etal. (2007), Sridhar
and Karamchand (2009),
Fiuza and Gusmão (2013b)
Porosphaerellopsis bipolaris Submerged decaying wood (stream) China Ranghoo etal. (2001)
Lulworthiomycetidae
Pisorisporiales
Pisorisporiaceae
Achroceratosphaeria potamia Submerged wood of Platanus species (stream) France Réblová etal. (2010)
Pisorisporium cymbiforme Submerged decorticated wood of Alnus glutinosa and Fraxi-
nus excelsior (brook) France Réblová etal. (2015)
Savoryellomycetidae
Conioscyphales
Conioscyphaceae
Conioscypha aquatica Submerged decaying wood (river) China Luo etal. (2019)
Conioscypha gracilis Submerged decaying wood (stream) Japan, Peru Minoura and Muroi (1978),
Zelski etal. (2015)
Conioscypha lignicola Submerged decaying wood (stream) Australia,
China Höhnel (1904), Mongkol etal.
(2009)
Conioscypha nakagirii Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal.
(2017)
Conioscypha peruviana Submerged decaying wood (stream) Peru Zelski etal. (2015)
Conioscypha submersa Submerged decaying wood (river) China Luo etal. (2019)
Conioscypha tenebrosa Submerged decaying wood (river) China Yang etal. (2020)
Conioscypha verrucosa Submerged decaying wood (stream) China Hyde etal. (2020b)
Vanakripa ellipsoidea Submerged decaying wood (pond) China Tsui etal. (2003a)
Vanakripa fasciata Submerged decaying wood (river) Cuba Castañeda etal. (2005)
Vanakripa gigaspora Submerged decaying wood (river) Philippines Cai etal. (2003a)
Vanakripa menglensis Submerged decaying wood (stream) China Hu etal. (2010a)
Vanakripa minutiellipsoidea Submerged dead petiole of Eleiodoxa conferta (peat
swamp), submerged decaying wood Thailand Pinnoi etal. (2003b), Pinnoi
etal. (2006), Yang etal.
(2020)
Fuscosporellales
Fuscosporellaceae
Bactrodesmiastrum obovatum Submerged decaying wood (reservoir) Spain Calduch etal. (2002)
Fuscosporella aquatica Submerged decaying wood (stream) Thailand Yang etal. (2017)
Fuscosporella pyriformis Submerged decaying wood (stream) Thailand Yang etal. (2016b)
Mucispora infundibulata Submerged decaying wood (stream) Thailand Hyde etal. (2020d)
Mucispora obscuriseptata Submerged decaying wood (stream) Thailand Yang etal. (2016b)
Mucispora phangngaensis Submerged decaying wood (stream) Thailand Yang etal. (2017)
Parafuscosporella aquatica Submerged decaying wood (river) Thailand Yang etal. (2020)
Parafuscosporella ellipsocon-
idiogena Submerged decaying wood (waterfall) Thailand Boonyuen etal. (2021)
Parafuscosporella garethii Submerged decaying wood (stream) Thailand Boonyuen etal. (2016)
Parafuscosporella monili-
formis Submerged decaying wood (stream) Thailand Yang etal. (2016b)
Parafuscosporella mucosa Submerged decaying wood (stream) Thailand Yang etal. (2016b)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Parafuscosporella obovata Submerged decaying wood (waterfall) Thailand Boonyuen etal. (2021)
Parafuscosporella pyriformis Submerged decaying wood (river) Thailand Yang etal. (2020)
Plagiascoma frondosum Submerged decorticated wood of Fraxinus excelsior (brook) France Réblová etal. (2016b)
Pseudoascotaiwania persoonii Submerged decaying wood (river) Canada Fallah etal. (1999a)
Pleurotheciales
Pleurotheciaceae
Coleodictyospora muriformis Submerged decaying wood (stream) Thailand Dong etal. (2021b)
Neomonodictys muriformis Submerged decaying wood (stream) Thailand Hyde etal. (2020b)
Phaeoisaria aguilerae Submerged decaying twig (river) Cuba Castaneda Ruiz etal. (2002)
Phaeoisaria annesophieae Submerged decaying wood Thailand Shi etal. (2021)
Phaeoisaria aquatica Submerged decaying wood China Luo etal. (2018a)
Phaeoisaria clematidis Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (lake, reservoir, river,
stream), bamboo (river), submerged decaying Licuala
longicalycata and Eleiodoxa conferta (peat swamp)
Brazil, China,
Philippines,
Thailand
Goh and Hyde (1999), Cai etal.
(2003a), Luo etal. (2004),
Pinnoi etal. (2006), Pinruan
etal. (2007), Barbosa etal.
(2013b), Luo etal. (2018a),
Chethana etal. (2021)
Phaeoisaria guttulata Submerged decaying wood (river) China Hyde etal. (2018)
Phaeoisaria filiformis Submerged decaying wood (waterfall) Thailand Luo etal. (2019)
Phaeoisaria sparsa Submerged decaying wood (stream) Brunei,
Malaysia Ho etal. (2001), Hyde (2007)
Pleurotheciella aquatica Submerged decaying wood (river) China Luo etal. (2018a)
Pleurotheciella centenaria Submerged decaying wood (lake) Canada Réblová etal. (2012)
Pleurotheciella fusiformis Submerged decaying wood (lake) China Luo etal. (2018a)
Pleurotheciella guttulata Submerged decaying wood (river) China Luo etal. (2018a)
Pleurotheciella krabiensis Submerged decaying wood (stream) Thailand Hyde etal. (2018)
Pleurotheciella lunata Submerged decaying wood (river) China Luo etal. (2018a)
Pleurotheciella nilotica Submerged decaying wood (river) China, Egypt,
Thailand Abdel-Aziz etal. (2020), Shi
etal. (2021)
Pleurotheciella rivularia Submerged decaying wood of Hedera helix (rivulets) France Réblová etal. (2012)
Pleurotheciella saprophytica Submerged decaying wood (river) China Luo etal. (2018a)
Pleurotheciella submersa Submerged decaying wood (river) China Luo etal. (2018a)
Pleurotheciella sympodia Submerged decaying wood Thailand Shi etal. (2021)
Pleurotheciella tropica Submerged decaying wood (stream) Thailand Hyde etal. (2018)
Pleurotheciella uniseptata Submerged decaying wood (river) China Cai etal. (2002b); Luo etal.
(2018a)
Pleurothecium aquaticum Submerged decaying wood (stream) China Luo etal. (2018a)
Pleurothecium floriforme Submerged decaying wood (stream) Thailand Hyde etal. (2017b)
Pleurothecium guttulatum Submerged decaying wood China Shi etal. (2021)
Pleurothecium pulneyense Submerged decaying wood (river, stream) China, India Sridhar and Sudheep (2011),
Luo etal. (2018a)
Pleurothecium recurvatum Submerged decaying wood (stream) China Luo etal. (2019)
Pleurothecium semifecundum Submerged decaying wood of Sambucus nigra (brook) France Réblová etal. (2012)
Saprodesmium dematio-
sporum Submerged decaying wood (stream) China Dong etal. (2021b)
Sterigmatobotrys macrocar-
pus Submerged decaying wood (river, stream) Canada, UK Hyde and Goh (1999a), Réblová
and Seifert (2011)
Sterigmatobotrys rudis Submerged decaying wood (water-cooling towers, lake) China, UK Jones etal. (2002), Luo etal.
(2004)
Sterigmatobotrys uniseptatus Submerged decaying wood (stream) China Chang (1991), Luo etal. (2019)
Pseudodactylariales
Pseudodactylariaceae
Pseudodactylaria aquatica Submerged decaying wood (stream) Thailand Bao etal. (2021a)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Pseudodactylaria campore-
siana Submerged decaying wood (stream) Thailand Hyde etal. (2020b)
Pseudodactylaria fusiformis Submerged decaying wood (stream) China Lu etal. (2020)
Pseudodactylaria hyalotuni-
cata Submerged decaying wood (stream, river) Brazil, China Tsui etal. (2000)
Savoryellales
Savoryellaceae
Ascotaiwania fusiformis Submerged decaying wood (stream) Thailand Yang etal. (2016b)
Ascotaiwania hsilio Submerged decaying wood (stream) China, USA Chang etal. (1998), Fallah etal.
(1999a), Raja etal. (2009b)
Ascotaiwania limnetica Submerged decaying wood (stream) China, France,
India Chang etal. (1998), Van
Vooren (2010), Patil and
Borse (2012a), Réblová etal.
(2016b)
Ascotaiwania mitriformis Submerged decaying wood (reservoir) China Ranghoo and Hyde (1998)
Ascotaiwania pallida Submerged decaying wood (river, stream) Australia,
Brunei, UK Hyde and Goh (1999a), Fryar
etal. (2004), Vijaykrishna and
Hyde (2006)
Ascotaiwania palmicola Submerged decaying wood (lake, stream) Australia,
China Hyde and Goh (1997), Luo
etal. (2004)
Ascotaiwania sawadae Submerged decaying wood (stream) China, Thai-
land Chang etal. (1998), Sivichai
(1998)
Ascotaiwania wulai Submerged decaying wood (stream) China Chang etal. (1998)
Canalisporium aquaticium Submerged decaying wood (stream) Thailand Hyde etal. (2020b)
Canalisporium caribense Submerged decaying wood (river, stream, waterfall), rain-
water from building roofs, decaying Licuala longicalycata
(peat swamp)
Brazil, Brunei,
China,
India, North
Maharashtra,
Malaysia,
Panama,
Poland,
Thailand,
USA
Goh etal. (1998a), Ho etal.
(2001), Raja etal. (2009b),
Sridhar etal. (2012), Pinruan
etal. (2014), Zhang etal.
(2014b), Patil etal. (2014)
Canalisporium elegans Submerged decaying wood (river, stream) Brunei,
China,
Malaysia,
Panama,
Thailand
Nawawi and Kuthubutheen
(1989a), Goh etal. (1998a),
Ferrer and Shearer (2005),
Zhang etal. (2014b), Goh and
Kuo (2021)
Canalisporium exiguum Submerged decaying wood (stream) Australia,
Brazil Goh etal. (1998a), Barbosa
etal. (2013b)
Canalisporium grenadoideum Submerged wood of Wrightia tomentosa (stream), unidenti-
fied wood (river) Egypt, Thai-
land Sri-indrasutdhi etal. (2010),
Bakhit (2021)
Canalisporium jinghongense Submerged decaying wood (river, stream, waterfall) China, Egypt,
Thailand Cai etal. (2003c), Zhang etal.
(2014b)
Canalisporium kenyense Submerged decaying wood (stream) Panama, USA Ferrer and Shearer (2005), Raja
etal. (2009b)
Canalisporium macrosporum Submerged decaying culm of Miscanthus floridulus (stream) China Goh and Kuo (2021)
Canalisporium nanhuaense Submerged decaying wood China Goh and Kuo (2021)
Canalisporium pallidum Submerged decaying wood China Goh etal. (1998a)
Canalisporium panamense Submerged decaying wood Panama Ferrer and Shearer (2005)
Canalisporium parvum Submerged decaying culm of Miscanthus floridulus (stream) China Goh and Kuo (2021)
Canalisporium paulopallidum Submerged decaying wood China Goh and Kuo (2021)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Canalisporium pulchrum Submerged decaying wood (reservoir, stream) Australia,
Brunei, China,
Malaysia,
Panama,
Thailand
Nawawi and Kuthubutheen
(1989a), Goh etal. (1998a),
Goh and Hyde (1999), Ferrer
and Shearer (2005), Zhang
etal. (2014b), Chethana etal.
(2021)
Canalisporium taiwanense Submerged decaying wood China Goh and Kuo (2021)
Canalisporium variabile Submerged decaying wood (lake), submerged decaying
Licuala longicalycata (peat swamp) Australia,
China, Thai-
land
Goh and Hyde (2000b), Luo
etal. (2004), Pinruan etal.
(2007)
Canalisporium waffleum Submerged decaying culm of Miscanthus floridulus (stream) China Goh and Kuo (2021)
Dematiosporium aquaticum Submerged decaying wood (lake) China Luo etal. (2019)
Helicoascotaiwania hughesii Submerged decaying wood (lake) USA Fallah etal. (1999a)
Savoryella aquatica Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) (creek, lake, river, stream)
and bamboo (river)
Australia, Bru-
nei, China,
Malaysia,
Philip-
pines, South
Africa,
Thailand
(Hyde (1993b), Ho etal.
(1997), Goh and Hyde (1999),
Cai etal. (2003a), Luo etal.
(2004)
Savoryella curvispora Submerged decaying wood (lake, river, stream) China,
Malaysia,
Mauritius,
Philippines,
South Africa
Ho etal. (1997), Luo etal.
(2004), Hyde (2007)
Savoryella fusiformis Submerged decaying wood Brunei Ho etal. (1997)
Savoryella grandispora Submerged decaying wood (river) Australia,
Malaysia Hyde (1994a), Ho etal. (1997)
Savoryella lignicola Submerged decaying wood (lake, river, stream, waterfall),
submerged pine test blocks (water-cooling towers) Australia, Bru-
nei, China,
Japan,
Malaysia,
South
Africa, Thai-
land, UK
Jones and Eaton (1969), Hyde
(1993b), Hyde and Goh
(1997), Ho etal. (1997),
Tsui etal. (2003c), Luo etal.
(2004), Kurniawati etal.
(2010)
Savoryella verrucosa Submerged decaying wood (e.g., Dipterocarpus alatus and
Xylia dolabriformis test blocks) Australia, Bru-
nei, Japan,
Malaysia,
Thailand
Ho etal. (1997), Sivichai etal.
(2002)
Rhexoacrodictys erecta Submerged decaying wood (lake, river, stream, waterfall)
and leaves (stream) Brazil, China,
Thailand Luo etal. (2004), Kurniawati
etal. (2010), Barbosa etal.
(2013b)
Savoryellomycetidae genera incertae sedis
Obliquifusoideum guttulatum Submerged decaying wood (stream) China Dong etal. (2021b)
Sordariomycetidae
Chaetosphaeriales
Chaetosphaeriaceae
Anacraspedodidymum sub-
mersum Submerged decaying dicotyledonous leaves (stream) China Zheng etal. (2021)
Arcuatospora novae-zelandiae Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Bahusutrabeeja dwaya Submerged decaying wood (river, stream) China, Malay-
sia Tsui etal. (2001b)
Brunneodinemasporium
jonesii Submerged decaying wood (stream) China Lu etal. (2016)
Catenularia angulospora Submerged decaying wood (stream) Brunei, China Ho etal. (2001)
Catenularia catenulata Submerged decaying wood China Luo etal. (2019)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Chaetosphaeria anglica Submerged decaying wood (stream) Brunei, Sudan,
UK Fisher and Petrini (1983), Ho
etal. (2001)
Chaetosphaeria aquatica Submerged decaying wood (river) China Luo etal. (2019)
Chaetosphaeria cubensis Submerged decaying wood (river) China Luo etal. (2019)
Chaetosphaeria guttulata Submerged decaying wood (stream) China Luo etal. (2019)
Chaetosphaeria hiugensis Decaying Nenga pumila (peat swamp) Thailand Pinruan etal. (2014)
Chaetosphaeria innumera Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Chaetosphaeria lentomita Submerged decaying wood (stream) China Ho etal. (2001)
Chaetosphaeria lignomollis Submerged decaying twig (stream) Brazil Barbosa etal. (2013a)
Chaetosphaeria myriocarpa Submerged decaying wood (river) China Luo etal. (2019)
Chaetosphaeria pulchriseta Submerged decaying wood and twig (stream) Brazil, Malay-
sia Kuthubutheen and Nawawi
(1991a), Barbosa etal.
(2013b)
Chaetosphaeria rivularis Submerged wood of Fagus sylvatica associated with Minuti-
sphaera japonica (brook) France Ariyawansa etal. (2015)
Chaetosphaeria submersa Submerged decaying wood (stream) China Luo etal. (2019)
Chaetosphaeria vermiculari-
oides Stemflow and throughfall of palm species India Ghate and Sridhar (2015)
Chloridium aquaticum Submerged decaying wood Thailand Wei etal. (2018)
Chloridium aseptatum Submerged decaying wood Thailand Wei etal. (2018)
Chloridium chloroconium Submerged decaying wood and leaves (stream) Brazil, China Ho etal. (2001), Barbosa etal.
(2013b)
Chloridium cylindrosporum Submerged decaying wood (river) Philippines Cai etal. (2003a)
Chloridium gonytrichii Submerged decaying wood (stream) China, Thai-
land Luo etal. (2019), Bao etal.
(2021a)
Chloridium lignicola Submerged decaying wood (stream) Brunei, China,
Malaysia Ho etal. (2001, 2002), Hyde
2007)
Chloridium matsushimae Submerged decaying wood (stream) China Tsui etal. (2000)
Chloridium obclaviforme Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Chloridium pachytrachelum Submerged decaying wood (stream) China Ho etal. (2002)
Chloridium submersum Submerged decaying wood (river) China Hyde etal. (2020d)
Chloridium transvaalense Submerged leaf Brazil da Silva and Gusmão (2015)
Chloridium virescens Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Codinaea aquatica Submerged decaying wood Brazil Oliveira etal. (2015c)
Codinaea lunulospora Submerged decaying twig (stream) Malaysia Kuthubutheen and Nawawi
(1991a)
Codinaea yunnanensis Submerged decaying wood (river) China Luo etal. (2019)
Craspedodidymum cubense Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Cryptophiale cucullata Submerged decaying leaves (stream) Brazil Kuthubutheen (1987b)
Cryptophiale kakombensis Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Cryptophiale manifesta Submerge decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Cryptophiale multiseptata Submerged decaying wood (river) Australia Goh and Hyde (1996d)
Cryptophiale udagawae Submerged decaying wood (e.g. Dipterocarpus alatus test
block) and leaves (stream) Brazil, Thai-
land Sivichai etal. (2002), Barbosa
etal. (2013b)
Cryptophialoidea fasciculata Submerged decaying twigs and petioles (stream) Brazil, Malay-
sia Kuthubutheen and Nawawi
(1994a), Barbosa etal.
(2013b)
Cryptophialoidea secunda Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Cryptophialoidea uncispora Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1987b)
Dictyochaeta aciculata Submerged petiole Brazil Silva and Gusmão (2013)
Dictyochaeta aquatica Submerged decaying wood Thailand Wei etal. (2018)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Dictyochaeta assamica Submerged decaying wood (reservoir, river, stream) China, Malay-
sia, UK Kuthubutheen and Nawawi
(1991a), Goh and Hyde
(1999), Hyde and Goh
(1999a)
Dictyochaeta britannica Submerged decaying petiole (stream) Brazil Barbosa etal. (2013b)
Dictyochaeta cangshanensis Submerged decaying wood (stream) China Luo etal. (2019)
Dictyochaeta coffeae Submerged decaying wood (river) China Tsui etal. (2001d)
Dictyochaeta curvispora Submerged bamboo (river) Philippines Cai etal. (2003a), Cai etal.
(2004b)
Dictyochaeta daphnioides Submerged decaying angiospermous petioles Malaysia Kuthubutheen and Nawawi
(1991b)
Dictyochaeta ellipsoidea Submerged decaying wood China Luo etal. (2019)
Dictyochaeta fertilis Submerged decaying leaves of unidentified dicotyledon plant
(stream), wood (stream, reservoir), Submerged decaying
leaves of Calophyllum brasiliense (stream)
Brazil, China,
Malaysia Kuthubutheen and Nawawi
(1991a), Barbosa etal.
(2013b), Fiuza etal. (2019)
Dictyochaeta gamundiae Decaying bark (river) Argentina Arambarri etal. (1987)
Dictyochaeta guadalcanalensis Submerged decaying twig of unidentified angiosperm Malaysia Kuthubutheen and Nawawi
(1991a)
Dictyochaeta gyrosetula Submerged decaying wood (e.g., Xylia dolabriformis test
block) (stream) Malaysia,
Thailand Kuthubutheen and Nawawi
(1991c), Sivichai etal. (2002)
Dictyochaeta heteroderae Submerged decaying wood (stream), and leaves of Calophyl-
lum brasiliense (stream) Brazil Barbosa etal. (2013b), Fiuza
etal. (2019)
Dictyochaeta lignicola Submerged decaying wood (river) China Luo etal. (2019)
Dictyochaeta longispora Submerged decaying leaves of unidentified dicotyledon
plant (stream) Malaysia Kuthubutheen and Nawawi
(1991a)
Dictyochaeta lunulospora Submerged decaying twig of unidentified dicotyledon plant Malaysia Kuthubutheen and Nawawi
(1991a)
Dictyochaeta malaysiana Submerged decaying leaves Malaysia Kuthubutheen (1987c)
Dictyochaeta microcylindros-
pora Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Dictyochaeta plovercovensis Submerged decaying wood and bamboo (lake, reservoir) China, Philip-
pines Goh and Hyde (1999) Cai etal.
(2003c), Luo etal. (2004)
Dictyochaeta pluriguttulata Submerged decaying wood (stream) Brazil, Malay-
sia Kuthubutheen and Nawawi
(1991c), Barbosa etal.
(2013b)
Dictyochaeta subfuscospora Submerged decaying wood Australia,
Malaysia Kuthubutheen and Nawawi
(1991c), Hyde and Goh
(1997)
Dictyochaeta ramulosetula Submerged decaying leaves, decaying Metroxylon sagu (peat
swamp) Malaysia,
Thailand Kuthubutheen (1987c), Pinruan
etal. (2014)
Dictyochaeta sampahia Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1991c)
Dictyochaeta simplex Submerged decaying wood and leaves (stream) and leaves
(stream), Submerged decaying leaves of Calophyllum
brasiliense (stream)
Brazil, Malay-
sia Kuthubutheen and Nawawi
(1991a), Barbosa etal.
(2013b), Fiuza etal. (2019)
Dictyochaeta subfuscospora Submerged decaying wood (stream) Australia,
Malaysia Kuthubutheen and Nawawi
(1991c), Hyde and Goh
(1997)
Dictyochaeta siamensis Submerged decaying wood (stream) Thailand Liu etal. (2016)
Dictyochaeta submersa Submerged decaying wood (stream) China Luo etal. (2019)
Dictyochaeta tumidoseta Submerged decaying culm of Miscanthus floridulus (stream) China Kuo and Goh (2019)
Dictyochaeta vittata Submerged decaying wood Malaysia Kuthubutheen and Nawawi
(1991c)
Dictyochaetopsis hamata Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1990)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Dictyochaetopsis gony-
trichodes Submerged decaying wood (stream, river) and leaves
(stream) Brazil, Malay-
sia, USA Shearer and Crane (1971),
Kuthubutheen and Nawawi
(1990), Barbosa etal. (2013b)
Dictyochaetopsis pahangensis Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1990)
Dictyochaetopsis polysetosa Submerged decaying leaves and petioles (stream) Brazil Barbosa etal. (2013b)
Dinemasporium lanatum Submerged decaying leaves and petiole (stream), decaying
Metroxylon sagu (peat swamp) Brazil, Thai-
land Barbosa etal. (2013b), Pinruan
etal. (2014)
Exserticlava aquatica Submerged decaying wood Brazil do Carmo etal. (2020)
Exserticlava globosa Submerged decaying wood (river) China Tsui etal. (2001a)
Exserticlava triseptata Submerged decaying wood (river, stream), submerged
decaying culm of Miscanthus floridulus (stream) Brazil, China,
Malaysia,
Seychelles
Kuthubutheen and Nawawi
(1994b), Hyde and Goh
(1998c), Tsui etal. (2001a),
Barbosa etal. (2013b), Kuo
and Goh (2019)
Exserticlava vasiformis Submerged decaying wood (river, stream) China, Malay-
sia, USA Schoknecht and Crane (1983),
Kuthubutheen and Nawawi
(1994b), Tsui etal. (2001a)
Exserticlava yunnanensis Submerged decaying wood (stream) China Cai and Hyde (2007a), Barbosa
etal. (2013b)
Gonytrichum macrocladum Submerged decaying wood (reservoir, stream, waterfall),
submerged decaying Eleiodoxa conferta (peat swamp) China, Thai-
land Goh and Hyde (1999), Pinnoi
etal. (2006), Kurniawati etal.
(2010)
Kionochaeta australiensis Submerged decaying wood (stream) Australia Goh and Hyde (1997a)
Kionochaeta pughii Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1988a)
Kionochaeta ramifera Submerged decaying leaves Malaysia Kuthubutheen and Nawawi
(1988a)
Kionochaeta setosimplicia Submerged decaying wood (rivulet) Thailand Nuankaew etal. (2021)
Kionochaeta spissa Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2019)
Miyoshiella triseptata Submerged decaying wood (lake, reservoir, stream) and
bamboo (river) Brazil, China,
Philippines Goh and Hyde (1999) Cai etal.
(2003a), Luo etal. (2004),
Barbosa etal. (2013b)
Menisporopsis multisetulata Submerged decaying wood (river) China Tsui etal. (1999a)
Menisporopsis pirozynskii Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Menisporopsis theobromae Submerged decaying wood Thailand Liu etal. (2016)
Nawawia oviformis Submerged leaves of an unidentified dicotyledonous plant
(stream) China Peng etal. (2016)
Nawawia quadrisetulata Submerged decaying wood Malaysia Goh etal. (2014)
Paliphora multiseptata Submerged leaf Brazil da Silva and Gusmão (2015)
Paliphora porosa Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2019)
Phaeonawawia diplocladiel-
loidea Submerged decaying wood Malaysia Goh etal. (2021b)
Phialogeniculata africana Submerged decaying wood (river) South Africa Hyde etal. (1998a)
Phialosporostilbe dendroidea Submerged pieces of stems of Phragmites, submerged
decaying culm of Miscanthus floridulus (stream) China, South
Africa Hyde etal. (1996a), Kuo and
Goh (2019)
Rattania aquatica Submerged decaying wood (river) Thailand Calabon etal. (2021a)
Sporoschisma australiense Submerged decaying wood (stream) Australia Goh etal. (1997b)
Sporoschisma chiangraiense Submerged decaying wood Thailand Bao etal. (2021a)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Sporoschisma hemipsilum Submerged decaying wood (e.g., Dipterocarpus alatus test
block) and herbaceous debris (lake, stream) Australia,
Brazil, Bru-
nei, China,
Malaysia,
Perú, South
Africa,
Thailand
Goh etal. (1997b), Sivichai
etal. (2000a, 2002), Luo
etal. (2004), Barbosa etal.
(2013b), Zelski etal. (2014),
Yang etal. (2016a), Luo etal.
(2016b)
Sporoschisma juvenile Submerged decaying wood (stream, river) and bamboo
(river) Brazil, China,
Perú,
Philippines,
Seychelles
Goh etal. (1997b), Cai etal.
(2003a) Barbosa etal.
(2013b), Zelski etal. (2014),
Luo etal. (2016b)
Sporoschisma longicatenatum Submerged decaying wood (stream) Thailand Yang etal. (2016a)
Sporoschisma mirabile Submerged decaying wood (river) China Luo etal. (2016b)
Sporoschisma nigroseptatum Submerged decaying wood (river, stream) China, India Sudheep and Sridhar (2011),
Luo etal. (2016b)
Sporoschisma palauense Submerged decaying wood (stream) Thailand Yang etal. (2016a)
Sporoschisma parcicuneatum Submerged decaying wood (stream) Philippines Goh etal. (1997b)
Sporoschisma phaeocentron Submerged decaying wood China, Malay-
sia Goh etal. (1997b), Ho etal.
(2001), Luo etal. (2016b)
Sporoschisma taitense Submerged decaying wood China Luo etal. (2016b)
Sporoschisma uniseptatum Submerged decaying wood (e.g., Dipterocarpus alatus test
block) (stream, reservoir) Brunei, China,
India,
Malay-
sia, Perú,
Seychelles,
Thailand
Goh and Hyde (1999), Ho etal.
(2001), Sivichai etal. (2002),
Sudheep and Sridhar (2011),
Zelski etal. (2014)
Tainosphaeria aquatica Submerged decaying wood (stream) Thailand Li etal. (2021a)
Tainosphaeria jonesii Submerged decaying wood (stream) China Lu etal. (2016)
Tainosphaeria siamensis Submerged decaying wood Thailand Liu etal. (2016)
Tainosphaeria lunata Submerged decaying wood (river) China Luo etal. (2019)
Tainosphaeria obclavata Submerged decaying wood (stream) Thailand Luo etal. (2019)
Tainosphaeria thailandensis Submerged decaying wood (stream) Thailand Li etal. (2021a)
Thozetella acerosa Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2018)
Thozetella canadensis Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2018)
Thozetella coronata Submerged decaying wood (stream) Brazil Monteiro etal. (2016a)
Thozetella cristata Submerged decaying wood and leaves (stream), submerged
decaying leaves of Calophyllum brasiliense (stream) Brazil Barbosa etal. (2013b), Fiuza
etal. (2018)
Thozetella cubensis Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Thozetella gigantea Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Thozetella nivea Submerged decaying wood (e.g., Dipterocarpus alatus
and Xylia dolabriformis test blocks), decaying Eleiodoxa
conferta, Licuala longicalycata (peat swamp)
Thailand Sivichai etal. (2002), Pinruan
etal. (2014)
Thozetella pinicola Submerged decaying leaves (stream), submerged decaying
leaves of Calophyllum brasiliense (stream) Brazil Barbosa etal. (2013b), Fiuza
etal. (2018)
Thozetella queenslandica Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2018)
Thozetella radicata Submerged decaying Licuala longicalycata (peat swamp),
submerged decaying leaves of Calophyllum brasiliense
(stream)
Brazil, Thai-
land Pinruan etal. (2007), Fiuza
etal. (2018)
Thozetella submersa Submerged decaying wood (stream) Brazil Barbosa etal. (2011), Barbosa
etal. (2013b)
Thozetella ypsiloidea Submerged decaying wood (stream) Brazil Monteiro etal. (2016a)
Zanclospora novae-zelandiae Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Helminthosphaeriaceae
Diplococcium aquaticum Submerged decaying wood (lake) Australia Goh etal. (1998d)
Diplococcium asperum Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Endophragmiella boothii Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Endophragmiella mexicana Submerged decaying wood (stream) China Ho etal. (2002)
Endophragmiella occidentalis Submerged decaying wood (stream) China Ho etal. (2002)
Endophragmiella profusa Submerged decaying decaying leaves of Bucida palustris
(pool) Cuba Castañeda Ruiz etal. (2010)
Endophragmiella ramificata Submerged decaying wood (stream) China Ho etal. (2002)
Endophragmiella rostrata Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Endophragmiella triseptata Submerged decaying wood (river) China Tsui etal. (2001c)
Hilberina breviseta Submerged decaying wood (lake, stream) Brunei, China Ho etal. (2001), Luo etal.
(2004)
Linocarpaceae
Linocarpon apiculatum Submerged decaying palm petiole (swamp) Indonesia Hyde (1997)
Linocarpon appendisporum Submerged decaying Pandanus leaves (swamp) Indonesia Hyde (1997)
Linocarpon bambusicola Submerged bamboo (river) Philippines Cai etal. (2004b)
Linocarpon elaeidis Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Linocarpon livistonae Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Linocarpon pandani Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Linocarpon pandanicola Submerged decaying Pandanus leaves Indonesia Hyde (1997)
Neolinocarpon inconspicuum Submerged decaying leaves (stream) China Ho etal. (2001)
Coniochaetales
Coniochaetaceae
Coniochaeta gigantospora Submerged decaying wood (river) France Raja etal. (2012a)
Coniochaeta hoffmannii Polluted water UK Weber etal. (2002)
Coniochaeta kellermanii Submerged decaying wood (river) USA Shearer (1972), Shearer and
Crane (1986)
Coniochaeta leucoplaca Submerged decaying wood (pond) USA Shearer and Crane (1986)
Coniochaeta ligniaria Submerged decaying wood (cooling tower) UK Eaton (1972)
Coniochaeta mutabilis Water (river) Germany Perdomo etal. (2011)
Coniochaeta renispora Submerged decaying wood (river) USA Crane and Shearer (1995)
Coniochaeta velutina Submerged decaying wood USA Shearer and Crane (1986)
Cordanales
Cordanaceae
Cordana abramovii Submerged decaying wood and herbaceous debris, bamboo
(river) Brunei, Perú,
Philippines
Thailand
Ho etal. (2001), Cai etal.
(2003a), Zelski etal. (2014),
Luo etal. (2019)
Cordana aquatica Submerged decaying wood (river) China Luo etal. (2019)
Cordana lignicola Submerged decaying wood (river) China Luo etal. (2019)
Cordana terrestris Submerged decaying wood and bamboo (river, stream) Brazil, China Cai and Hyde (2007a), Luo
etal. (2019)
Cordana uniseptata Submerged decaying Phyllostachys bambusoides (stream) China Cai etal. (2004a)
Phyllachorales
Phyllachoraceae
Ascovaginospora stellipala Submerged dead stems and leaves of Carex limosa USA Fallah etal. (1997)
Phyllachora therophila Juncus filiformis Austria Magnes and Hafellner (1991)
Tamsiniella labiosa Submerged decaying wood Australia Wong etal. (1998b)
Phyllachorales genera incertae sedis
Neoxylomyces multiseptatus Submerged decaying wood (stream) Thailand Calabon etal. (2021a)
Sordariales

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Chaetomiaceae
Arcopilus aureus Submerged decaying wood USA Shearer and Crane (1986)
Chaetomium cochliodes Phragmites australis, balsa wood UK, USA Apinis etal. (1972b)
Chaetomium elatum Beech wood UK Eaton (1972)
Chaetomium globosum Submerged decaying balsa wood, submerged dead stems of
Carex oligosperma (bog, lake) China, USA,
UK Eaton (1972), Fallah and
Shearer (2001), Luo etal.
(2019)
Chaetomium trilaterale Balsa wood USA Shearer and Crane (1986)
Corynascella inquinata sewage sludge Japan Udagawa and Ueda (1979)
Collariella carteri Water (pond, stream) Korea Nguyen etal. (2019a)
Dichotomopilus funicola Wood USA Shearer and Crane (1986)
Collariella carteri Water (pond, stream) Korea Nguyen etal. (2019a)
Dichotomopilus funicola Wood USA Shearer and Crane (1986)
Humicola asteroidea Submerged decaying wood (stream) USA Raja etal. (2009b)
Humicola homopilata Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Humicola seminuda Submerged leaf litter USA Shearer and Crane (1986)
Staphylotrichum longicolle Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Trichocladium englandense Submerged decaying wood (river) UK Hyde and Goh (1999a)
Trichocladium lignicola Submerged decaying wood (lake, river) China, China Cai etal. (2002b), Kane etal.
(2002)
Trichocladium linderi Submerged decaying wood (river) USA Crane and Shearer (1978)
Trichocladium lobatum Submerged decaying wood (swamp) USA Crane and Shearer (1978)
Trichocladium moenitum Submerged decaying wood (lake, swamp) USA Crane and Shearer (1978)
Lasiosphaeriaceae
Apiosordaria hamata Sediment (lake) China Wu etal. (2016)
Arnium apiculatum Equisetum fluviatile USA Shearer etal. (1980)
Bombardia arachnoidea Balsa wood USA Shearer (1972)
Cercophora appalachianensis Submerged decaying wood (stream) China Ho etal. (2001)
Cercophora aquatica Submerged decaying wood of Quercus (stream) France Chaudhary etal. (2007)
Cercophora caudata Submerged decaying wood (river) China Luo etal. (2019)
Cercophora costaricensis Submerged decaying wood (stream) China Tsui etal. (2000)
Cercophora squamulosa Submerged decaying wood (brook, stream) France Crous etal. (2016a)
Cercophora sulphurella Submerged decaying wood (lake) USA Fallah and Shearer (2001)
Cercophora vinosa Submerged decaying wood (stream) France Crous etal. (2016b)
Cladorrhinum australe Submerged decaying stem Korea Nam etal. (2020)
Cladorrhinum foecundissimum Submerged decaying wood (river) UK Hyde and Goh (1999a)
Cladorrhinum samala Submerged decaying wood (stream) Brunei Ho etal. (2001)
Echria gigantospora Submerged decaying wood (lake) USA Kruys etal. (2015)
Immersiella immersa Submerged decaying wood (stream) China Tsui etal. (2000)
Zopfiella cephalothecoidea Submerged dead twig Iraq Guarro etal. (1996)
Zopfiella latipes Submerged decaying wood (lake) China, Egypt Cai etal. (2002b), Luo etal.
(2004), Abdel-Aziz (2016b)
Zopfiella leucotricha Submerged decaying wood (river) Egypt Abdel-Aziz (2016b)
Zopfiella lundqvistii Submerged decaying wood (Ochroma pyramidale) (swamp) USA Shearer and Crane (1978)
Zopfiella submersa Submerged dead culms of Phragmites sp. (river) Iraq Guarro etal. (1997)
Podosporaceae
Triangularia setosa Submerged beech wood UK Eaton and Jones (1971b)
Sordariaceae
Neurospora retispora Submerged decaying wood (lake) China Luo etal. (2004)
Sordariales genera incertae sedis
Ascolacicola aquatica Submerged decaying wood (lake) China Ranghoo and Hyde (1998)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Cuspidatispora xiphiago Submerged decaying wood (river) China Luo etal. (2019)
Lockerbia striata Submerged decaying wood (lake, river) USA Raja and Shearer (2008)
Lunulospora curvula Foam (lake, spring, river, stream), submerged decaying
leaves of Alnus glutinosa and Salix, submerged leaves of
Coffea arabica and Hevea brasiliensis (river, stream), rain-
water from building roofs, water (river, stream), sediment
(dam, stream), stemflow and throughfall of tree canopies
Australia,
Austria,
Brazil, Cuba,
France,
India,
Jamaica,
Malaysia,
Nigeria,
Poland,
Portugal,
Spain, UK,
USA, Ven-
ezuela
Ingold (1942), Ingold (1943a),
Ranzoni (1953), Hudson
and Ingold (1960), Crane
(1968), Alasoadura (1968a),
Marvanová and Marvan
(1969), Ingold and Webster
(1973), Iqbal and Webster
(1973), Sridhar and Kaveri-
appa (1988), Czeczuga and
Orłowska (1997), Marvanová
(2000), Gulis and Suberkropp
(2003), Rajashekhar and
Kaveriappa (2003), Hosoya
etal. (2006), Smits etal.
(2007), Schoenlein-Crusius
etal. (2009), Sridhar and Kar-
amchand (2009), Duarte etal.
(2012), Sudheep and Sridhar
(2012), Menéndez etal.
(2012), Cornut etal. (2014),
Raposeiro etal. (2018),
Pietryczuk etal. (2018)
Lunulospora cymbiformis Submerged decaying leaves of dicotyledon (river), sub-
merged leaves of Coffea arabica and Hevea brasiliensis
(stream), sediment (dam, stream), foam (stream, river)
Australia,
Brazil,
India, Japan,
Malaysia,
Thailand
Miura (1972b), Sridhar and
Kaveriappa (1988), Sridhar
and Kaveriappa (1992), Saka-
yaroj etal. (2005), Hosoya
etal. (2006), Hosoya and
Tanaka (2007), Schoenlein-
Crusius etal. (2009), Sudheep
and Sridhar (2012), Duarte
etal. (2015)
Sordariomycetidae genera incertae sedis
Hanliniomyces hyaloapicalis Submerged decaying wood USA Raja and Shearer (2008)
Hydromelitis pulchella Submerged decaying wood Costa Rica Ferrer etal. (2012)
Rhexodenticula aquatica Submerged decaying wood (stream) Thailand Dong etal. (2021c)
Xylariomycetidae
Amphisphaeriales
Amphisphaeriaceae
Amphisphaeria uniseptata Submerged decaying wood (river) China Tsui etal. (2001f)
Capsulospora frondicola Submerged decaying Eleiodoxa conferta and Metroxylon
sagu (peat swamp) Thailand Pinnoi etal. (2006), Pinruan
etal. (2014)
Lepteutypa aquatica Submerged decaying wood (stream) Thailand Luo etal. (2019)
Pestalosphaeria austroameri-
cana Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Apiosporaceae
Arthrinium aquaticum Submerged decaying wood (river) China Luo etal. (2019)
Arthrinium arundinis Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Arthrinium sinense Submerged decaying wood (lake) China Luo etal. (2004)
Beltraniaceae
Beltrania africana Submerged decaying leaves of Calophyllum Brasiliense
(stream) Brazil Fiuza etal. (2019)

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Beltrania rhombica Foam (river, stream), submerged decaying leaves and peti-
oles (stream) Brazil, India,
Thailand,
Venezuela
Rajashekhar and Kaveriappa
(2003), Sakayaroj etal.
(2005), Smits etal. (2007),
Barbosa etal. (2013b)
Beltraniella botryospora Submerged decaying leaves of Calophyllum Brasiliense
(stream) Brazil Fiuza etal. (2019)
Beltraniella fertilis Submerged decaying leaves of Calophyllum Brasiliense
(stream) Brazil Fiuza etal. (2019)
Beltraniella portoricensis Submerged decaying leaves (stream), foam/water (stream) Brazil, Ven-
ezuela Smits etal. (2007), Barbosa
etal. (2013b)
Beltraniopsis aquatica Submerged decaying leaves (stream_ Cuba Castañeda-Ruiz etal. (2006)
Hemibeltrania decorosa Submerged decaying leaves of Calophyllum Brasiliense
(stream) Brazil Fiuza etal. (2019)
Hyponectriaceae
Arecomyces bruneiensis Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Arecomyces epigenus Submerged decaying Licuala longicalycata and Nenga
pumila (peat swamp) Thailand Pinruan etal. (2007)
Arecomyces frondicola Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Atractium holubovae Submerged decaying wood (e.g., Xylia dolabriformis test
block) (river, stream) China, Philip-
pines, South
Africa,
Thailand
Seifert etal. (1995), Ho etal.
(2001), Sivichai etal. (2002)
Physalospora aquatica Submerged Eleocharis palustris (lake) UK Ingold (1955)
Physalospora limnetica Submerged herbaceous debris (pond) USA Raja and Shearer (2008)
Physalospora vaccinii Submerged partially decomposed leaves of Vaccinium
oxycoccus (lake) USA Fallah and Shearer (2001)
Pseudomassariaceae
Leiosphaerella leonensis Submerged twig Africa Fisher and Petrini (1983)
Pseudomassaria sepincoli-
formis Submerged decaying wood China Chen etal. (2011)
Sporocadaceae
Bartalinia robillardoides Water (pond, stream) Korea Nguyen etal. (2019a)
Discosia rubi Submerged plant litter Korea Goh etal. (2020)
Discostroma corticola Scots pine wood UK Eaton and Jones (1971a)
Hymenopleella lakefuxian-
ensis Submerged decaying wood (lake) China Jeewon etal. (2003)
Pestalotiopsis submersa Submerged decaying rotten needles of Pinus roxburghii India Sati and Tiwari (1993)
Robillarda sessilis Submerged plant litter Korea Goh etal. (2020)
Seiridium aquaticum Submerged decaying wood (stream) China Luo etal. (2019)
Xylariales
Castanediellaceae
Castanediella cagnizarii Submerged decaying leaves (stream) Brazil, Mexico Barbosa etal. (2013b),
Castañeda-Ruiz etal. (2005)
Castanediella ramosa Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Clypeosphaeriaceae
Apioclypea apiosporioides Submerged decaying Eleiodoxa conferta (peat awamp) Thailand Pinnoi etal. (2006)
Clypeosphaeria mamillana Scots pine wood UK Eaton and Jones (1971b)
Diatrypaceae
Peroneutypa lignicola Submerged decaying wood (stream) China Luo etal. (2019)
Hypoxylaceae
Daldinia kretzschmarioides Submerged decaying wood (stream) Brunei Ho etal. (2001)
Hypoxylon lignicola Submerged decaying wood (river) China Luo etal. (2019)
Microdochiaceae

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Selenodriella fertilis Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Oxydothidaceae
Oxydothis angustispora Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Oxydothis atypica Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Oxydothis frondicola Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Oxydothis grisea Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis hoehnelii Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis licualae Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis livistonae Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis livistonica Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis oraniopsidis Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Oxydothis rattanicola Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Zygosporiaceae
Vesiculozygosporium echino-
sporum Submerged decaying leaves and petiole (stream) Brazil Barbosa etal. (2013b)
Zygosporium masonii Submerged decaying wood (stream), submerged decaying
leaves of Calophyllum brasiliense (stream) Brazil Barbosa etal. (2013b), Fiuza
etal. (2018)
Zygosporium minus Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Xylariaceae
Anthostomella aquatica Submerged decaying wood of Dipterocarpus alatus, Xylia
dolabriformis and unidentified wood (reservoir, stream) Australia,
China, Thai-
land
Hyde and Goh (1998b), Goh
and Hyde (1999), Sivichai
etal. (2002)
Anthostomella palmaria Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Astrocystis eleiodoxae Submerged decaying petiole of Eleiodoxa conferta and
Licuala longicalycata (peat swamp) Thailand Pinnoi etal. (2010), Pinruan
etal. (2014)
Astrocystis rachidis Decaying Licuala longicalycata and Metroxylon sagu (peat
swamp) Thailand Pinruan etal. (2014)
Rosellinia corticium Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2007)
Vamsapriyaceae
Vamsapriya aquatica Submerged decaying bamboo culms China Bao etal. (2021a)
Vamsapriya indica Submerged decaying bamboo culms (river) China Bao etal. (2021a)
Xylariales genera incertae sedis
Circinotrichum papakurae Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Gyrothrix circinata Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Gyrothrix magica Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Gyrothrix microsperma Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Gyrothrix podosperma Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Gyrothrix verticiclada Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Lanceispora amphibia Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Melanographium citri Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Melanographium palmicola Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Polyancora globosa Submerged decaying leaves (peat swamp) Malaysia Voglmayr and Yule (2006)
Roselymyces brasiliensis Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2018)
Stilbohypoxylon elaeidicola Decaying Eleiodoxa conferta, Licuala longicalycata (peat
swamp) Thailand Pinruan etal. (2014)
Stilbohypoxylon moelleri Submerged decaying Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006)
Xylariomycetidae family incertae sedis
Cainiaceae
Endocalyx melanoxanthus Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Myelospermataceae

Fungal Diversity
1 3
Table 9 (continued)
Taxon Substrate Distribution References
Myelosperma tumidum Submerged decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Sordariomycetes genera incertae sedis
Aquadulciospora rhomboidia Submerged decomposed stems of Scirpus validus (lake) USA Fallah and Shearer (2001)
Aquasphaeria dimorphospora Submerged decaying wood Australia Hyde (1995d)
Ascoyunnania aquatica Submerged bamboo (stream) China Cai etal. (2005)
Ellisembia acicularia Submerged decaying culms of Miscanthus floridulus
(stream) China Kuo and Goh (2019)
Ellisembia bambusicola Submerged decaying wood (stream) Brazil, China Ho etal. (2001), Barbosa etal.
(2013b)
Ellisembia brachypus Submerged decaying Dipterocarpus alatus and Xylia dola-
briformis test block Thailand Sivichai etal. (2002)
Ellisembia flagelliformis Submerged leaves Brazil da Silva and Gusmão (2015)
Ellisembia opaca Submerged decaying wood (stream) China Ho etal. (2001)
Ellisembia plovercovensis Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Ellisembia repentioriunda Submerged decaying wood (lake, reservoir) China Goh and Hyde (1999), Luo
etal. (2004)
Ellisembia vaginata Submerged twig Brazil da Silva and Gusmão (2015)
Flammispora bioteca Submerged dead leaves of Licuala longicalycata (peat
swamp) Thailand Pinruan etal. (2004b), Pinruan
etal. (2007)
Flammispora pulchra Submerged decaying wood USA Raja and Shearer (2008)
Phialemoniopsis endophytica Submerged plant litter (stream) Korea Goh etal. (2020)
Pleurophragmium bitunicatum Submerged decaying wood (reservoir, river) and bamboo
(river) China, Philip-
pines Goh and Hyde (1999), Cai etal.
(2003a)
Pleurophragmium malay-
sianum Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Potamomyces armatisporus Submerged decaying wood (river) Australia,
Malaysia Hyde (1995d), Hyde (2007)
Paoayensis lignicola Submerged decaying wood (lake) Philippines Cabanela etal. (2007)
Plagiosphaera nilotica Submerged twig Sudan Monod and Fisher (1983)
Rivulicola aquatica Submerged decaying wood (reservoir) China Ranghoo etal. (2000)
Rivulicola cygnea Submerged decaying wood (lake) USA Raja etal. (2009a)
Rivulicola incrustata Submerged decaying wood Australia Hyde etal. (1997)
Saccardoella allequashensis Submerged decaying wood (lake) USA Fallah and Shearer (2001)
Saccardoella horizontalis Submerged decaying wood (river) Philippines,
USA Fallah and Shearer (2001), Cai
etal. (2003c)
Saccardoella lacustris Submerged decaying wood (lake) USA Fallah and Shearer (2001)
Saccardoella minuta Submerged bamboo and wood (river) Philippines Cai etal. (2002a, 2003b)
Stanjehughesia clavata Submerged decaying bark Brazil de Almeida etal. (2014)
Stanjehughesia curviapicis Submerged decaying wood (stream, river) Australia,
China Ho etal. (2001), Goh and Hyde
(2018)
Stanjehughesia kaohsiun-
gensis Submerged decaying wood (stream) China Hsieh etal. (2021b)
Selenosporella aristata Submerged decaying leaves (stream) Malaysia Kuthubutheen and Nawawi
(1988b)
Selenosporella curvispora Submerged decaying leaves of Calophyllum brasiliense
(stream) Brazil Fiuza etal. (2018)
Selenosporella minima Submerged decaying leaves of Calophyllum brasiliens Brazil Fiuza etal. (2015c)
Selenosporella perramosa Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)

Fungal Diversity
1 3
(1905–2010) who was the first pioneer of this fungal group
and for his recognition as one of the most British distin-
guished mycologists of the twentieth century (Money 2010).
Ingoldian fungi are an ecologically heterogeneous group of
freshwater asexual morphs of Ascomycota or Basidiomy-
cota. Uniquely, they produce relatively modified conidia
(i.e., hyaline, filiform, tetraradiate, multiradiate, scolecoid,
sigmoid or branched as shown in Fig.7) for decomposition
on submerged plants and for their reproduction in aquatic
ecosystems (Ingold 1975; Descals 2005; Shearer etal. 2007;
Seifert etal. 2011). Ingoldian fungi are often found in fast-
flowing water, well-aerated forest streams and humid envi-
ronments with ubiquitous distribution, but mostly in temper-
ate zones of the earth (Bärlocher 1992; Jones and Pang 2012;
Duarte etal. 2016). Currently, there are over 95 genera with
approximately 330 species discovered, and ca 28 genera are
monotypic (Descals 2005; Baschien etal. 2006; Shearer
etal. 2007; Campbell etal. 2009; Fiuza etal. 2017a; Tarda
etal. 2019). In Table14, most of which are in Ascomycota
i.e., Helotiales and Leotiales (Leotiomycetes), some of them
are phylogenetically related to Jahnulales and Pleosporales
(Dothideomycetes), Orbiliales (Orbiliomycetes), Hypocre-
ales, Sordariales and Microascales (Sordariomycetes), while
rarely few taxa are Basidiomycota i.e., Cantharellales and
Atheliales (Shearer etal. 2007).
Classication offreshwater fungal basal
clades
Early diverging lineages of the fungal tree of life consist
of zoosporic and zygomycetous (zygosporic) fungi (James
etal. 2006; Voigt etal. 2021). They comprise saprobic and
parasitic fungi isolated from various niches, such as soil
and freshwater habitats (Ishida etal. 2015). Among basal
fungi, the most common in freshwater habitats are the zoo-
sporic ones, such as Chytridiomycota, Blastocladiomycota,
Monoblepharomycota, Rozellomycota and Aphelidiomycota
(Calabon etal. 2020b; Voigt etal. 2021). Zoosporic fungal
parasites control the numbers of numerous phytoplankton
species such as diatoms, green algae, cyanobacteria and
dinoflagellates (Kagami etal. 2007, 2014; Ishida etal. 2015).
Estimation of fungal numbers has always been a debated
research topic. Recent estimates of fungal species range
from 2.2 to 3.8 million, of which only 100,000 to 143,273
have been described (Hawksworth and Lücking 2017; Hyde
etal. 2020c). Most of the described fungal species are from
terrestrial habitats, while the discovery and description of
aquatic fungi are significantly lower, and even smaller for
basal fungi. The diversity and abundance of basal fungi in
aquatic niches have recently started to gain the spotlight in
mycological research. For instance, studies have shown that
Fig. 6 Major families among different orders of Sordariomycetes taxa from freshwater

Fungal Diversity
1 3
Table 10 Classification of freshwater lichens from different fungal classes with information on the type of habitats, substrates, and geographical
distribution of fungal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
ARTHONIOMYCETES
Arthoniales
Arthoniaceae
Arthonia caudata* River watershed Canada Maloles etal. (2018)
Arthonia helvola* River watershed Canada Maloles etal. (2018)
Arthonia radiata * River watershed Canada Maloles etal. (2018)
Chrysotrichaceae
Chrysothrix caesia* River watershed Canada Maloles etal. (2018)
Chrysothrix granulosa* River Canada Björk etal. (2009)
Lecanographaceae
Alyxoria varia* River watershed Canada Maloles etal. (2018)
Opegraphaceae
Cresponea chloroconia* River watershed Canada Maloles etal. (2018)
Roccellaceae
Enterographa osagensis#Creek Central North America Morse (2013)
Enterographa zonata*Waterfall Canada Halda (2011)
Gyrographa gyrocarpa*North America Earland-Bennett and Seaward (1974)
Pseudoschismatomma rufescens* River watershed Canada Maloles etal. (2018)
Schismatomma cretaceum*Nature reserve Ukraine Khodosovtsev etal. (2016)
CANDELARIOMYCETES
Candelariales
Candelariaceae
Candelaria concolor* River watershed Canada Maloles etal. (2018)
Candelariella aurella* River watershed Canada Maloles etal. (2018)
Candelariella efflorescens* River watershed Canada Maloles etal. (2018)
Candelariella vitellina* River watershed Canada Maloles etal. (2018)
CONIOCYBOMYCETES
Coniocybales
Coniocybaceae
Chaenotheca balsamconensis* River watershed Canada Maloles etal. (2018)
Chaenotheca ferruginea* River watershed Canada Maloles etal. (2018)
Chaenotheca furfuracea* River watershed Canada Maloles etal. (2018)
Chaenotheca xyloxena* River watershed Canada Maloles etal. (2018)
DOTHIDEOMYCETES
Capnodiales
Stigmidium rivulorum#River Bulgaria Shivarov etal. (2018)
Collemopsidiales
Xanthopyreniaceae
Collemopsidium japonicum* Nature reserves Japan Harada (1999)
Collemopsidium monense* Stream Europe Gilbert and Giavarini (1997)
Zwackhiomyces lacustris#Stream and lake margin Europe Orange (2002)
Lichenotheliales
Lichenotheliaceae
Endococcus rugulosus#Stream and lake margin Europe Orange (2002)
Endococcus verrucisporus#Stream and lake margin Europe Orange (2002)
Monoblastiales
Monoblastiaceae
Acrocordia cavata* River watershed Canada Maloles etal. (2018)
Pleosporales
Mycoporaceae

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Mycoporum antecellens* Twig (waterfall) Canada Björk etal. (2009)
Phaeosphaeriaceae
Didymocyrtis pseudeverniae#Nature reserve Ukraine, UK Khodosovtsev etal. (2016)
Trypetheliales
Polycoccaceae
Polycoccum sp.#Stream and lake margin Europe Orange (2002)
Trypetheliaceae
Arthopyrenia fallaciosa* River watershed Canada Maloles etal. (2018)
EUROTIOMYCETES
Collemopsidiales
Xanthopyreniaceae
Sagediopsis lomnitzensis#Stream and lake margin Europe Orange (2002)
Mycocaliciales
Mycocaliciaceae
Chaenothecopsis debilis* River watershed Canada Maloles etal. (2018)
Chaenothecopsis pusiola* River watershed Canada Maloles etal. (2018)
Chaenothecopsis savonica* River watershed Canada Maloles etal. (2018)
Phaeocalicium curtisii* River watershed Canada Maloles etal. (2018)
Phaeocalicium polyporaeum* River watershed Canada Maloles etal. (2018)
Phaeocalicium populneum* River watershed Canada Maloles etal. (2018)
Sphinctrina anglica#River watershed Canada Maloles etal. (2018)
Stenocybe pullatula* River watershed Canada Maloles etal. (2018)
Verrucariales
Adelococcaceae
Sagediopsis lomnitzensis#Stream and lake margin Europe Orange (2002)
Verrucariaceae
Agonimia allobata* Stream Lithuania Motiejunaite (2003)
Agonimia tristicula* Stream UK Gilbert and Giavarini (1997)
Catapyrenium lachneum* Stream UK Gilbert and Giavarini (1997)
Catapyrenium squamulosum* River watershed Canada Maloles etal. (2018)
Catillaria cf. chalybeia* Stream Gwynedd Pentecost (1977)
Dermatocarpon arnoldianum* River and stream Europe Gilbert and Giavarini (1997), Shiv-
arov etal. (2018)
Dermatocarpon intestiniforme* Stream UK Orange (1998)
Dermatocarpon leptophyllodes* Stream UK Gilbert and Giavarini (1997)
Dermatocarpon luridum* Stream UK Pentecost (1977), Gilbert and Gia-
varini (1997)
Dermatocarpon meiophyllizum* Stream UK Gilbert and Giavarini (1997)
Dermatocarpon miniatum* River Bulgaria Shivarov etal. (2018)
Dermatocarpon rivulorum* Stream and river Bulgaria, UK Gilbert and Giavarini (1997), Shiv-
arov and Lőkös (2015)
Endocarpon adscendens* Stream UK Gilbert and Giavarini (1997)
Endocarpon globosum* Stream China Harada and Wang (1996)
Hydropunctaria orae* Stream Orange (2013), Thüs etal. (2014)
Hydropunctaria rheitrophila* River and stream Poland, alpine Australia Gilbert and Giavarini (1997),
McCarthy (2002), Matura and
Krzewicka (2015)
Hydropunctaria scabra* Siliceous rock (stream) Bulgaria Shivarov etal. (2017)
Muellerella pygmaea#Stream and lake margin Europe Orange (2002)
Placopyrenium cinereoatratum#Siliceous rocks (brook, lake, river, stream) Bulgaria, Finland, Norway,
Sweden, UK Orange (2009)
Placopyrenium formosum#Rock (river) Finland, Iceland, UK Orange (2009)
Polyblastia ardesiaca* – Italy Nascimbene and Nimis (2006)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Psoroglaena stigonemoides* Twig (waterfall) Canada Björk etal. (2009)
Sporodictyon cruentum* River and stream Alpine Australia, Italy, UK Gilbert and Giavarini (1997), McCa-
rthy (2002), Nascimbene and
Nimis (2006), Hafellner (2010)
Staurothele acarosporoides* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele arenaria* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele chlorospora* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele clopimoides* River Italy, USA Thomson (1991), Nascimbene and
Nimis (2006)
Staurothele fauriei* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele fissa* Lakes and stream Europe Lindsay (1856), Gilbert and Gia-
varini (1997), Nascimbene and
Nimis (2006), Shivarov (2017)
Staurothele fuscocuprea* – Italy Nascimbene and Nimis (2006)
Staurothele honghensis* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele kwapiensis* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele muliensis* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele ochroplaca* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele pachystroma* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele paraguayensis* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele sinensis* Nature reserves Vietnam Gueidan etal. (2014)
Staurothele succedens* Stream Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006)
Staurothele yunnanica* Nature reserves Vietnam Gueidan etal. (2014)
Thelidium aquaticum* Stream Poland Matura and Krzewicka (2015)
Thelidium carbonaceum* Nature reserves Australia McCarthy and Kantvilas (2020)
Thelidium circumspersellum* Stream Poland Matura etal. (2017)
Thelidium epomphalum* Creek and river Europe Thüs and Nascimbene (2008)
Thelidium fontigenum* – Europe Thüs and Nascimbene (2008)
Thelidium klementii* Stream Poland Matura etal. (2017)
Thelidium lahmianum* – Europe Thüs and Nascimbene (2008)
Thelidium methorium* – Europe Thüs and Nascimbene (2008)
Thelidium minutulum* Stream Poland, UK Gilbert and Giavarini (1997),
Matura and Krzewicka (2015)
Thelidium nigricans* Siliceous rocks (creek's splash water zone) Switzerland Thüs and Nascimbene (2008)
Thelidium papulare* Stream UK Pentecost (1977), Gilbert and Gia-
varini (1997)
Thelidium pertusatii* – Italy Nascimbene and Nimis (2006)
Thelidium pluvium* Stream Poland, UK Gilbert and Giavarini (1997),
Matura etal. (2017)
Thelidium rehmii* Stream Poland Matura etal. (2017)
Thelidium rivulicola* – Europe Thüs and Nascimbene (2008)
Thelidium spadanum* – Europe Thüs and Nascimbene (2008)
Thelidium sublacteum* – Europe Thüs and Nascimbene (2008)
Thelidium submethorium* – Europe Thüs and Nascimbene (2008)
Thelidium suzaeanum* – Europe Thüs and Nascimbene (2008)
Thelidium zwackhii* Stream Poland Matura and Krzewicka (2015)
Verrucaria aethiobola* Stream Italy, UK Pentecost (1977), Nascimbene and
Nimis (2006)
Verrucaria alpicola* Pebble (river) Bulgaria Shivarov etal. (2017)
Verrucaria amnica* Mountain stream New Zealand McCarthy and Johnson (1997)
Verrucaria aquatilis* Nature reserve, river, stream Australia, Italy, UK Gilbert and Giavarini (1997),
McCarthy (2002), Nascimbene
etal. (2009)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Verrucaria austroalpina* Creeks and rivers Australia McCarthy (1995)
Verrucaria basaltica* Streams and lakes Poland Krzewicka (2012)
Verrucaria calkinsiana* River watershed Canada Maloles etal. (2018)
Verrucaria consociata* Stream UK Orange (2015)
Verrucaria conturmatula#Stream and lake margin Europe Orange (2002)
Verrucaria cootapatambensis* River Alpine Australia McCarthy (2002)
Verrucaria devensis* Stream Poland Matura and Krzewicka (2015),
Matura etal. (2017)
Verrucaria dolosa* Stream Poland, UK Gilbert and Giavarini (1997),
Matura and Krzewicka 2015)
Verrucaria elaeina* Stream Poland Matura and Krzewicka (2015)
Verrucaria elaeomelaena* Nature reserve, stream Italy, Poland, UK Pentecost (1977), Nascimbene and
Nimis (2006), Nascimbene etal.
(2009), Matura and Krzewicka
(2015)
Verrucaria funckii* Stream France, Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006), Coste
etal. (2016)
Verrucaria glaucina* Stream UK Gilbert and Giavarini (1997)
Verrucaria hydrela* Stream UK Pentecost (1977), Gilbert and Gia-
varini (1997)
Verrucaria hydrophila* Stream Poland, Europe Orange (2013), Matura and Krze-
wicka (2015)
Verrucaria igii* River Alpine Australia McCarthy (2002)
Verrucaria inconstans* Creeks and rivers Eastern Australia McCarthy (1995)
Verrucaria latebrosa* Stream Italy, Poland, UK Gilbert and Giavarini (1997),
Nascimbene and Nimis (2006),
Matura and Krzewicka (2015)
Verrucaria maas-geesterani* Lake Switzerland Servít (1954)
Verrucaria margacea* Creeks, rivers, stream Eastern Australia, UK Gilbert and Giavarini (1997), McCa-
rthy (2003)
Verrucaria morosa* River Alpine Australia McCarthy (2002)
Verrucaria mundula* Creeks, rivers, stream Eastern Australia Valcárcel etal. (2010)
Verrucaria nodosa* Stream Europe Orange (2013)
Verrucaria pachyderma* Stream Australia, Italy, Europe Nascimbene and Nimis (2006),
Krzewicka (2012)
Verrucaria phaeoderma* Creeks, rivers, and stream Eastern Australia, France, UK McCarthy (1999, 2004)
Pentecost (1977), Coste etal. (2016)
Verrucaria placida* Stream Europe Orange (2013)
Verrucaria praetermissa* River and stream Italy, Europe Gilbert and Giavarini (1997),
Nascimbene and Nimis (2006),
Orange (2014)
Verrucaria puncticulata* Creeks and rivers Eastern Australia McCarthy (2002)
Verrucaria rosula* Stream UK, Europe Orange (2013), Orange (2015)
Verrucaria ruttneri* River Alpine Australia McCarthy (2002)
Verrucaria submersella* Stream Poland Matura and Krzewicka (2015)
Verrucaria sublobulata* Stream UK Orange (2015)
Verrucaria tasmanica* River, creeks, and rivers Alpine Australia, Eastern Australia McCarthy (1995, 2002)
Verrucaria tholocarpa* Creeks and rivers Eastern Australia McCarthy (1995)
Willeya australis* Nature reserves Vietnam Gueidan etal. (2014)
Willeya diffractella* Nature reserves Vietnam Gueidan etal. (2014)
Willeya fusca* Nature reserves Vietnam Gueidan etal. (2014)
Willeya iwatsukii* Nature reserves, river Japan, Vietnam Harada (1992), Gueidan etal.
(2014)
Willeya japonica* Nature reserves Vietnam Gueidan etal. (2014)
Willeya laevigata* Nature reserves Vietnam Gueidan etal. (2014)
Willeya malayensis* Nature reserves Vietnam Gueidan etal. (2014)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Willeya microlepis* Nature reserves Vietnam Gueidan etal. (2014)
Willeya pallidipora* Nature reserves Vietnam Gueidan etal. (2014)
Willeya protrudens* Nature reserves Vietnam Gueidan etal. (2014)
Willeya rimosa* Nature reserves Vietnam Gueidan etal. (2014)
LECANOROMYCETES
Acarosporales
Acarosporaceae
Acarospora fuscata* River watershed Canada Maloles etal. (2018)
Acarospora schleicheri*Waterfall Canada Hawksworth (2000), Björk etal.
(2009)
Acarospora smaragdula* Stream UK Gilbert and Giavarini (1997)
Sarcogyne regularis* River watershed Canada Maloles etal. (2018)
Arctomiales
Arctomiaceae
Arctomia delicatula* Twig (lake) Canada Björk etal. (2009)
Baeomycetales
Baeomycetaceae
Baeomyces rufus* Stream UK Gilbert and Giavarini (1997)
Caliciales
Caliciaceae
Amandinea punctata* River watershed Canada Maloles etal. (2018)
Buellia aethale* Stream UK Gilbert and Giavarini (1997)
Cyphelium tigillare* River watershed Canada Maloles etal. (2018)
Diplotomma venustum* River watershed Canada Maloles etal. (2018)
Physciaceae
Heterodermia albicans* Stream USA Mason (1984)
Hyperphyscia adglutinata* River watershed Canada Maloles etal. (2018)
Phaeophyscia adiastola* River watershed Canada Maloles etal. (2018)
Phaeophyscia orbicularis* Stream UK Gilbert and Giavarini (1997)
Phaeophyscia pusilloides* River watershed Canada Maloles etal. (2018)
Phaeophyscia rubropulchra* River watershed Canada Maloles etal. (2018)
Phaeophyscia sciastra* Stream UK Gilbert and Giavarini (1997)
Physcia caesia* Stream UK Gilbert and Giavarini (1997)
Physcia dubia* River watershed Canada Maloles etal. (2018)
Physcia millegrana* River watershed Canada Maloles etal. (2018)
Physcia stellaris* River watershed Canada Maloles etal. (2018)
Physciella chloantha* River watershed Canada Maloles etal. (2018)
Physconia detersa* River watershed Canada Maloles etal. (2018)
Physconia enteroxantha* River watershed Canada Maloles etal. (2018)
Physconia perisidiosa* River watershed Canada Maloles etal. (2018)
Rinodina fimbriata* Stream UK Gilbert and Giavarini (1997)
Rinodina freyi* River watershed Canada Maloles etal. (2018)
Rinodina teichophila* Stream UK Gilbert and Giavarini (1997)
Rinodina rivularis* River China Harada and Wang (2004)
Hymeneliaceae
Hymenelia cyanocarpa* Stream UK Gilbert and Giavarini (1997)
Ionaspis lacustris* Stream and river Alpine Australia, France, UK Pentecost (1977), Gilbert and Gia-
varini (1997), McCarthy (2002),
Coste etal. (2016)
Ionaspis odora* – Italy Nascimbene and Nimis (2006)
Ionaspis sp.* River Alpine Australia McCarthy (2002)
Protothelenellaceae
Thrombium thelostomum* Stream UK Gilbert and Giavarini (1997)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Trapeliaceae
Placopsis gelida* Stream UK Gilbert and Giavarini (1997)
Sarea resinae* River watershed Canada Maloles etal. (2018)
Trapelia coarctata* Stream UK Gilbert and Giavarini (1997)
Trapelia involuta* Stream UK Gilbert and Giavarini (1997)
Trapelia obtegens* Stream UK Gilbert and Giavarini (1997)
Trapelia placodioides* Stream, river watershed Canada, UK Gilbert and Giavarini (1997)
Maloles etal. (2018)
Xylographaceae
Xylographa vitiligo* Nature reserve Ukraine Khodosovtsev etal. (2016)
Lecanorales
Pilocarpaceae
Micarea peliocarpa* Stream UK Gilbert and Giavarini (1997)
Micarea prasina* River watershed Canada Maloles etal. (2018)
Micarea viridiatra* Nature reserve Ukraine Khodosovtsev etal. (2016)
Micarea viridileprosa* Nature reserve Ukraine Khodosovtsev etal. (2016)
Catillariaceae
Catillaria chalybeia* Stream UK Gilbert and Giavarini (1997)
Catillaria nigroclavata* River watershed Canada Maloles etal. (2018)
Cladoniaceae
Cladonia cervicornis* River watershed Canada Maloles etal. (2018)
Cladonia chlorophaea* River watershed Canada Maloles etal. (2018)
Cladonia fimbriata* River watershed Canada Maloles etal. (2018)
Cladonia furcata* River watershed Canada Maloles etal. (2018)
Cladonia macilenta* River watershed Canada Maloles etal. (2018)
Cladonia ochrochlora* River watershed Canada Maloles etal. (2018)
Cladonia pocillum* River watershed Canada Maloles etal. (2018)
Cladonia pyxidata* River watershed Canada Maloles etal. (2018)
Cladonia rangiferina* River watershed Canada Maloles etal. (2018)
Cladonia rei* River watershed Canada Maloles etal. (2018)
Lepraria lobificans* River watershed Canada Maloles etal. (2018)
Stereocaulon saxatile* River watershed Canada Maloles etal. (2018)
Haematommataceae
Haematomma ochroleucum* Waterfall Canada Björk etal. (2009)
Lecanoraceae
Claurouxia chalybeioide* Stream UK Gilbert and Giavarini (1997)
Lecanora campestris* Stream UK Gilbert and Giavarini (1997)
Lecanora epanora* Stream UK Gilbert and Giavarini (1997)
Lecanora gangaleoides* Stream UK Gilbert and Giavarini (1997)
Lecanora hybocarpa* River watershed Canada Maloles etal. (2018)
Lecanora pulicaris* River watershed Canada Maloles etal. (2018)
Lecanora soralifera* Stream UK Gilbert and Giavarini (1997)
Lecanora symmicta* River watershed Canada Maloles etal. (2018)
Lecidella carpathica* Stream UK Gilbert and Giavarini (1997)
Lecidella scabra* Stream UK Gilbert and Giavarini (1997)
Lecidella stigmatea* Stream UK Gilbert and Giavarini (1997)
Lepra amara* River watershed Canada Maloles etal. (2018)
Myriolecis hagenii* River watershed Canada Maloles etal. (2018)
Polyozosia dispersa* Stream UK Gilbert and Giavarini (1997)
Polyozosia sambuci* River watershed Canada Maloles etal. (2018)
Polyozosia semipallida* River watershed Canada Maloles etal. (2018)
Protoparmeliopsis muralis* River watershed Canada Maloles etal. (2018)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Verseghya thysanophora* River watershed Canada Maloles etal. (2018)
Parmeliaceae
Evernia mesomorpha* River watershed Canada Maloles etal. (2018)
Flavoparmelia caperata* River watershed Canada Maloles etal. (2018)
Flavopunctelia flaventior* River watershed Canada Maloles etal. (2018)
Flavopunctelia soredica* River watershed Canada Maloles etal. (2018)
Hypogymnia oceanica*Lake, river, waterfall Canada Björk etal. (2009)
Hypogymnia physodes* River watershed Canada Maloles etal. (2018)
Melanelixia subaurifera* River watershed Canada Maloles etal. (2018)
Parmelia sulcata* River watershed Canada Maloles etal. (2018)
Protoparmelia hypotremella* River watershed Canada Maloles etal. (2018)
Punctelia rudecta* River watershed Canada Maloles etal. (2018)
Usnea hirta* River watershed Canada Maloles etal. (2018)
Xanthoparmelia conspersa* Italy Gilbert and Giavarini (1997)
Xanthoparmelia cumberlandia* River watershed Canada Maloles etal. (2018)
Psoraceae
Protoblastenia rupestris* River watershed Canada Maloles etal. (2018)
Ramalinaceae Stream UK Gilbert and Giavarini (1997)
Bacidia arnoldiana* Stream UK Gilbert and Giavarini (1997)
Bacidia fuscoviridis* Stream UK Gilbert and Giavarini (1997)
Bacidia inundata* Stream UK Gilbert and Giavarini (1997)
Bacidina delicata* Stream UK Motiejunaite (2003)
Bacidina inundata* Stream UK Gilbert and Giavarini (1997)
Bilimbia sabuletorum* River watershed Canada Maloles etal. (2018)
Lecania croatica* River watershed Canada Maloles etal. (2018)
Lecania erysibe* Stream UK Gilbert and Giavarini (1997)
Lecania hutchinsiae* Stream Lithuania Motiejunaite (2003)
Ramalina intermedia* River watershed Canada Maloles etal. (2018)
Scoliciosporaceae
Scoliciosporum chlorococcum* River watershed Canada Maloles etal. (2018)
Scoliciosporum umbrinum* Stream UK Gilbert and Giavarini (1997)
Stereocaulaceae
Stereocaulon pileatum* Stream UK Řídká etal. (2013)
Lecideales
Lecideaceae
Bryobilimbia ahlesii* Stream Poland Matura etal. (2017)
Koerberiella wimmeriana* – Italy Nascimbene and Nimis (2006)
Porpidia crustulata* Stream UK Gilbert and Giavarini (1997)
Porpidia hydrophila* stream France, Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006), Coste
etal. (2016)
Porpidia macrocarpa* River watershed Canada Maloles etal. (2018)
Porpidia platycarpoides* Stream UK Gilbert and Giavarini (1997)
Porpidia soredizodes* Stream Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006)
Porpidia tuberculosa* Stream UK Gilbert and Giavarini (1997)
Ostropales
Coenogoniaceae
Coenogonium pineti* River watershed Canada Maloles etal. (2018)
Graphidaceae
Graphis scripta* River watershed Canada Maloles etal. (2018)
Gyalectaceae
Gyalecta jenensis* Stream UK Gilbert and Giavarini (1997)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Gyalidea roseola* Stream UK Gilbert and Giavarini (1997)
Porinaceae
Porina ahlesiana* Stream UK Gilbert and Giavarini (1997)
Porina lectissima* Stream Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006)
Porina otagensis* Semi-aquatic New Zealand McCarthy (1999)
Pseudosagedia chlorotica* Stream Lithuania, UK Gilbert and Giavarini (1997), Motie-
junaite (2003)
Pseudosagedia guentheri*Nature reserve Ukraine, UK Khodosovtsev etal. (2016), Thüs
etal. (2014)
Pseudosagedia interjungens* Stream Ukraine Gilbert and Giavarini (1997)
Porina grandis* - - Thüs etal. (2014)
Porina rivalis* Stream, river UK Orange (2015)
Pertusariales
Megasporaceae
Aspicilia aquatica* Stream UK Gilbert and Giavarini (1997)
Aspicilia caesiocinerea* Stream UK Gilbert and Giavarini (1997)
Aspicilia inornata* Stream UK Gilbert and Giavarini (1997)
Aspicilia laevata* Stream UK Gilbert and Giavarini (1997)
Aspicilia laevatoides* Stream UK Gilbert and Giavarini (1997)
Lobothallia melanaspis* – Italy Nascimbene and Nimis (2006)
Microcaliciaceae
Microcalicium ahlneri* River watershed Canada Maloles etal. (2018)
Ochrolechiaceae
Ochrolechia arborea* River watershed Canada Maloles etal. (2018)
Pertusariaceae
Loxosporopsis corallifera* River Canada Björk etal. (2009)
Pertusaria macounii* River watershed Canada Maloles etal. (2018)
Peltigerales
Collemataceae
Collema elenkii* – – Thüs etal. (2014)
Collema flaccidum* Stream UK Gilbert and Giavarini (1997)
Collema ramenskii* – Russia Kulikova etal. (2008)
Enchylium bachmanianum* Stream UK Gilbert and Giavarini (1997)
Enchylium tenax* Stream UK Gilbert and Giavarini (1997)
Lathagrium dichotomum* Stream UK Gilbert and Giavarini (1997)
Leptogium cyanescens* River watershed Canada Maloles etal. (2018)
Leptogium teretiusculum* Stream UK Gilbert and Giavarini (1997)
Scytinium gelatinosum* Stream UK Gilbert and Giavarini (1997)
Scytinium lichenoides* River watershed Canada Maloles etal. (2018)
Scytinium massiliense* Stream UK Gilbert and Giavarini (1997)
Scytinium plicatile* Stream UK Gilbert and Giavarini (1997)
Scytinium rivale* River USA Glavich and Geiser (2007)
Lobariaceae
Pseudocyphellaria crocata* Twig (waterfall) Canada Björk etal. (2009)
Pseudocyphellaria mallota* Twig (river) Canada Björk etal. (2009)
Massalongiaceae
Massalongia carnosa* Stream UK Gilbert and Giavarini (1997)
Peltigeraceae
Nephroma occultum* Bark (lake, river) Canada Björk etal. (2009)
Peltigera canina* River watershed Canada Maloles etal. (2018)
Peltigera elisabethae* River watershed Canada Maloles etal. (2018)
Peltigera evansiana * River watershed Canada Maloles etal. (2018)
Peltigera horizontalis* Stream UK Gilbert and Giavarini (1997)

Fungal Diversity
1 3
Table 10 (continued)
Taxon Substrate Distribution References
Peltigera hydrothyria* River USA Glavich and Geiser (2007), Thüs
etal. (2014)
Peltigera praetextata* Stream UK Gilbert and Giavarini (1997)
Peltigera rufescens* River watershed Canada Maloles etal. (2018)
Placynthiaceae
Placynthium flabellosum* Stream Italy, UK Gilbert and Giavarini (1997), Nas-
cimbene and Nimis (2006)
Placynthium nigrum* River watershed Canada Maloles etal. (2018)
Placynthium rosulans* – Italy Nascimbene and Nimis (2006)
Placynthium tantaleum* Stream UK Gilbert and Giavarini (1997)
Rhizocarpales
Rhizocarpaceae
Rhizocarpon amphibium* Stream UK Gilbert and Giavarini (1997)
Rhizocarpon lavatum* Stream UK Pentecost (1977), Gilbert and Gia-
varini (1997)
Rhizocarpon sublavatum* Stream Poland Matura etal. (2017)
Teloschistales
Teloschistaceae
Athallia holocarpa* River watershed Canada Maloles etal. (2018)
Athallia pyracea* River watershed Canada Maloles etal. (2018)
Blastenia ferruginea* River watershed Canada Maloles etal. (2018)
Caloplaca borreri* Nature reserve Ukraine Khodosovtsev etal. (2016)
Caloplaca flavovirescens* River Brazil Thüs etal. (2014)
Caloplaca obscurella* Stream UK Gilbert and Giavarini (1997)
Flavoplaca oasis* Stream UK Gilbert and Giavarini (1997)
Ostropomycetidae genera incertae sedis
Dictyocatenulata alba* River watershed Canada Maloles etal. (2018)
Gyalolechia flavovirescens* River watershed Canada Maloles etal. (2018)
Polycauliona polycarpa* River watershed Canada Maloles etal. (2018)
Rufoplaca arenaria* River watershed Canada Maloles etal. (2018)
Xanthomendoza fallax* River watershed Canada Maloles etal. (2018)
Xanthoria elegans* River watershed Canada Maloles etal. (2018)
Xanthoria parietina* River watershed Canada Maloles etal. (2018)
Umbilicariales
Ophioparmaceae
Hypocenomyce scalaris* River watershed Canada Maloles etal. (2018)
Umbilicariaceae
Umbilicaria deusta* Stream UK Gilbert and Giavarini (1997)
LICHINOMYCETES
Lichinomycetidae
Lichinales
Lichinaceae
Ephebe brasiliensis* Submerged rocks (river, stream) Brazil, Uruguay Thüs etal. (2014), Gumboski etal.
(2019)
Ephebe lanata* Submerged rocks (river, stream) UK Pentecost (1977), Gilbert and Gia-
varini (1997), Hawksworth (2000)
Ephebe ocellata* Semi-inundated rocks Australia, South America Kantvilas (2018), Gumboski etal.
(2019)
Ephebe perspinulosa* Stream UK Gilbert and Giavarini (1997)
Jenmania goebelii* Submerged rocks (river, stream) Central & South America,
Venezuela
Thüs etal. (2014)
Jenmania osorioi* Submerged rocks (river, stream) Uruguay Thüs etal. (2014)
Lempholemma botryosum* Damp limestone, seepage tracks UK Gilbert (1996)
Lempholemma cladodes* Block bellow limestone (waterfalls) UK Gilbert (1996)
Lempholemma polyanthes* Damp limestone, seepage tracks UK Gilbert (1996), Reeb etal. (2004)

Fungal Diversity
1 3
Taxon Substrate Distribution References
Porocyphus coccodes* Stream UK Gilbert and Giavarini (1997)
Porocyphus kenmorensis* Stream UK Gilbert and Giavarini (1997)
Psorotichia schaereri* Submerged rocks (limestone river) UK Gilbert (1996)
Pterygiopsis australiensis* Submerged stone (river) Australia Thüs etal. (2014)
Pterygiopsis coracodiza* Submerged rocks (limestone river) UK Gilbert (1996)
Pterygiopsis lacustris* Submerged rocks (limestone river) UK Gilbert (1996)
Pterygiopsis sp. Stream UK Schultaz and Büdel (2002)
Pterygiopsis submersa* Granite covered by running water (stream) South Africa Henssen etal. (1985)
Pyrenocarpon cf. flotowianum* Submerged rocks (limestone river) UK Gilbert (1996)
Pyrenopsis furfurea* – Italy Nascimbene and Nimis (2006)
Pyrenopsis subareolata* Stream Italy, UK Pentecost (1977), Nascimbene and
Nimis (2006)
Thelignya lignyota* Moist rocks (lakes, river) USA McCune (2016)
Zahlbrucknerella calcarea* – Italy Nascimbene and Nimis (2006)
Peltulaceae
Peltula euploca* Drainage surface with periodical rain Morocco, South Africa Egea and Rowe (1988), Thüs etal.
(2014)
Peltula obscurans* Drainage surface with periodical rain Morocco Egea and Rowe (1988)
Peltula omphaliza* Drainage surface with periodical rain Morocco Egea and Rowe (1988)
SORDARIOMYCETES
Hypocreomycetidae
Coronophorales
Nitschkiaceae
Rhagadostoma lichenicola#Lake USA Hafellner (2014)
Hypocreales
Bionectriaceae
Ovicuculispora parmeliae* River watershed Canada Maloles etal. (2018)
Hypocreales genera incertae sedis
Illosporiopsis christiansenii#River watershed Canada Maloles etal. (2018)
Illosporium carneum* River watershed Canada Maloles etal. (2018)
Pezizomycotina genera incertae sedis
Psammina stipitata#Nature reserve Western Ukraine, UK Khodosovtsev etal. (2016)
Pyrenocollema strontianense* Stream UK Gilbert and Giavarini (1997)
Table 10 (continued)
Number sign symbol (#) indicate Lichenicolous taxa while asterisk sign (*) are lichens and lichens-like

Fungal Diversity
1 3
Table 11 Classification of freshwater Ascomycota incertae sedis with information on the type of habitats, substrates, and geographical distribution of fungal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
Ascomycota genera incertae sedis
Actinocladium rhodosporum Submerged decaying twig (stream), water-filled
treeholes Brazil, Hungary Barbosa etal. (2013b) Gönczöl and Révay (2003)
Actinocladium verruculosum Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Alatosessilispora bibrachiata Rainwater from leaves of Quercus mongolica var.
grosseserrata Japan Ando and Tubaki (1984b)
Angulospora aquatica Submerged decaying leaves, rainwater from build-
ing roofs, foam (stream) India, Poland, Venezuela Nilsson (1962b), Czeczuga and Orłowska (1997),
Rajashekhar and Kaveriappa (2003)
Anisospadicoides macroobovata Submerged bark Brazil da Silva and Gusmão (2015)
Anungitopsis speciosa Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2019)
Anungitopsis triseptata Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2019)
Arachnophora combuensis Submerged decaying wood Brazil Monteiro etal. (2014b)
Arachnophora longa Submerged decaying wood Australia Fryar and Hyde (2018)
Arborispora dolichovirga Foam (stream)stemflow and throughfall of tree
canopies Canada, India Marvanová and Bärlocher (2001), Sridhar and
Karamchand (2009)
Arborispora palma Rainwater: building roofs, draining from leaves of
Quercus, sediment (dam) India, Japan, Poland Ando and Kawamoto (1986), Czeczuga and
Orłowska (1997), Sudheep and Sridhar (2012)
Arborispora paupera Spume (river), submerged decaying leaves (stream),
stemflow and throughfall of tree canopies Canada, India Marvanová and Bärlocher (1989), Sridhar and Bär-
locher (1993), Sridhar and Karamchand (2009)
Ardhachandra aequilatera Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2017b), Fiuza etal. (2019)
Arthrotaeniolella aquatica Submerged decaying leaves and twigs (stream) Brazil Monteiro etal. (2017a)
Ascomauritiana lignicola Submerged decaying wood (stream) Mauritius Ranghoo and Hyde (1999)
Biflagellospora gracilis Submerged decaying wood of Dipterocarpus ala-
tus and Xylia dolabriformis test block (stream) Thailand Sivichai and Hywel-Jones (1999)
Biflagellospora japonica Submerged decaying wood of Xylia dolabriformis
test block (stream) Thailand Sivichai and Hywel-Jones (1999), Sivichai etal.
(2002)
Biflagellospora papillata Submerged decaying wood of Xylia dolabriformis
test block (stream) Thailand Sivichai and Hywel-Jones (1999)
Biflagellospora siamensis Submerged decaying wood of Xylia dolabriformis
test block (stream) Thailand Sivichai and Hywel-Jones (1999)
Blastoheterospora catenata Submerged decaying wood (river) Brazil Monteiro etal. (2017b)
Blodgettia indica Submerged decaying leaves, foam (stream) Brazil, India Rajashekhar and Kaveriappa (2003), Moreira and
Schoenlein-Crusius (2012)
Brachydesmiella anthostomelloidea Submerged decaying wood (stream) and petiole
(stream) Australia, Brazil Sivichai etal. (1998)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Brachydesmiella biseptata Submerged test block (Anisoptera oblonga) Thailand Sivichai etal. (1998)
Brachydesmiella caudata Submerged decaying wood (Machilus velutina)
and petiole (stream) Brazil, China, India, Thailand Sivichai etal. (1998), Barbosa etal. (2013b)
Brachydesmiella orientalis Submerged test block (Anisoptera oblonga), sub-
merged decaying wood (stream) China, Thailand Sivichai etal. (1998), Mongkol etal. (2009)
Brachydesmiella verrucosa Submerged wood Malaysia, Thailand Sivichai etal. (1998), Hyde (2007)
Brevicatenospora enteroproliferata Submerged decaying leaves of Andira inermis
(stream) Cuba Castañeda-Ruiz etal. (2006)
Cacumisporium capitulatum Water-filled treeholes Hungary Gönczöl and Révay (2003)
Cacumisporium pleuroconidiopho-
rum Submerged decaying twig and bark (stream) Brazil Barbosa etal. (2013b)
Cacumisporium rugosum Submerged decaying wood (river) China Tsui etal. (2001b)
Cacumisporium sigmoideum Submerged decaying twig (stream) Brazil Barbosa etal. (2013b)
Cacumisporium uniseptatum Submerged decaying wood (stream) China Wongsawas etal. (2009a)
Calcarispora hiemalis Foam (stream), submerged decaying leaves of
Fagus sylvatica Czechoslovakia, France Gessner etal. (1993), Marvanová and Bärlocher
(2001)
Camposporidium cristatum Submerged decaying angiospermous leaves
(stream) Brazil, Malaysia Nawawi and Kuthubutheen (1988c), Barbosa etal.
(2013b)
Camposporidium hughesii Submerged decaying bark (stream) Cuba Castaneda Ruiz and Guarro (1998)
Candelabrum clathrosphaeroides Submerged decaying wood (stream), submerged
decaying leaves of Fagus sylvatica (peat hole) Austria, Thailand Voglmayr (1998), Chuaseeharonnachai etal. (2013)
Candelabrum desmidiaceum Submerged decaying leaves of Fagus sylvatica
(peat hole) Austria Voglmayr (1998)
Candelabrum macrosporum Submerged decaying leaves of broad-leaved tree
(stream) Japan Matsushima (1996)
Candelosynnema ranunculosporum Submerged decaying wood (waterfall) Australia Hyde and Seifert (1992)
Casaresia sphagnorum Foam (lake, spring, stream, river), submerged
decaying leaves of Quercus and angiosperm
(lake), water (stream)
Canada, France, Ireland, USA Petersen (1963a), Crane (1968), Abdullah and Web-
ster (1980), Bärlocher (1987), Gulis and Suberk-
ropp (2003), Cornut etal. (2014)
Ceuthospora gaeumannii Submerged decaying wood (stream) Brunei Ho etal. (2001)
Chaetendophragmia triangularis Foam (stream) India, Malaysia Nawawi (1985b), Rajashekhar and Kaveriappa
(2003)
Chaetopsis probosciophora Submerged decaying wood (reservoir) China Goh and Hyde (1999)
Cheiromyces lignicola Submerged decaying wood (river) China Ho etal. (2000)
Cladoconidium articulatum Foam (stream) Canada Marvanová and Bärlocher (2001)
Clavariana aquatica Foam (stream), rainwater from building roofs.
Sediment (stream) Malaysia, India, Poland Descals etal. (1976), Czeczuga and Orłowska
(1997), Sudheep and Sridhar (2012)
Colispora cavincola Submerged leaves, twigs, and cupules of Fagus
sylvatica in wet tree hollow Hungary Gönczöl and Révay (1996), Gönczöl and Révay
(2003)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Colispora elongata Rainwater from building roofs, water (stream),
foam (rivulet) Austria, Poland, Slovakia Marvanová (1988), Voglmayr (1996), Czeczuga and
Orłowska (1997)
Colispora curvata Submerged decaying angiospermic petioles
(stream) Malaysia Nawawi and Kuthubutheen (1989b)
Condylospora flexuosa Submerged decaying twig (stream), foam (stream) Brazil, Malaysia, Venezuela Nawawi and Kuthubutheen (1988d), Smits etal.
(2007), Fiuza etal. (2015a)
Condylospora gigantea Submerged decaying twig and leaves of Calophyl-
lum brasiliense (stream), foam (stream), Brazil, Malaysia Nawawi and Kuthubutheen (1988c), Hosoya etal.
(2006), Fiuza and Gusmão (2013a), Fiuza etal.
(2017a)
Condylospora spumigena Foam (stream, river), submerged decaying twigs,
throughfall of palm species Brazil, India, Japan, Malaysia, Thailand Nawawi (1976c), Nawawi and Kuthubutheen
(1988c), Sridhar and Kaveriappa (1992), Saka-
yaroj etal. (2005), Hosoya and Tanaka (2007),
Fiuza etal. (2015a), Ghate and Sridhar (2015)
Conioscyphopsis australiensis Submerged decaying of angiospermous wood Australia Goh and Hyde (1998)
Corynesporella superioramifera Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Corynesporopsis antillana Submerged twig, submerged decaying leaves of
Calophyllum brasiliense (stream) Brazil da Silva and Gusmão (2015), Fiuza etal. (2019)
Culicidospora aquatica Foam (lake, spring, stream, rivulet, river), sub-
merged decaying leaves (stream), submerged leaf
litter of Alnus glutinosa (stream), scum (stream),
water (stream)
Austria, Canada, France, Malaysia, USA Ingold (1960), Petersen (1963a), Crane (1968), Bär-
locher (1987), Gessner etal. (1993), Sridhar and
Bärlocher (1993), Voglmayr (1996), Marvanová
(2000), Hosoya etal. (2006), Cornut etal. (2014)
Culicidospora gravida Foam (river, stream, rivulet), submerged decaying
leaves (river, stream), submerged leaf litter of
Alnus glutinosa (stream)
Austria, Brazil, Canada, France, Japan, USA,
Venezuela Petersen (1963a), Bärlocher (1987), Gessner etal.
(1993), Sridhar and Bärlocher (1993), Voglmayr
(1996), Marvanová (2000), Hosoya and Tanaka
(2007), Smits etal. (2007), Fiuza and Gusmão
(2013a)
Curucispora ombrogena Rainwater from leaves of Phyllostachys bambu-
soides Japan Ando and Tubaki (1984b)
Curucispora ponapensis Stemflow and throughfall of tree canopies India Sridhar and Karamchand (2009)
Delortia palmicola Submerged decaying Licuala longicalycata and
Eleiodoxa conferta (peat swamp) Thailand Pinnoi etal. (2006), Pinruan etal. (2007)
Dendrospora erecta Foam (lake, stream, spring, rivulet, river),
submerged decaying leaves of Alnus glutinosa
(stream), scum (stream), water (stream)
Austria, Brazil, Canada, France, India, Ireland,
UK, USA Ingold (1943a), Ingold (1960), Petersen (1962),
Crane (1968), Bärlocher (1987), Sridhar and
Kaveriappa (1992), Gessner etal. (1993), Vogl-
mayr (1996), Harrington (1997), Marvanová
(2000), Gulis and Suberkropp (2003), Schoenlein-
Crusius etal. (2009), Cornut etal. (2014)
Dendrospora fastuosa Foam (river) Canada, UK Descals and Webster (1980), Bärlocher (1987)
Dendrospora fusca Foam (river), submerged leaf litter of Alnus gluti-
nosa (stream) France, Ireland, UK Descals and Webster (1980), Gessner etal. (1993),
Harrington (1997)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Dendrospora juncicola Submerged decaying culms of Juncus effusus
(river), foam (stream), stemflow India, UK Iqbal (1972a), Sridhar and Kaveriappa (1992),
Ghate and Sridhar (2015)
Dendrospora moniliformis Submerged decaying wood (Fagus sylvatica, Picea
abies), foam (stream, rivulet), water (stream) Austria, Canada, UK Descals and Webster (1983), Bärlocher (1987),
Voglmayr (1996), Marvanová (2000)
Dendrospora nana Spume (river), foam (river), submerged decaying
leaves (stream), stemflow Canada, India, UK Descals and Webster (1980), Bärlocher (1987),
Sridhar and Bärlocher (1993), Ghate and Sridhar
(2015)
Dendrospora polymorpha Foam (river, stream) Austria, Spain Roldán etal. (1987), Marvanová (2000)
Dendrospora tenella Foam (stream, rivulet), submerged decaying twigs
(river) Austria, Canada, Portugal, UK Descals and Webster (1980), Bärlocher (1987),
Voglmayr (1996), Marvanová (2000), Duarte etal.
(2015)
Dendrospora torulosa Submerged rotting cupules of Castanea (river),
stemflow India, UK Descals and Webster (1980), Ghate and Sridhar
(2015)
Dendrospora yessemreddyi Foam (stream) India Sreekala and Bhatt (2002)
Dendrosporium lobatum Submerged decaying Xylia dolabriformis test
block, foam (river, stream), submerged decaying
leaves of Calophyllum brasiliense (stream)
Brazil, India, Thailand Sivichai etal. (2002), Rajashekhar and Kaveriappa
(2003), Fiuza and Gusmão (2013a), Fiuza etal.
(2019)
Dendryphiosphaera parvula Submerged decaying twig (stream) Malaysia Nawawi and Kuthubutheen (1988e)
Denticularia limoniformis Submerged decaying wood (stream) Brunei Ho etal. (2001)
Dictyoaquaphila appendiculata Submerged decaying wood (river, stream) Brazil Monteiro etal. (2016b)
Dictyotrichocladium aquaticum Submerged decaying leaves of Calophyllum Bra-
siliense (river) Brazil Fiuza etal. (2017c)
Diplocladiella appendiculata Foam (river), submerged decaying wood Malaysia, Thailand Nawawi (1987b), Sakayaroj etal. (2005)
Diplocladiella aquatica Submerged decaying wood Brunei Lee etal. (1998)
Diplocladiella scalaroides Foam (stream, river, rivulet), water-filled treeholes Austria, Hungary, India, Malaysia, Thailand,
Venezuela Voglmayr (1996), Marvanová (2000), Gönczöl
and Révay (2003), Rajashekhar and Kaveriappa
(2003), Sakayaroj etal. (2005), Hosoya etal.
(2006), Smits etal. (2007)
Diplocladiella tricladioides Submerged decaying wood Malaysia Nawawi (1985d)
Drepanospora viridis Stemflow and throughfall of palm species India Ghate and Sridhar (2015)
Dwayaangam colodena Foam (stream) Canada Sokolski etal. (2006)
Dwayaangam cornuta Foam (stream, rivulet), stemflow and throughfall
of tree canopies and palm species, water-filled
treeholes
Austria, Brazil, Canada, Hungary, India, UK Descals and Webster (1982), Bärlocher (1987),
Voglmayr (1996), Marvanová (2000), Gönczöl and
Révay (2003), Sridhar and Karamchand (2009),
Fiuza etal. (2015a), Ghate and Sridhar (2015)
Dwayaangam dichotoma Spume (river), stemflow and throughfall of tree
canopies India, Malaysia Nawawi (1985d), Sridhar and Karamchand (2009)
Elegantimyces sporidesmiopsis Submerged decaying wood China Goh etal. (1998e)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Enantioptera bialata Foam (stream) Czech Republic Marvanová (2004)
Enantioptera tetra-alata Foam (river, stream) Ireland, UK Descals and Webster (1983), Harrington (1997)
Endophragmia boewei Submerged decaying plant material (swamp) USA Crane (1972)
Everhartia aquatica Submerged decaying wood (stream) Australia Goh and Hyde (1997b)
Everhartia tumidiapicis Submerged decaying wood (stream) Australia Goh and Hyde (1997b)
Flabellocladia gigantea Foam (stream) Malaysia Nawawi (1985e)
Flabellospora acuminata Foam (river), submerged leaf litter of Alnus gluti-
nosa (stream) France, UK, Venezuela Descals and Webster (1982), Gessner etal. (1993),
Smits etal. (2007)
Flabellospora amphibia Dead insects (pond), foam (stream), submerged
decaying wood Portugal, Uzbekistan, UK Descals and Webster (1982), Sagdullaeva etal.
(1989), Marvanová etal. (2003)
Flabellospora crassa Submerged decaying leaves of Phoenix dactyl-
ifera, Coffea arabica and Hevea brasiliensis,
stemflow and throughfall, foam (stream, river)
Brazil, India, Nigeria, Thailand, USA, Ven-
ezuela Alasoadura (1968b), Ranzoni (1979), Sridhar
and Kaveriappa (1988), Sridhar and Kaveriappa
(1992), Rajashekhar and Kaveriappa (2003), Saka-
yaroj etal. (2005), Smits etal. (2007), Schoenlein-
Crusius etal. (2009), Sridhar and Karamchand
(2009), Ghate and Sridhar (2015)
Flabellospora multiradiata Foam (river, stream), stemflow and throughfall of
tree canopies India, Malaysia, Thailand Nawawi (1976b), Sridhar and Kaveriappa (1992),
Sakayaroj etal. (2005), Sridhar and Karamchand
(2009)
Flabellospora tetracladia Foam (stream) Brazil, India, Japan, Malaysia, Venezuela Nawawi (1985e), Sridhar and Kaveriappa (1992),
Hosoya and Tanaka (2007), Smits etal. (2007),
Fiuza and Gusmão (2013a)
Flabellospora verticillata Submerged decaying leaves of Coffea arabica and
Phoenix dactylifera, stemflow and throughfall
of tree canopies and palm species, foam (river,
stream)
Brazil, India, Nigeria, Thailand, Venezuela Alasoadura (1968b), Sridhar and Kaveriappa
(1988), Sridhar and Kaveriappa (1992), Sakayaroj
etal. (2005), Smits etal. (2007), Sridhar and
Karamchand (2009), Fiuza and Gusmão (2013a),
Ghate and Sridhar (2015)
Frigidispora colnensis Submerged decaying wood (river) UK Hyde and Goh (1999a)
Fusticeps laevisporus Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2019)
Gemmulina botryosa Foam (stream) UK Descals and Webster (1982)
Helicoubisia coronata Submerged decaying angiospermous twig, decay-
ing Eleiodoxa conferta (peat swamp) Malaysia, Thailand Kuthubutheen and Nawawi (1994b), Pinnoi etal.
(2006)
Goidanichiella fusiformis Submerged decaying Eleiodoxa conferta (peat
swamp) Thailand Pinnoi etal. (2006)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Heliscella stellata Foam (stream, rivulet), submerged decaying
leaves of Alnus glutinosa (stream), stemflow
and throughfall of tree canopies and pal species,
water (stream)
Austria, France, India, Ireland, Malaysia,
Spain, Sweden, UK, USA Ingold and Cox (1957), Willen (1958), Descals and
Webster (1982), Nawawi (1985b), Gessner etal.
(1993), Voglmayr (1996), Harrington (1997),
Marvanová (2000), Gulis and Suberkropp (2003),
Sridhar and Karamchand (2009), Menéndez etal.
(2012), Cornut etal. (2014), Ghate and Sridhar
(2015)
Heliscella stellatacula Foam (rivulet) Austria, Malaysia Nawawi (1985b), Voglmayr (1996)
Heliscina antennata Decaying leaf litter of Alnus glutinosa (stream),
foam, (stream), spring, water (stream) Austria, Czech Republic, France, Spain, USA Marvanová (1980), Gessner etal. (1993), Descals
(1997), Marvanová (2000), Gulis and Suberkropp
(2003)
Heliscina campanulata Foam (river, stream), submerged leaf litter of
Alnus glutinosa (stream), water (stream) Austria, Canada, France, India, Ireland, USA Descals and Webster (1982), Bärlocher (1987),
Gessner etal. (1993), Harrington (1997),
Marvanová (2000), Gulis and Suberkropp (2003),
Rajashekhar and Kaveriappa (2003), Cornut etal.
(2014)
Henicospora amazonensis Submerged decaying leaves (stream) Brazil Monteiro etal. (2014a)
Henicospora coronata Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2019)
Henicospora longissima Submerged decaying litter of Saraca thaipingensis Malaysia Kuthubutheen and Nawawi (1994b)
Hormiactis ontariensis Water-filled treeholes Hungary Gönczöl and Révay (2003)
Hyalocamposporium acutum Submerged decaying wood (stream) Hungary Révay and Gönczöl (2007)
Hyalocamposporium longiflagel-
latum Submerged decaying wood (stream) China, Hungary Révay and Gönczöl (2007), Hu etal. (2013b)
Hydrometrospora symmetrica Foam/water (stream), stemflow India, Venezuela Smits etal. (2007), Ghate and Sridhar (2015)
Inesiosporium longispirale Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b), Fiuza etal. (2017b)
Ityorhoptrum verruculosum Submerged decaying bamboo (stream, river,
swamp) Brazil, Philippines, USA Crane (1972), Cai etal. (2003a) Barbosa etal.
(2013b)
Isthmolongispora asymmetrica Decaying leaves (river) Argentina Arambarri etal. (1987)
Isthmolongispora geniculata Submerged decaying angiospermic leaves Malaysia Nawawi and Kuthubutheen (1988a)
Isthmolongispora lanceata Water Netherlands Hoog and Hennebert (1983)
Isthmolongispora minima Foam (river), water-filled treeholes Hungary, Ireland Harrington (1997) Gönczöl and Révay (2003)
Isthmolongispora rotundata Submerged decaying leaves China Matsushima (1987)
Isthmophragmospora laevispora Submerged decaying leaves (stream) Brazil Monteiro etal. (2014a)
Isthmophragmospora verruculosa Submerged decaying angiospermous twigs
(stream) Malaysia Kuthubutheen and Nawawi (1992)
Isthmotricladia britannica Foam (stream, river rivulet) Austria, Canada, Ireland, UK Descals and Webster (1982), Bärlocher (1987),
Voglmayr (1996), Harrington (1997), Marvanová
(2000)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Isthmotricladia gombakiensis Foam (stream, river), stemflow and throughfall India, Malaysia, Thailand, Venezuela Nawawi (1975), Sridhar and Kaveriappa (1992),
Sakayaroj etal. (2005), Smits etal. (2007), Ghate
and Sridhar (2015)
Isthmotricladia laeensis Stemflow and throughfall of tree canopies and
palm species, foam (stream, river) India, Malaysia, Thailand Nawawi (1985b), Sridhar and Kaveriappa (1992),
Sakayaroj etal. (2005), Sridhar and Karamchand
(2009)
Iyengarina asymmetrica Submerged decaying angiospermous twigs
(stream) Malaysia Kuthubutheen and Nawawi (1992)
Iyengarina furcata Submerged decaying twig (stream) Malaysia Kuthubutheen and Nawawi (1992)
Jerainum triquetrum Submerged decaying twig (stream) Malaysia Nawawi and Kuthubutheen (1992)
Kameshwaromyces globosus Submerged decaying wood (stream) China Ho etal. (2001)
Kiliophora fusispora Submerged decaying angiospermous leaves Malaysia Kuthubutheen and Nawawi (1993b)
Kiliophora ubiensis Submerged decaying leaves Malaysia Kuthubutheen and Nawawi (1993b)
Kontospora halophila Submerged leaves of Populus euphratica (stream) Spain Roldán etal. (1990)
Kostermansinda magna Submerged decaying culm of Miscanthus floridu-
lus (stream) China Kuo and Goh (2019)
Kostermansinda minima Decaying bark (river) Argentina Arambarri etal. (1987)
Lambdasporium viridense Spume (river) Malaysia Nawawi (1985d)
Lateriramulosa quadriradiata Foam (stream), stemflow and throughfall of tree
canopies Canada, India Marvanová and Bärlocher (2001), Sridhar and
Karamchand (2009)
Lateriramulosa uniinflata Foam (stream, river, rivulet), sediment (dam,
river), stemflow and throughfall of tree canopies,
water (stream)
Austria, Canada, France, India, Ireland, Japan,
Malaysia, USA Bärlocher (1987), Voglmayr (1996), Harrington
(1997), Gulis and Suberkropp (2003), Hosoya
etal. (2006), Hosoya and Tanaka (2007), Sridhar
and Karamchand (2009), Sudheep and Sridhar
(2012), Cornut etal. (2014)
Linkosia aquatica Submerged decaying twig (stream) Brazil Conceição etal. (2016)
Listeromyces insignis Submerged decaying wood (stream) China Ho etal. (2001)
Magdalaenaea monogramma Foam (stream) Canada Bärlocher (1987)
Megaloseptoria mirabilis Submerged decaying wood (stream) Brunei Ho etal. (2001)
Melanocephala australiensis Submerged decayingwood (river, stream) Brazil. China, USA Shearer and Crane (1971), Ho etal. (2001), Barbosa
etal. (2013b)
Melanocephala triseptata Submerged decaying wood USA Shearer etal. (1976)
Mirandina corticola Rainwater from building roofs, submerged decay-
ing woods (river), water-filled treeholes Brazil, Hungary, Poland Czeczuga and Orłowska (1997), Gönczöl and Révay
(2003), Barbosa etal. (2013b)
Mirandina taiwanensis Submerged decaying wood China Matsushima (1987)
Mirandina uncinata Submerged decaying leaves of Calophyllum
brasiliense Brazil Fiuza etal. (2016)
Neopodoconis ampullacea Submerged decaying wood (river, stream) Brunei, Philippines Ho etal. (2001), Cai etal. (2003a)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Neta angliae Submerged decaying wood (river) UK Hyde and Goh (1999a)
Neta lignicola Submerged decaying wood (river) USA Shearer (1974)
Neta mexicana Submerged decaying leaves (stream) Mexico Castañeda-Ruiz etal. (2005)
Neta patuxentica Submerged balsa wood (river), rainwater from
building roofs Poland, USA Shearer and Crane (1971), Czeczuga and Orłowska
(1997)
Neta salmonicolor Submerged decaying balsa wood (river) USA Shearer (1974)
Nidulispora quadrifida Submerged decaying wood (river) Brazil, Malaysia Nawawi and Kuthubutheen (1990b), Hyde (2007),
Fiuza etal. (2017b)
Obeliospora basispira Submerged decaying twigs (stream) Malaysia Nawawi and Kuthubutheen (1990c)
Obeliospora microappendiculata Submerged decaying twigs (river, stream) Brazil Cantillo-Pérez etal. (2018)
Obeliospora nitida Submerged decaying angiospermous twigs
(stream) Malaysia Kuthubutheen etal. (1992), Cantillo-Pérez etal.
(2018)
Obeliospora triappendiculata Submerged decaying wood, foam (stream) Brazil, Malaysia Kuthubutheen and Nawawi (1994b)
Obstipispora chewaclensis Submerged decaying leaves of Populus deltoides
(stream) USA Sinclair and Morgan-Jones (1979b)
Ordus tribrachiatus Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Pachycladina hispanica Foam (stream) Spain Roldán etal. (1990)
Pachycladina mutabilis Foam (stream) Czech Republic Marvanová (1986)
Pachycladina parva Foam (stream) Canada Marvanová and Bärlocher (1998c)
Paraceratocladium malaysianum Submerged decaying wood (river) Malaysia Goh and Hyde (2000a)
Hyde (2007)
Paracryptophiale kamaruddinii Submerged decaying twigs (stream) Malaysia Kuthubutheen and Nawawi (1994c)
Paraulocladium fabisporum Submerged decaying angiospermous wood Malaysia Kuthubutheen and Nawawi (1994b)
Phaeomonilia aquatica Submerged decaying wood (stream) China Huang etal. (2017)
Phaeomonilia pleiomorpha Submerged decaying palm petioles (stream) Mexico Castañeda-Ruíz etal. (2007)
Phaeotrichoconis aquatica Decaying branch (river) Argentina Arambarri etal. (1987)
Phaeotrichoconis minigelatinosa Decaying branch (river) Argentina Arambarri etal. (1987)
Pleuropedium macrum Foam (stream) Canada Marvanová and Bärlocher (1998c)
Pleuropedium multiseptatum Submerged decaying Juncus effusus (stream) UK Marvanová and Descals (1996)
Pleuropedium tricladioides Submerged rotten culms of Juncus effusus, foam
(stream, rivulet), water (stream) Austria, Canada, UK, USA Marvanová and Iqbal (1973), Bärlocher (1987),
Voglmayr (1996), Gulis and Suberkropp (2003)
Polycladium equiseti Submerged dead inter-nodes of Equisetum fluvia-
tile UK Ingold (1959b)
Porocladium aquaticum Foam (river) UK Descals etal. (1976)
Pseudoanguillospora gracilis Submerged decaying leaves of Platanus occiden-
talis (stream), foam (stream) USA Sinclair and Morgan-Jones (1979a)
Pseudoanguillospora prolifera Submerged decaying leaves of Poaceae and culms
of Juncus effusus USA Iqbal (1974)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Pseudoanguillospora stricta Submerged decaying leaves of Poaceae and culms
of Juncus effusus USA Iqbal (1974)
Pseudoclathrosphaerina evamariae Submerged decaying wood (stream) and leaves
(creek, water-filled channel, lake) Thailand, USA Voglmayr and Krisai-Greilhuber (1997a), Chuasee-
haronnachai etal. (2013)
Pseudoclathrosphaerina spiralis Submerged decaying wood (stream) Thailand Chuaseeharonnachai etal. (2013)
Pseudofuscophialis lignicola Submerged decaying wood (stream) China Cai and Hyde (2007a)
Pseudotracylla dentata Submerged decaying wood and leaves (stream) Brazil Barbosa etal. (2013b)
Pyramidospora casuarinae Submerged decaying leaves of Calophyllum
brasiliense and branches of Casuarina, foam
(stream),
Brazil, Cuba, India, USA, Venezuela Marvanová and Marvan (1969), Ranzoni (1979),
Sridhar and Kaveriappa (1992), Schoenlein-Cru-
sius etal. (2009), Fiuza etal. (2017a)
Pyramidospora constricta Submerged decaying leaves of Dicotyledonae
(stream), foam (stream) India, Sierra Leone Singh (1972), Sridhar and Kaveriappa (1992)
Pyramidospora densa Submerged decaying leaves (stream) and wood
(river) Nigeria Alasoadura (1968a)
Pyramidospora fluminea Submerged decaying leaves (stream) and wood
(river) Japan, Malaysia Hyde (2007)
Miura and Kudo (1971)
Pyramidospora herculiformis Submerged decaying leaves Sierra Leona Singh (1976)
Pyramidospora quadricellularis Submerged decaying leaves (stream) Brazil Oliveira etal. (2015a)
Pyramidospora ramificata Submerged decaying leaves (stream) Japan Miura and Kudo (1971)
Pyramidospora robusta Submerged decaying leaves of Caesalpinia echi-
nata and Campomanesia phaea (lake) Brazil Moreira and Schoenlein-Crusius (2012)
Pyramidospora stellata Submerged decaying leaves of Acer saccharinum,
Quercus alba, and Salix sp. USA Sinclair and Morgan-Jones (1979a)
Quadracaea stauroconidia Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Quadricladium aquaticum Submerged deciduous bark (stream) Malaysia Nawawi and Kuthubutheen (1989c)
Ramicephala sphaerospora Submerged decaying wood (river) Cuba Voglmayr and Delgado-Rodríguez (2003)
Satchmopsis brasiliensis Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Scutisporus brunneus Foam (river, stream), submerged decaying petioles
(stream), submerged decaying wood (e.g.,
Dipterocarpus alatus and Xylia dolabriformis
test blocks), submerged leaves of Calophyllum
brasiliense (stream)
Brazil, Malaysia, Thailand, Venezuela Sivichai etal. (2002), Smits etal. (2007), Fiuza and
Gusmão (2013a), Barbosa etal.( 2013b), Fiuza
etal. (2017a), Hosoya etal. (2019)
Septotrullula bacilligera Submerged decaying wood (river) UK Hyde and Goh (1999a)
Sirothecium triseriale Submerged decaying bamboo culms China Hu etal. (2007)
Solheimia costispora Decaying Licuala longicalycata (peat swamp) Thailand Pinruan etal. (2014)
Sphaerulomyces coralloides Parasitic on conidia of Anguillospora crassa from
foam (brook) Slovakia Marvanová (1977a)
Spiropes caaguazuensis Submerged decaying wood (river) Philippines Cai etal. (2003a)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Spirosphaera beverwijkiana Submerged decaying leaves of Quercus peduncu-
lata, Quercus sp., Q. coccinea, Fagus sylvatica,
Betula verrucosa, Populus tremula (lake, stag-
nant water)
Belgium, Ireland, Netherlands Hennebert (1968), Abdullah and Webster (1980)
Spirosphaera carici-graminis Submerged decaying leaves and culms of Calama-
grostis epigejos, Juncus effusus and Carex
brizides (puddle)s, Submerged decaying leaves
of Calophyllum brasiliense (stream)
Austria, Brazil Voglmayr (1997), Fiuza etal. (2018)
Spirosphaera dimorpha Foam (stream) Canada Marvanová and Bärlocher (1998b)
Spirosphaera floriformis Submerged decaying Betula leaves, submerged
decaying wood (river) Netherlands, Malaysia van Beverwijk (1953), Ho etal. (2001)
Spirosphaera lignicola Submerged decaying twigs (stream) Italy Abdullah etal. (1998)
Spirosphaera minuta Submerged decaying leaves of Betula alba, B.
verrucosa, Fagus sylvatica, Populus termula,
and Corylus avellana, Quercus pedunculata,
Quercus sp.
Belgium, Netherlands Hennebert (1968)
Spirosphaera verruculosa Submerged decaying leaves (pond) Japan Abdullah etal. (1986)
Spondylocladiopsis cupulicola Water-filled treeholes Hungary Gönczöl and Révay (2003)
Subulispora biappendiculata Submerged decaying leaves (stream) Malaysia Marvanová and Laichmanová (2007)
Subulispora longirostrata Submerged decaying angiospermous leaves and
petiole (stream) Brazil, Malaysia Nawawi and Kuthubutheen (1987c), Barbosa etal.
(2013b)
Subulispora malaysiana Submerged decaying leaves (stream), foam
(stream) Malaysia Nawawi and Kuthubutheen (1990a), Hosoya etal.
(2019)
Subulispora procurvata Submerged decaying petioles, foam (stream),
Submerged decaying leaves of Calophyllum
brasiliense (stream)
Brazil, India Barbosa etal. (2013b) Rajashekhar and Kaveriappa
(2003), Fiuza etal. (2018)
Stenocladiella neglecta Foam (stream, rivulet), submerged decaying leaves
of Alnus glutinosa, Alnus sp. (stream), water
(stream)
Austria, France, Slovakia, Spain Descals and Webster (1982), Marvanová and
Descals (1985), Gessner etal. (1993), Voglmayr
(1996), Marvanová (2000), Menéndez etal.
(2012), Cornut etal. (2014)
Sympodiella goidanichii Water-filled treeholes Hungary Gönczöl and Révay (2003)
Sympodiocladium frondosum Foam (stream) Canada, UK Descals and Webster (1982), Marvanová and Bär-
locher (2001)
Synnematophora constricta Sediment (stream), submerged decaying leaves of
Mangifera indica (river), stemflow and through-
fall of tree canopies
India Sridhar and Kaveriappa (2002), Sridhar and Karam-
chand (2009), Sudheep and Sridhar (2012)
Synchaetomella aquatica Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2015b)
Tetrabrachium elegans Submerged decaying leaves (stream), submerged
decaying angiosperm submerged (stream) Canada, Malaysia Nawawi and Kuthubutheen (1987a), Sridhar and
Bärlocher (1993)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Tetranacrium gramineum Submerged decaying wood (river) USA Shearer and Crane (1971)
Tetrabrunneospora ellisii Foam (stream), submerged decaying Betula leaves
(brook) Canada, USA Dyko (1978), Marvanová and Bärlocher (2001)
Titaea callispora Foam (stream) Malaysia Nawawi (1985b)
Titaea clarkeae Rainwater from building roofs, stemflow and
throughfall of tree canopies India, Poland Czeczuga and Orłowska (1997), Sridhar and Kara-
mchand (2009)
Triramulispora gracilis Foam (rivulet) Austria Voglmayr (1996)
Triramulispora obclavata Foam (rivulet) Austria Voglmayr (1996)
Triscelosporium verrucosum Foam (stream), submerged decaying angiospermic
leaves (stream) Malaysia Nawawi and Kuthubutheen (1987d), Hosoya etal.
(2019)
Tricellula aquatica Submerged leaves of Alnus glutinosa, Castanea
sativa, Hedera helix (river), foam (stream),
rainwater from building roofs
Austria, Poland, UK Marvanová (2000)
Webster (1959), Czeczuga and Orłowska (1997)
Tricellula inaequalis Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Tricellula ornata Foam (stream) Czech Republic Marvanová (2004)
Tricladiopsis flagelliformis Foam (stream), submerged decaying leaves of
Quercus (river) Portugal, UK Descals and Webster (1982), Marvanová etal.
(2003)
Tricladiopsis foliosa Foam (stream) UK Descals and Webster (1982)
Tricladiospora longissima Submerged decaying angiospermic leaves (stream) Malaysia Nawawi and Kuthubutheen (1988f)
Tricladiospora stricta Submerged decaying angiospermic leaves (stream) Malaysia Nawawi and Kuthubutheen (1988e)
Trifurcospora irregularis Foam (stream), stemflow and throughfall Canada, India Marvanová and Bärlocher (2001), Ghate and Srid-
har (2015)
Triglyphium alabamense Foam (stream) Canada Marvanová and Bärlocher (2001)
Triscelophorus acuminatus Foam (stream, river), submerged leaves of Cin-
namomum zeylanicum, Coffea arabica, and
Hevea brasiliensis, unindentified leaves (stream,
lake), sediment (dam, stream), stemflow and
throughfall of tree canopies and palm species.
Submerged decaying leaves of Calophyllum
brasiliense (stream)
Austria, Brazil, Canada, India, Ireland, Italy,
Malaysia, Portugal, Spain, Thailand, Ven-
ezuela
Descals and Webster (1982), Nawawi (1985b),
Sridhar (1987), Sridhar and Bärlocher (1993),
Harrington (1997), Marvanová (2000), Rajashek-
har and Kaveriappa (2003), Sivichai and Boony-
ene (2004), Sakayaroj etal. (2005), Smits etal.
(2007), Sridhar and Karamchand (2009), Moreira
and Schoenlein-Crusius (2012), Sudheep and
Sridhar (2012), Ghate and Sridhar (2015), Duarte
etal. (2015), Fiuza etal. (2018), Raposeiro etal.
(2018), Hosoya etal. (2019)
Triscelophorus curviramifer Foam/water (stream) Brazil, Venezuela Fiuza etal. (2015a) Smits etal. (2007)
Triscelophorus konajensis Submerged rotten leaves of Ficus benghalensis,
sediment (dam, stream), stemflow and through-
fall of tree canopies and palm species, foam
(stream), water (stream)
India, USA Sridhar and Kaveriappa (1987), Sridhar and
Kaveriappa (1992), Gulis and Suberkropp (2003),
Sridhar and Karamchand (2009), Sudheep and
Sridhar (2012), Ghate and Sridhar (2015)

Fungal Diversity
1 3
Table 11 (continued)
Taxon Substrate Distribution References
Triscelophorus magnificus Submerged leaves (stream), foam (stream) Canada, USA Petersen (1962), Bärlocher (1987)
Triscelophorus monosporus Submerged decaying leaves (river), submerged
leaves of Coffea arabica, and Hevea, water
(river), foam, scum (stream, spring, river, lake),
sediment (dam, stream), stemflow and through-
fall of tree canopies, submerged leaves of Calo-
phyllum brasiliense (stream), water (stream)
Australia, Brazil, Canada, Cuba, France,
Ghana, India, Ireland, Jamaica, Malaysia,
Nigeria, Poland, Portugal, UK, USA, Ven-
ezuela
Ingold (1943a), Ingold (1943b), Ranzoni (1953),
Dixon (1959), Hudson and Ingold (1960)ngold
(1960, Petersen (1962), Petersen (1963b), Crane
(1968), Alasoadura (1968a), Marvanová and
Marvan (1969), Ingold and Webster (1973),
Nawawi (1985b), Bärlocher (1987), Sridhar and
Kaveriappa (1988), Czeczuga etal. (1990), Har-
rington (1997), Gulis and Suberkropp (2003),
Smits etal. (2007), Schoenlein-Crusius etal.
(2009), Sridhar and Karamchand (2009), Sudheep
and Sridhar (2012), Cornut etal. (2014), Duarte
etal. (2015), Fiuza etal. (2017a), Raposeiro etal.
(2018)
Triscelophorus ponapensis Submerged decaying leaves of Pandanus Micronesia, Thailand Matsushima (1981), Sivichai and Boonyene (2004)
Triscelophorus septatus Submerged decaying leaves USA Parker and Roane (1976)
Trisulcosporium acerinum Stemflow and throughfall of tree canopies, foam
(river, stream) Brazil, India, Japan, Malaysia Hosoya etal. (2006), Hosoya and Tanaka (2007),
Sridhar and Karamchand (2009), Fiuza and
Gusmão (2013a)
Uberispora heteroseptata Submerged decaying wood (river) Malaysia Ho etal. (2001), Hyde (2007)
Umbellidion radulans Submerged decaying leaves (stream) Brazil Barbosa etal. (2013b)
Vanbeverwijkia spirospora Submerged decaying wood (river) USA Shearer and Crane (1971)
Veramyces manuensis Rainwater from building roofs Poland Czeczuga and Orłowska (1997)
Verticicladus subiculifer Submerged decaying leaves of Calophyllum bra-
siliense (stream) Brazil Fiuza etal. (2015b), Fiuza etal. (2017b)
Virgariella atra Submerged decaying wood (stream) Brazil, Brunei Ho etal. (2001), Barbosa etal. (2013b)
Virgariella globigera Submerged decaying wood (stream) Brazil Barbosa etal. (2013b)
Xiuguozhangia appendiculata Submerged decaying wood Thailand Sivichai and Boonyene (2004)
Xylohyphopsis aquatica Submerged decaying wood (stream) China Huang etal. (2016)
Yinmingella mitriformis Submerged decaying wood (river) China Goh etal. (1998f)
Zelotetraploa aquatica Submerged decaying wood (river) Brazil Monteiro etal. (2017b)

Fungal Diversity
1 3
Table 12 Classification of freshwater Basidiomycota with information on the type of habitats, substrates, and geographical distribution of fungal
taxa
Taxon Substrate Distribution References
AGARICOMYCOTINA
AGARICOMYCETES
Agaricales
Agaricales genera incertae sedis
Mycocalia denudata Decaying Eleiodoxa conferta (peat
swamp) Thailand Pinruan etal. (2014)
Mycocalia reticulata Submerged wood Australia Hyde and Goh (1998a)
Limnoperdaceae
Limnoperdon incarnatum Soaked hardwood twigs, marsh water USA Escobar etal. (1976)
Limnoperdon sp. Submerged wood and date palm
rachis (river) Egypt Abdel-Aziz (2016b)
Niaceae
Flagelloscypha sp. Submerged decaying wood (stream) Japan Yamaguchi etal. (2009)
Peyronelina glomerulata Submerged decaying Cyperus sp.
culm, unidentified pod, twig,
litter (pond, stream, waterfall)
submerged pinnules of Pteridium
aquilinum
China, Japan, Thailand, UK Fisher etal. (1976), Mongkol etal.
(2009), Yamaguchi etal. (2009),
Chuaseeharonnachai etal. (2013)
Physalacriaceae
Gloiocephala aquatica Decaying culms of Scirpus californi-
cus (lakes, eutrophic ponds) Argentina Desjardin etal. (1995)
Psathyrella aquatica Submerged wood or sediment USA Frank etal. (2010)
Atheliales
Atheliaceae
Leptosporomyces crucelliger Foam (stream) UK Marvanová and Stalpers (1987)
Leptosporomyces galzinii Foam, submerged decaying Acer
leaves (stream), water UK Nawawi etal. (1977a)
Taeniospora gracilis Foam (river, rivulet, stream), sub-
merged decaying leaves (stream),
water (stream)
Austria, Canada, France, UK, USA Marvanová and Stalpers (1987),
Bärlocher (1987), Sridhar and
Bärlocher (1993), Voglmayr
(1996), Marvanová (2000), Gulis
and Suberkropp (2003), Cornut
etal. (2014)
Taeniospora gracilis var.
gracilis Foam (stream), submerged leaf litter
of Alnus glutinosa (stream), water Austria, France, UK Marvanová and Stalpers (1987),
Gessner etal. (1993), Marvanová
(2000)
Taeniospora nasifera Foam (river) Canada Marvanová and Bärlocher (1988)
Cantharellales
Ceratobasidiaceae
Ceratobasidium sp. Submerged plant root (lake) Norway Kohout etal. (2012)
Ceratorhiza sp. Submerged wood and date palm
rachis Egypt Abdel-Aziz (2016b)
Hydnaceae
Ingoldiella fibulata Foam (stream), submerged decaying
leaves, throughfall of palm species India, Malaysia Nawawi (1973b, 1985b), Ghate and
Sridhar (2015)
Ingoldiella nutans Mixed litter of Pinus contorta and
Sphagnum (peat bog) Canada Bandoni and Marvanová (1989)
Rogersiomyces okefeno-
keensis Submerged decaying leaves of angio-
sperms and Taxodium ascendens
(swamp)
USA Crane and Schoknecht (1978)

Fungal Diversity
1 3
Table 12 (continued)
Taxon Substrate Distribution References
Sistotrema hamatum Submerged decaying petioles of
Hevea brasiliensis, submerged uni-
dentified leaves (river), submerged
leaves (Calophyllum brasiliense,
Cinnamomum zeylanicum and
Hevea brasiliensis) (stream), sedi-
ment (stream), foam (river, stream)
Australia, Brazil, India, Malaysia,
Thailand Shaw (1972), Ingold and Webster
(1973), Nawawi and Webster
(1982b), Sridhar (1987), Sridhar
and Kaveriappa (1992), Sakayaroj
etal. (2005), Sudheep and Sridhar
(2012), Barbosa etal. (2013b),
Fiuza etal. (2019), Hosoya etal.
(2019)
Agaricomycetes genera incertae sedis
Akenomyces costatus Roots of rotting Triticum, submerged
leaves of Cladium muhscus France, UK Arnaud (1954), Hornby (1984),
Voglmayr and Krisai-Greilhuber
(1997b)
Cruciger lignatilis Rotting wood Germany Kirschner and Oberwinkler (1999)
Dendrosporomyces prolifer Foam (river), submerged decaying
leaves of Calophyllum brasiliense
(stream)
Brazil, Malaysia Nawawi etal. (1977b), Fiuza etal.
(2019)
Dendrosporomyces splendens Submerged water leaves (river) Malaysia Nawawi (1973c)
Fibulotaeniella canadensis Foam (river) Canada Marvanová and Bärlocher (1988)
Titaeella capnophila Rainwater on needles of Pinus densi-
flora, foam (stream), stemflow and
throughfall of tree canopies
India, Japan, Portugal, Spain Ando and Tubaki (1985), Descals
(1997), Marvanová etal. (2003),
Sridhar and Karamchand (2009)
Tricladiomyces geniculatus Immersed rotten angiospermic leaves
(stream) Malaysia Nawawi and Kuthubutheen (1988g)
Tricladiomyces malaysianus Foam and scum (river), stemflow and
throughfall of tree canopies India, Malaysia Nawawi (1974), Nawawi (1985a),
Sridhar and Karamchand (2009)
BARTHELETIOMYCETES
Polyporales
Polyporales genera incertae sedis
Bulbillomyces farinosus Decaying bark of the fallen branch of
Alnus glutinosa Ukraine Ordynets (2009)
Sebacinales
Sebacinaceae
Chaetospermum camelliae Submerged decaying Licuala
longicalycata (peat swamp), foam
(river)
Thailand Sakayaroj etal. (2005), Pinruan
etal. (2007)
Chaetospermum chaeto-
sporum Foam and scum (stream), submerged
decaying leaves India, USA Anastasiou (1964), Rajashekhar and
Kaveriappa (2003)
Chaetospermum indicum Scum (river), submerged stems
of Fimbristylis quinquangularis
(river)
India Talde (1981)
Trechisporales
Hydnodontaceae
Subulicystidium longispo-
rum Wood (channel) UK Abdullah and Fisher (1984), Hyde
and Goh (1999a)
TREMELLOMYCETES
Tremellales
Tremellales genera incertae sedis
Xenolachne flagellifera Hymenium of Hyaloscypha atomaria
growing on the lower side of damp
logs (Libocedrus decurrens, Pinus
Strobus, Pseudotsuga taxifolia)
USA Rogers (1947)
Agaricomycotina incertae sedis
Nodulospora inconstans Foam (river) Canada Marvanová and Bärlocher (2000)

Fungal Diversity
1 3
Table 12 (continued)
Taxon Substrate Distribution References
PUCCINIOMYCOTINA
ATR ACT IELLOMYCETES
Atractiellales
Phleogenaceae
Helicogloea angustispora Foam – Kirschner (2004)
CLASSICULOMYCETES
Classiculales
Classiculaceae
Classicula fluitans Scirpus microcarpus leaf litter (bank
of roadside drainage channel),
foam (stream)
Canada, Germany Marvanová and Bandoni (1987),
Bauer etal. (2003)
Jaculispora submersa Submerged rotten leaves (stream) Jamaica Hudson and Ingold (1960)
MICROBOTRYOMYCETES
Kriegeriales
Camptobasidiaceae
Camptobasidium hydrophi-
lum (Crucella subtilis)Submerged decaying leaves (stream),
water (stream) France USA Marvanová and Suberkropp (1990),
Cornut etal. (2014)
Microbotryales
Microbotryales incertae sedis
Reniforma strues Biological film (waste-water treat-
ment plant) USA Pore and Sorenson (1990)
USTILAGINOMYCOTINA
EXOBASIDIOMYCETES
Doassansiales
Doassansiaceae
Burrillia narasimhanii Leaves of Caldesia reniformis (pond) India Thirumalachar and Mundkur (1951)
Pseudodermatosorus
alismatis-oligococci Leaves of Caldesia oligococca Sri Lanka Vánky (1999)
Rhamphosporaceae
Rhamphospora nymphaeae Leaves of Nymphaea nouchali, N.
lotus, N. rubra, Caldesia reni-
formis, (pond)
India Cunningham (1888), Lotz-Winter
etal. (2011)
USTILAGINOMYCETES
Urocystidales
Doassansiopsidaceae
Doassansiopsis caldesiae Leaves of Caldesia reniformis in a
marsh close to
lakeshore
Cameroon Piatek etal. (2008)
Doassansiopsis deformans Leaves, petioles, peduncles, pedicels
and ovaries of Sagittaria variabilis USA Engler etal. (1900)
Doassansiopsis euryales Leaves, petioles and ovaries of
Euryale ferox India Verma and Jha (1999)
Doassansiopsis furva Leaves of Sagittaria rigida USA Davis (1919), Vánky (1981)
Doassansiopsis hydrophila Leaves of Potamogeton spp. Spain, USA, UK Regents of the University of the
State of New York (1884),
Fragoso (1917), Lavrov (1937)
Doassansiopsis nymphaeae Petioles of Nymphaea stellata India Sydow and Sydow (1912)
Doassansiopsis nymphoides Leaves of Nymphoides rautaneni
(lake) Zambia Vánky (2002)
Doassansiopsis occulta Ovaries of Potamogeton natans Germany Engler etal. (1900)
Doassansiopsis ticonis Leaves of Nymphaea blanda Costa Rica Piepenbring (1995)

Fungal Diversity
1 3
the abundance of Chytridiomycota and Rozellomycota is
higher than that of Dikarya in some aquatic habitats (Rojas-
Jimenez etal. 2017). The results are mostly derived from
next generation sequencing, but the isolation and proper
description of these fungi remains to be done. Some of the
basal lineages, such as Chytridiomycota have mostly been
isolated from freshwater habitats, such as ponds and lakes
(Blaalid and Khomich 2021). The ecological roles of some
of these fungi have been studied but for many, their roles are
still unclear (Shearer etal. 2007; Rojas-Jimenez etal. 2017).
Hence, comprehensive studies keeping track the number of
taxa isolated from various niches are important to under-
stand their biology and ecology.
Through the years, ecological research on zoosporic
fungi has been conducted mostly in moist and cool fresh-
water ecosystems such as Douglas Lake in the USA, and
Lake Windemere in the English Lake District. Other hot-
spots include countries such South Americas, Australia,
Central Europe, and Japan (Gleason etal. 2010; Blaalid and
Khomich 2021). These fungi are thought to be ubiquitous
in ecosystems which are rich in organic matter but have also
been periodically found in extreme conditions (Gleason
etal. 2010). Hence, it is important to know the ecological
factors that affect the distribution and abundance of these
fungi. Waterborne oxygen content, one the abiotic factors
that influence the distribution and development of zoosporic
fungi, regulates the abundance of these taxa in varying water
depths. Temperature also influences the amount of dissolved
oxygen in water, hence, affecting the growth, reproduction,
and distribution of aquatic fungi (Voronin 2008).
The morphological description of basal fungi has been
well documented, but many taxa lack molecular data, which
limits knowledge of their evolutionary relationship. Many
previously described species either do not have a valid type
specimen or the specimen is unusable. It is also difficult to
know the exact number of species with the ranks currently
circumscribed. These problems can be seen in genera such as
Rhizophydium and Synchytrium wherein the number of spe-
cies present within that genera has been specified in some
studies but acquiring additional information on these taxa is
difficult (Wijayawardene etal. 2020). This section provides a
resource of information of freshwater basal fungi, their sub-
strate, country of isolation and status on DNA-based sequence
data. The list, though not exhaustive, provides an idea on the
number of basal fungi isolated from freshwater habitats.
In this paper, we list 684 species of basal fungi under 11
phyla, 29 orders, 85 families, and 243 genera (Table15).
Chytridiomycota has the largest number with 333 species
(in 97 genera), followed by Rozellomycota (221 species,
105 genera), Blastocladiomycota (47 species, 10 genera),
Monoblepharomycota (29 species, 6 genera), Mucoromycota
(19 species, 10 genera), and Aphelidiomycota (15 species, 3
genera). Other phyla have fewer than 10 species and include
Entomophthoromycota (6 species, 4 genera), Mortierello-
mycota (5 species, 3 genera), Olpidiomycota (4 species, 1
genus), Zoopagomycota (3 species, 2 genera), and Sanchy-
triomycota (2 species, 2 genera).
Microsporidia
The number microsporidia from all environments ranges
from 118 genera (Sprague etal. 1992), 144 in Sprague and
Becnel (1999) to 200 listed in Becnel etal. (2014), and 217
in Wijayawardene etal. (2020). Species numbers may exceed
1,500 and this may be a conservative estimate. The website
www. marin efungi. org lists 62 genera with 138 species of
marine microsporidia (Jones etal. 2019). Data on freshwa-
ter microsporidia is spread widely in the literature with no
definitive list of actual species and their hosts. Table16 lists
105 genera of freshwater microsporidia with 221 species in 4
orders and 24 families, but this undoubtedly is an underesti-
mate as new taxa are continuously being introduced.
Table 12 (continued)
Taxon Substrate Distribution References
Doassansiopsis tomasii Leaves of submerged Nymphaea
nouchali var. caerulea in lake
margins
Cameroon, Ethiopia, Uganda Piatek etal. (2008)
Basidiomycota genera incertae sedis
Anguillomyces acadiensis Foam (river) Canada Marvanová and Bärlocher (2000)
Anguillomyces acadiensis Foam (river) Canada Marvanová and Bärlocher (2000)
Arcispora bisagittaria Foam (stream) Canada (Marvanová and Bärlocher 1998a)
Microstella pluvioriens Rainwater from leaves of Metase-
quoia glyptostroboides Japan Ando and Tubaki (1984b)
Stauriella aquatica Submerged test blocks of Diptero-
carpus alatus (stream) Thailand Sivichai and Jones (2004)

Fungal Diversity
1 3
Table 13 Classification of freshwater yeasts with information on the type of habitats, substrates, and geographical distribution of fungal taxa
Taxon Substrate Distribution References
ASCOMYCOTA
DOTHIDEOMYCETES
Dothideomycetidae
Dothideomycetes incertae sedis
Endosporiaceae
Endosporium aviarium Phytotelmata of bromeliad tanks, rock holes Brazil Morais etal. (2020)
EUROTIOMYCETES
Chaetothyriomycetidae
Chaetothyriales
Herpotrichiellaceae
Exophiala alcalophila Phytotelmata of bromeliad tank, tap water Brazil Novak Babič etal.
(2016), Morais etal.
(2020)
Exophiala placitae Phytotelmata of bromeliad tank Brazil Morais etal. (2020)
Nadsoniella nigra Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
LEOTIOMYCETES
Leotiomycetidae
Helotiales
Sclerotiniaceae
Monilia zeylanica Freshwater everglades USA Fell etal. (2011)
SACCHAROMYCETES
Saccharomycetidae
Saccharomycetales
Debaryomycetaceae
Babjeviella inositovora Surfaces of freshwater turtles Podocnemis unifilis Brazil Morais etal. (2010,
2020)
Debaryomyces delbrueckii Phytotelmata of bromeliad tanks, water (pond) Brazil, Slovakia Sláviková and Vadker-
tiová (1993), Morais
etal. (2020)
Debaryomyces hansenii Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, water (lake, pond), tapwater, glacial meltwa-
ter (river)
Argentina, Bangla-
desh, Brazil, Poland,
Slovakia
Sláviková and Vad-
kertiová (1995b),
Khan etal. (2002), de
García etal. (2007),
Morais etal. (2010),
Biedunkiewicz etal.
(2013)
Debaryomyces maramus Water (lake) Poland Biedunkiewicz etal.
(2013)
Debaryomyces nepalensis Freshwater mud Egypt Moubasher etal.
(2018)
Debaryomyces occidentalis var.
occidentalis Surfaces of freshwater turtles, P. unifilis Brazil Morais etal. (2010)
Lodderomyces elongisporus Water (lake) Poland Biedunkiewicz etal.
(2013)
Meyerozyma caribbica Phytotelmata of bromeliad tanks, tapwater, mud (river,
canal), water (lake) Brazil, Slovenia,
Egypt Brandão etal. (2010),
Novak Babič etal.
(2016), Moubasher
etal. (2018), Morais
etal. (2020)
Meyerozyma carpophila Freshwater Everglades USA Fell etal. (2011)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Meyerozyma guilliermondii Water (lake, pond, rivers), phytotelmata of bromeliad
tanks, surfaces of freshwater turtles Podocnemis
expansa, P. unifilis, tapwater, mud (river, canal)
Brazil, Egypt, Poland,
Slovakia, Slovenia,
South Africa, USA
van Uden and Ahearn
(1963), Sláviková
etal. (1992), Slávik-
ová and Vadkertiová
(1993), Brandão etal.
(2010), Medeiros
etal. (2012), Van
Wyk etal. (2012),
Biedunkiewicz etal.
(2013), Brilhante
etal. (2016), Novak
Babič etal. (2016),
Moubasher etal.
(2018), Morais etal.
(2020)
Millerozyma farinosa Water (lake) Poland Biedunkiewicz etal.
(2013)
Priceomyces carsonii Water (lake) Poland Biedunkiewicz etal.
(2013)
Scheffersomyces coipomensis Water (lake) Argentina Libkind etal. (2009)
Scheffersomyces segobiensis Water (lake) Poland Biedunkiewicz etal.
(2013)
Schwanniomyces occidentalis Water (lake) Poland Biedunkiewicz etal.
(2013)
Schwanniomyces polymorphus Water (lake) Poland Biedunkiewicz etal.
(2013)
Schwanniomyces pseudopolymor-
phus Water (lake) Poland Biedunkiewicz etal.
(2013)
Schwanniomyces vanrijiae Freshwater Everglades, surfaces of freshwater turtles
Podocnemis expansa, P. unifilis Brazil, USA Morais etal. (2010),
Fell etal. (2011)
Suhomyces xylopsoci Water (lake) Brazil Brandão etal. (2010)
Yamadazyma akitaensis Water (lake) Poland Biedunkiewicz etal.
(2013)
Yamadazyma tenuis Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis Brazil Morais etal. (2010)
Dipodascaceae
Dipodascus armillariae Water (lake) Poland Biedunkiewicz etal.
(2013)
Dipodascus fermentans Polluted water (river) USA Woollett and Hedrick
(1970)
Dipodascus geotrichum River water, freshwater mud (river, canal), water (lake,
pond), tapwater Brazil, Egypt, Slova-
kia, Slovenia, USA Lamore and Goos
(1978), Sláviková
etal. (1992), Slávik-
ová and Vadkertiová
(1995b), Medeiros
etal. (2012), Novak
Babič etal. (2016),
Moubasher etal.
(2018)
Geotrichum europaeum Mud (river, canal) Egypt Moubasher etal.
(2018)
Magnusiomyces capitatus Water (lake) Poland Biedunkiewicz etal.
(2013)
Saprochaete capitata Polluted water (river) USA Woollett and Hedrick
(1970)
Saprochaete sericea Polluted water (river) USA Woollett and Hedrick
(1970)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Lipomycetaceae
Lipomyces lipofer Water (lake) Poland Biedunkiewicz etal.
(2013)
Metschnikowiaceae
Clavispora lusitaniae Water (lake, river), tapwater South Africa, Slovenia Van Wyk etal. (2012),
Novak Babič etal.
(2016)
Kodamaea ohmeri Water (lake) Brazil Brilhante etal. (2016)
Metschnikowia pulcherrima Water (lake) Poland, Slovakia, USA van Uden and Ahearn
(1963), Sláviková
etal. (1992), Bie-
dunkiewicz etal.
(2013)
Metschnikowia reukaufii Water (lake, river) Poland, Brazil Medeiros etal. (2012),
Biedunkiewicz etal.
(2013)
Phaffomycetaceae
Barnettozyma wickerhamii Surfaces of freshwater turtles Podocnemis expansa Brazil Morais etal. (2010)
Cyberlindnera fabianii Water (lake) Brazil Brandão etal. (2010)
Cyberlindnera jadinii Water (lake), freshwater mud, polluted water (lake) Egypt, Poland, USA Woollett and Hedrick
(1970), Biedunkie-
wicz etal. (2013),
Moubasher etal.
(2018)
Cyberlindnera japonica Water (lake) Brazil Brandão etal. (2010)
Cyberlindnera rhodanensis Water (lake) Brazil Brandão etal. (2010)
Cyberlindnera saturnus Polluted water (river), water, sediment (lake) Colombia, USA Woollett and Hedrick
(1970), Silva-Bedoya
etal. (2014)
Starmera stellimalicola Water (lake) Brazil Brandão etal. (2010)
Pichiaceae
Brettanomyces nanus Water (lake) Poland Biedunkiewicz etal.
(2013)
Dekkera bruxellensis Water (lake), polluted water (river) Brazil, Poland, USA Woollett and Hedrick
(1970), Brandão etal.
(2010), Biedunkie-
wicz etal. (2013)
Komagataella pastoris Surfaces of freshwater turtles Brazil Morais etal. (2010)
Kregervanrija fluxuum Freshwater Everglades USA Fell etal. (2011)
Ogataea polymorpha Polluted water (river) USA Woollett and Hedrick
(1970)
Pichia fermentans Water (lake, pond), tapwater Brazil, Slovakia,
Slovenia Sláviková etal. (1992),
Sláviková and
Vadkertiová (1995b),
Brandão etal. (2010),
Novak Babič etal.
(2016)
Pichia galeiformis Water (lake) Brazil Brandão etal. (2010)
Pichia kluyveri Freshwater Everglades USA Fell etal. (2011)
Pichia mandshurica Water (lake) Brazil Brandão etal. (2010)
Pichia membranifaciens Water (lake), Surfaces of freshwater turtles Podocnemis
unifilis Brazil, Poland Morais etal. (2010),
Biedunkiewicz etal.
(2013)
Saturnispora diversa Water, sediment (lake) Colombia Silva-Bedoya etal.
(2014)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Saturnispora silvae Water (lake, wetland) Poland, USA Fell etal. (2011),
Biedunkiewicz etal.
(2013)
Saccharomycetaceae
Citeromyces matritensis Water (lake, river) South Africa, Poland Van Wyk etal. (2012),
Biedunkiewicz etal.
(2013)
Cyniclomyces guttulatus Water (lake) Poland Biedunkiewicz etal.
(2013)
Grigorovia transvaalensis Water (lake) Poland Biedunkiewicz etal.
(2013)
Hagleromyces aurorensis Phytotelmata of bromeliad tank Brazil Sousa etal. (2014),
Morais etal. (2020)
Issatchenkia orientalis Tropical freshwater, water (lake, pond), mud (river
canal), wetland Brazil, Egypt, Poland,
Slovakia, USA Sláviková etal. (1992),
Sláviková and
Vadkertiová (1995b),
Brandão etal. (2010),
Fell etal. (2011),
Medeiros etal.
(2012), Biedunkie-
wicz etal. (2013),
Moubasher etal.
(2018)
Issatchenkia scutulata Water (river) Brazil Medeiros etal. (2012)
Kazachstania africana Water (lake) Poland Biedunkiewicz etal.
(2013)
Kazachstania exigua Water (lake), polluted water (river) Poland, USA Biedunkiewicz etal.
(2013)
Kazachstania lodderae Water (lake) Poland Biedunkiewicz etal.
(2013)
Kluyveromyces aestuarii Water (lake, wetland) Poland, USA Fell etal. (2011),
Biedunkiewicz etal.
(2013)
Kluyveromyces lactis Water (lake) Poland, Slovakia Sláviková etal. (1992),
Biedunkiewicz etal.
(2013)
Kluyveromyces marxianus Water (lake, river) Brazil, Poland, Medeiros etal. (2012)
Kluyveromyces wickerhamii Water (lake), surfaces of freshwater turtles Podocnemis
unifilis Brazil, Poland Morais etal. (2010),
Biedunkiewicz etal.
(2013)
Lachancea cidri Water (lake) Poland Biedunkiewicz etal.
(2013)
Lachancea kluyveri Water (lake) Slovakia Sláviková etal. (1992)
Lachancea meyersii Water (wetland) USA Fell etal. (2011)
Lachancea thermotolerans Water (lake) Poland Biedunkiewicz etal.
(2013)
Lachancea waltii Water (lake) Poland Biedunkiewicz etal.
(2013)
Nakaseomyces bacillisporus Water (lake) Poland Biedunkiewicz etal.
(2013)
Naumovozyma castellii Water (lake) Bangladesh, Poland Biedunkiewicz etal.
(2013)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Saccharomyces cerevisiae Water (lake), polluted water (river), mud (river, canal),
sediment (lake) Colombia, Egypt,
Poland, USA Woollett and Hedrick
(1970), Khan etal.
(2002), Biedunkie-
wicz etal. (2013),
Silva-Bedoya etal.
(2014), Moubasher
etal. (2018)
Tetrapisispora phaffii Water (lake) Poland Biedunkiewicz etal.
(2013)
Torulaspora delbrueckii Water, sediment (lake) Colombia, Poland Biedunkiewicz etal.
(2013), Silva-Bedoya
etal. (2014)
Torulaspora globosa Water (pond) Slovakia Sláviková and Vadker-
tiová (1995b)
Torulaspora pretoriensis Freshwater muds, water, sediment (lake) Colombia, Egypt Silva-Bedoya etal.
(2014), Moubasher
etal. (2018)
Vanderwaltozyma polyspora Water (lake) Poland Biedunkiewicz etal.
(2013)
Vanderwaltozyma yarrowii Water (lake) Poland Biedunkiewicz etal.
(2013)
Zygosaccharomyces bailii Water (lake) Poland Biedunkiewicz etal.
(2013)
Zygosaccharomyces mellis Water (lake) Poland Biedunkiewicz etal.
(2013)
Zygosaccharomyces rouxii Water (lake) Poland Biedunkiewicz etal.
(2013)
Saccharomycodaceae
Hanseniaspora guilliermondii Water (lake, river) Brazil, Slovakia Sláviková etal. (1992),
Medeiros etal.
(2012)
Hanseniaspora opuntiae Phytotelmata of bromeliad tank, water (lake) Argentina, Brazil Libkind etal. (2009),
Morais etal. (2020)
Hanseniaspora osmophila Water (lake) Poland Biedunkiewicz etal.
(2013)
Hanseniaspora uvarum Freshwater Everglades, polluted water (river), water
(lake, pond), sediment (lake) Colombia, Poland,
Slovakia, USA Woollett and Hedrick
(1970), Sláviková
etal. (1992), Slávik-
ová and Vadkertiová
(1993), Fell etal.
(2011), Biedunkie-
wicz etal. (2013),
Silva-Bedoya etal.
(2014)
Hanseniaspora vineae Water (lake) Poland Biedunkiewicz etal.
(2013)
Saccharomycodes ludwigii Water (lake) Poland Biedunkiewicz etal.
(2013)
Saccharomycopsidaceae Water (lake)
Saccharomycopsis capsularis Water (lake) Poland Biedunkiewicz etal.
(2013)
Saccharomycopsis crataegensis Water (river) Brazil Medeiros etal. (2012)
Saccharomycopsis fermentans Water (lake) Poland Biedunkiewicz etal.
(2013)
Saccharomycopsis fibuligera Water (lake) Slovakia Sláviková etal. (1992)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Saccharomycopsis javanensis Water (lake) Poland Biedunkiewicz etal.
(2013)
Trichomonascaceae
Sugiyamaella smithiae Water (lake) Poland Biedunkiewicz etal.
(2013)
Trichomonascus ciferrii Water (lake), polluted water (river) Brazil, Poland, USA,
Slovakia Woollett and Hedrick
(1970), Sláviková
etal. (1992), Bril-
hante etal. (2016)
Trichomonascus farinosus Water (lake) Poland Biedunkiewicz etal.
(2013)
Wickerhamiella pararugosa Water (river), tapwater Brazil, Slovenia Medeiros etal. (2012),
Novak Babič etal.
(2016)
Zygoascus tannicola Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis Brazil Morais etal. (2010)
Wickerhamomycetaceae
Wickerhamomyces anomalus Tropical freshwater, surfaces of freshwater turtles,
water (lake, pond), mud (river, canal) Brazil, Egypt, Poland,
Slovakia Sláviková etal. (1992),
Sláviková and
Vadkertiová (1995b),
Morais etal. (2010),
Biedunkiewicz etal.
(2013), Moubasher
etal. (2018)
Wickerhamomyces bisporus Water (lake) Poland Biedunkiewicz etal.
(2013)
Wickerhamomyces bovis Water (lake) Poland Biedunkiewicz etal.
(2013)
Wickerhamomyces canadensis Polluted water (river) USA Woollett and Hedrick
(1970)
Wickerhamomyces hampshirensis Freshwater everglades USA Fell etal. (2011)
Wickerhamomyces pijperi Water (river) Brazil Medeiros etal. (2012)
Wickerhamomyces patagonicus Glacial meltwater (river) Argentina de García etal. (2012)
Wickerhamomyces rabaulensis Water (lake)
Wickerhamomyces subpelliculosus Polluted water (lake, river) USA Woollett and Hedrick
(1970)
Wickerhamomyces sydowiorum Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, water (wetland) Brazil, USA Morais etal. (2010),
Fell etal. (2011)
Saccharomycetales genera incertae sedis
Candida albicans Phytotelmata of bromeliad tanks, polluted water (lake,
river), sediment (lake) Brazil, Colombia,
Egypt, Poland, USA Woollett and Hedrick
(1970), Brandão etal.
(2010), Biedunkie-
wicz etal. (2013),
Silva-Bedoya etal.
(2014), Moubasher
etal. (2018), Morais
etal. (2020)
Candida amphicis Freshwater Everglades USA Fell etal. (2011)
Candida asiatica Freshwater Everglades USA Fell etal. (2011)
Candida atlantica Water (lake) Brazil Brandão etal. (2010)
Candida atmosphaerica Polluted water (river) USA Woollett and Hedrick
(1970)
Candida blankii Water (river) Brazil Medeiros etal. (2012)
Candida boidinii Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, water (pond) Brazil, Slovakia Sláviková and Vadker-
tiová (1995b), Morais
etal. (2010)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Candida buenavistaensis Phytotelmata of bromeliad tanks, rock holes Brazil Morais etal. (2020)
Candida dendronema Freshwater Everglades USA Fell etal. (2011)
Candida diffluens Polluted water (river) USA Woollett and Hedrick
(1970)
Candida duobushaemulonis Phytotelmata of bromeliad tanks, rock holes Brazil Morais etal. (2020)
Candida ethanolica Water (lake, river) Brazil Medeiros etal. (2012)
Candida fluviatilis Aquatic system, glacial water Argentina Libkind etal. (2014)
Candida glabrata Phytotelmata of bromeliad tanks, rock holes, polluted
water (lake, river), sediment (lake) Brazil, Colombia,
Poland, USA Woollett and Hedrick
(1970), Brandão etal.
(2010), Biedunkie-
wicz etal. (2013),
Silva-Bedoya etal.
(2014), Morais etal.
(2020)
Candida heveicola Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Candida inconspicua Water (lake, pond), polluted water (river) Brazil, Slovakia, USA Sláviková and Vad-
kertiová (1995b),
Brandão etal. (2010)
Candida intermedia Water (lake, pond), Phytotelmata of bromeliad tanks,
rock holes, polluted water (river) Brazil, Poland, Slova-
kia, USA Woollett and Hedrick
(1970), Sláviková
and Vadkertiová
(1993), Biedunkie-
wicz etal. (2013),
Morais etal. (2020)
Candida leandrae Phytotelmata of bromeliad tanks, rock holes Brazil Morais etal. (2020)
Candida maris Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, wetland Brazil, USA Morais etal. (2010),
Fell etal. (2011)
Candida maritima Glacial meltwater (river) Argentina de García etal. (2012)
Candida melibiosica Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, water (lake, river, wetland) Brazil, Poland, USA Fell etal. (2011),
Medeiros etal.
(2012), Biedunkie-
wicz etal. (2013)
Candida mesenterica Polluted water (river), Glacial meltwater (river) USA, Argentina Woollett and Hedrick
(1970) de García
etal. (2012)
Candida nivariensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Candida oregonensis Polluted water (river) USA Woollett and Hedrick
(1970)
Candida orthopsilosis Phytotelmata of bromeliad tanks, rock holes, tapwater Brazil, Slovenia Morais etal. (2020)
Candida palmioleophila Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis, water (river) Brazil Medeiros etal. (2012),
Novak Babič etal.
(2016), Morais etal.
(2020)
Candida parapsilosis Tropical freshwater, phytotelmata of bromeliad tanks,
rock holes, surfaces of freshwater turtles Podocne-
mis expansa, P. unifilis, polluted water (lake, river),
groundwater, tapwater
Brazil, Slovenia, USA Woollett and Hedrick
(1970), Brandão etal.
(2010), Morais etal.
(2010), Medeiros
etal. (2012), Morais
etal. (2020)
Candida pseudointermedia Phytotelmata of bromeliad tanks, rock holes, tapwater Brazil, Slovenia Novak Babič etal.
(2016), Morais etal.
(2020)
Candida pseudolambica Aquatic system, glacial melt water, mud (river, canal),
water (lake, wetland), sediment (lake) Argentina, Colombia,
Egypt, USA Fell etal. (2011),
Libkind etal. (2014),
Silva-Bedoya etal.
(2014), Moubasher
etal. (2018)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Candida pinus Polluted water (lake, river) USA Woollett and Hedrick
(1970)
Candida saitoana Water (lake), tapwater Poland, Slovenia Biedunkiewicz etal.
(2013), Novak Babič
etal. (2016)
Candida sake Acidic water (river)ater (lake), glacial melt water,
surfaces of freshwater turtles Podocnemis expansa,
P. unifilis
Argentina, Brazil Russo etal. (2008),
Morais etal. (2010),
Van Wyk etal.
(2012), Biedunkie-
wicz etal. (2013),
Libkind etal. (2014)
Candida santamariae Glacial meltwater (river) Italy Branda etal. (2010)
Candida sharkensis Water (wetland) USA Fell etal. (2011)
Candida silvanorum Water (wetland) USA Fell etal. (2011)
Candida silvicultrix Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis Brazil Morais etal. (2010)
Candida solani Water (lake), polluted water (river) Poland, USA Woollett and Hedrick
(1970), Biedunkie-
wicz etal. (2013)
Candida sonorensis Water (lake) Brazil Brandão etal. (2010)
Candida suecica Polluted water (river) Bangladesh, USA Woollett and Hedrick
(1970), Khan etal.
(2002)
Candida tropicalis Tropical freshwater, water (lake, pond), mud (river
canal) Brazil, Egypt, Poland,
Slovakia, South
Africa
Sláviková etal.
(1992), Sláviková
and Vadkertiová
(1995b), Brandão
etal. (2010), Van
Wyk etal. (2012),
Biedunkiewicz etal.
(2013), Moubasher
etal. (2018)
Candida ubatubensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Candida zeylanoides Water (lake) Poland Biedunkiewicz etal.
(2013)
Diutina catenulata Water (lake. river), polluted water (river) South Africa, Poland,
USA Woollett and Hedrick
(1970), Biedunkie-
wicz etal. (2013)
Diutina rugosa Tropical freshwater, water (river), polluted water (river) Brazil, South Africa,
USA Woollett and Hedrick
(1970), Medeiros
etal. (2012), Van
Wyk etal. (2012)
Hyphopichia burtonii Water (lake, pond) Brazil, Slovakia Sláviková etal. (1992),
Sláviková and
Vadkertiová (1993),
Brandão etal. (2010)
Middelhovenomyces tepae Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis Brazil Morais etal. (2010)
Nakazawaea holstii Polluted water (river) USA Woollett and Hedrick
(1970)
Nakazawaea ernobii Polluted water (river) USA Woollett and Hedrick
(1970)
Yarrowia galli Surfaces of freshwater turtles Podocnemis expansa, P.
unifilis Brazil Morais etal. (2010)
Yarrowia lipolytica Water (lake), tapwater Colombia, Poland,
Slovenia Biedunkiewicz etal.
(2013), Silva-Bedoya
etal. (2014)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
BASIDIOMYCOTA
AGARICOSTILBOMYCETES
Agaricostilbomycetes incertae sedis
Agaricostilbales
Agaricostilbaceae
Sterigmatomyces elviae Surfaces of freshwater turtles Podocnemis expansa Brazil Morais etal. (2010)
MONILIELLOMYCETES
Moniliellomycetes incertae sedis
Moniliellales
Moniliellaceae
Moniliella spathulata Water (lake) Poland Biedunkiewicz etal.
(2013)
CYSTOBASIDIOMYCETES
Cystobasidiomycetes incertae sedis
Cystobasidiales
Cystobasidiaceae
Cystobasidium calyptogenae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Cystobasidium minutum Freshwater Everglades, polluted water (lake, river),
water (lake, pond) Argentina, Slovakia,
USA Woollett and Hedrick
(1970), Sláviková
etal. (1992), Slávik-
ová and Vadkertiová
(1995b), Libkind
etal. (2009), Fell
etal. (2011)
Cystobasidium laryngis Glacial melt water (river), water (lake) Argentina de García etal. (2007),
Libkind etal. (2009)
Cystobasidium pinicola Water (lake) Argentina, Brazil Libkind etal. (2003),
Brandão etal. (2011)
Cystobasidium slooffiae Tapwater, water (lake, wetland) Argentina, Brazil,
Slovenia, USA Libkind etal. (2003),
Brandão etal. (2011),
Fell etal. (2011),
Novak Babič etal.
(2016)
Occultifur brasiliensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Occultifur externus Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Erythrobasidiales
Erythrobasidiaceae
Erythrobasidium elongatum Phytotelmata of bromeliad Brazil Morais etal. (2020)
Erythrobasidium hasegawae Phytotelmata of bromeliad Brazil Morais etal. (2020)
Cyrenella elegans Sand near lake USA Sampaio (2010)
Hasegawazyma lactosa Phytotelmata of bromeliad tanks, polluted water (river) Brazil, USA Woollett and Hedrick
(1970), Morais etal.
(2020)
Symmetrospora marina Phytotelmata of bromeliad tanks, water (wetland) Brazil, USA Fell etal. (2011),
Morais etal. (2020)
Sakaguchia cladiensis Water (wetland) USA Fell etal. (2011)
Sakaguchia meli Glacial melt water Argentina Libkind etal. (2014)
EXOBASIDIOMYCETES
Exobasidiomycetidae
Exobasidiales
Brachybasidiaceae
Meira geulakonigae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Georgefischeriales

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Gjaerumiaceae
Gjaerumia minor Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Tilletiariaceae
Phragmotaenium oryzicola Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Microstromatales
Microstromatales genera incertae sedis
Sympodiomycopsis paphiopedili Water (river) Brazil Medeiros etal. (2012)
MICROBOTRYOMYCETES
Kriegeriales
Kriegeriaceae
Phenoliferia himalayensis Water (lake) Himalayas Shivaji etal. (2008)
Leucosporidiales
Leucosporidiaceae
Leucosporidium creatinivorum Glacial melt water (river), water (lake) Argentina de García etal. (2007),
Libkind etal. (2009)
Leucosporidium fragarium Glacial melt water (river) Argentina de García etal. (2007)
Leucosporidium golubevii Water (river) Sampaio etal. (2003)
Leucosporidium intermedium Glacial meltwater (river) Argentina de García etal. (2012)
Leucosporidium muscorum Water (lake), polluted water (river) Argentina, USA Woollett and Hedrick
(1970), Libkind etal.
(2009)
Leucosporidium scottii River water, polluted water (lake, river), water (lake,
pond) Slovakia, USA Woollett and Hedrick
(1970), Summerbell
(1983), Sláviková
etal. (1992), Slá-
viková and Vadker-
tiová (1993)
Microbotryales
Ustilentylomataceae
Microbotryozyma collariae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Microbotryomycetes families incertae sedis
Chrysozymaceae
Bensingtonia yamatoana Glacial meltwater (river) Argentina de García etal. (2012)
Colacogloeaceae
Colacogloea foliorum Phytotelmata of bromeliad tanks, polluted water (lake,
river) Brazil, USA Woollett and Hedrick
(1970), Morais etal.
(2020)
Microbotryomycetes genera incertae sedis
Reniforma strues Wastewater Poland Pore and Sorenson
(1990)
Sampaiozyma ingeniosa Water (lake), polluted water (river) USA Woollett and Hedrick
(1970), Sampaio
(2011)
Sampaiozyma vanillica Freshwater Everglades USA Fell etal. (2011)
Slooffia cresolica Freshwater Everglades USA Fell etal. (2011)
Udeniozyma ferulica Water, glacial lakes and rivers, water (wetland) Argentina, USA Sampaio and Van
Uden (1991), Kurtz-
man and Fell (1998),
Libkind etal. (2003),
Fell etal. (2011)
Sporidiobolales
Sporidiobolaceae

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Rhodosporidiobolus colostri Glacial meltwater (lakes, river), acidic water (river) Argentina de García etal. (2007),
Russo etal. (2008)
Rhodosporidiobolus lusitaniae Water (lake) Poland Biedunkiewicz etal.
(2013)
Rhodosporidiobolus ruineniae Freshwater Everglades, phytotelmata of bromeliad
tanks Brazil, USA Fell etal. (2011),
Morais etal. (2020)
Rhodotorula alborubescens Water (pond) Slovakia Sláviková and Vadker-
tiová (1993)
Rhodotorula babjevae Water (lake), acidic water (river) Argentina Libkind etal. (2003),
Russo etal. (2008)
Rhodotorula diobovata Freshwater Everglades, water (lake) Argentina, USA Libkind etal. (2009),
Fell etal. (2011)
Rhodotorula evergladensis Water (wetland) USA Fell etal. (2011)
Rhodotorula glutinis Water (lake), polluted water (lake, river) Bangladesh, South
Africa, Slovakia,
USA
van Uden and Ahearn
(1963), Woollett
and Hedrick (1970),
Sláviková etal.
(1992), Khan etal.
(2002), Van Wyk
etal. (2012)
Rhodotorula graminis Polluted water (lake, river) USA Woollett and Hedrick
(1970)
Rhodotorula kratochvilovae Glacial melt water (river) Argentina Libkind etal. (2003),
de García etal.
(2007)
Rhodotorula mucilaginosa Glacial meltwater (lake, river), tropical freshwaters,
phytotelmata of bromeliad tanks, rock holes, surfaces
of freshwater turtles Podocnemis expansa, ground-
water, mud (river, canal), water (lake, wetland, river),
acidic water (lake, river), sediment (lake)
Argentina, Bangla-
desh, Brazil, Colom-
bia, Egypt, Poland,
Slovakia, Slovenia,
USA,
van Uden and Ahearn
(1963), Sláviková
etal. (1992), Khan
etal. (2002), de
García etal. (2007),
Libkind etal. (2009),
Morais etal. (2010),
Fell etal. (2011),
Sampaio (2011), Van
Wyk etal. (2012),
Silva-Bedoya etal.
(2014), Brilhante
etal. (2016), Novak
Babič etal. (2016),
Moubasher etal.
(2018), Pietryczuk
etal. (2018), Morais
etal. (2020)
Rhodotorula paludigena Phytotelmata of bromeliad tanks, water (wetland) Brazil, USA Fell etal. (2011),
Morais etal. (2020)
Rhodotorula sphaerocarpa Water Portugal Sampaio (2011)
Rhodotorula taiwanensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Rhodotorula toruloides Water (lake), acidic water (river), phytotelmata of
bromeliad tanks Argentina, Brazil,
Poland Russo etal. (2008),
Biedunkiewicz etal.
(2013), Morais etal.
(2020)
Sporobolomyces beijingensis Freshwater Everglades, Water (lake) USA Fell etal. (2011)
Sporobolomyces blumeae Water (wetland) USA Fell etal. (2011)
Sporobolomyces carnicolor Water (wetland) USA Fell etal. (2011)
Sporobolomyces japonicus Water (wetland) USA Fell etal. (2011)
Sporobolomyces longiusculus Water (lake) Argentina Libkind etal. (2005,
2009)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Sporobolomyces marcillae Water (lake) Argentina Libkind etal. (2009)
Sporobolomyces patagonicus Water (lake) Argentina Libkind etal. (2005)
Sporobolomyces roseus Freshwater Everglades, glacial waters and rivers, water
(pond) Argentina, USA,
Slovakia Sláviková and Vad-
kertiová (1995b),
Libkind etal. (2005),
Fell etal. (2011)
Sporobolomyces ruberrimus Freshwater Everglades, Glacial meltwater (river) USA, Argentina Libkind etal. (2009),
Fell etal. (2011), de
García etal. (2012)
Sporobolomyces salmonicolor Water (lake) Argentina, Slovakia Sláviková etal. (1992),
Libkind etal. (2005)
Sporobolomyces shibatanus Water (wetland) USA Fell etal. (2011)
Sporidiobolus salmonicolor Groundwater Slovenia Novak Babič etal.
(2016)
TREMELLOMYCETES
Tremellomycetes incertae sedis
Filobasidiales
Filobasidiaceae
Filobasidium floriforme Freshwater Everglades, Surfaces of freshwater turtles
Podocnemis expansa, P. unifilis USA, Brazil Morais etal. (2010),
Fell etal. (2011),
Morais etal. (2020)
Filobasidium magnum Water (lake, river, wetland) Brazil, Slovakia, USA Sláviková etal. (1992),
Brandão etal. (2011),
Fell etal. (2011),
Medeiros etal.
(2012)
Filobasidium oeirens Glacial meltwater (river) Italy Branda etal. (2010)
Filobasidium stepposum Water (lake), Glacial meltwater (river) Brazil, Italy Branda etal. (2010),
Brandão etal. (2011)
Filobasidium wieringae Water (lake), Glacial meltwater (river) Argentina, Brazil,
Italy Branda etal. (2010),
Brandão etal. (2011),
de García etal.
(2012)
Goffeauzyma agrionensis Glacial lakes and rivers Argentina de García etal. (2007)
Goffeauzyma gastrica Glacial meltwater (river), water (lake) Argentina, Italy, USA, van Uden and Ahearn
(1963), Libkind etal.
(2009), Branda etal.
(2010)
Goffeauzyma gilvescens Glacial melt water Argentina Turchetti etal. (2008)
Naganishia adeliensis Glacial melt water (river) Argentina, Italy de García etal. (2007),
Branda etal. (2010)
Naganishia albida Water (lake, pond), rock pools, surfaces of freshwater
turtles Podocnemis expansa, P. unifilis, polluted water
(river)
Argentina, Brazil,
Slovakia, USA van Uden and Ahearn
(1963), Woollett
and Hedrick (1970),
Sláviková and
Vadkertiová (1995b),
Libkind etal. (2009),
Morais etal. (2010)
Naganishia albidosimilis Water (pond), glacial meltwater (river), acidic water
(river) Argentina, Italy,
Portugal Gadanho etal. (2006),
Russo etal. (2008),
Branda etal. (2010)
Naganishia antarctica Water (lake), acidic water (lake) Argentina Russo etal. (2008),
Libkind etal. (2009)
Naganishia bhutanensis Acidic water (lake) Argentina Russo etal. (2008)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Naganishia diffluens Freshwater Everglades, polluted water (lake, river),
water (lake) USA van Uden and Ahearn
(1963), Woollett and
Hedrick (1970), Fell
etal. (2011)
Naganishia liquefaciens Water (wetland) USA Fell etal. (2011)
Piskurozymaceae
Piskurozyma cylindrica Glacial melt water (river) Argentina de García etal. (2007)
Solicoccozyma aeria Polluted water (river), water (lake), glacial meltwater
(river) Italy, Slovakia, USA Woollett and Hedrick
(1970), Sláviková
etal. (1992), Branda
etal. (2010)
Solicoccozyma terrea Freshwater Everglades USA Fell etal. (2011)
Solicoccozyma terricola Polluted water (river), glacial meltwater (river) Argentina, USA Woollett and Hedrick
(1970), de García
etal. (2012)
Holtermanniales
Holtermanniaceae
Holtermanniella festucosa Glacial meltwater (river) Argentina de García etal. (2007),
Wuczkowski etal.
(2011), Libkind etal.
(2014)
Tremellales
Bulleraceae
Bullera alba Rock holes, water (lake, pond) Brazil, Slovakia Sláviková etal. (1992),
Sláviková and
Vadkertiová (1995b),
Medeiros etal.
(2012)
Bulleribasidiaceae
Dioszegia butyracea Glacial meltwater (river) Argentina de García etal. (2012)
Dioszegia crocea Glacial melt water (river) Argentina de García etal. (2007)
Dioszegia fristingensis Glacial melt water (river), water (lake) Argentina de García etal.
(2007),Libkind etal.
(2009)
Dioszegia hungarica Glacial melt water (river), water (lake) Argentina de García etal. (2007),
Libkind etal. (2009)
Dioszegia zsoltii Freshwater Everglades USA Fell etal. (2011)
Hannaella luteola River water, phytotelmata of bromeliad tanks, rock
pools, polluted water (lake) Brazil, USA Woollett and Hedrick
(1970), Medeiros
etal. (2012), Morais
etal. (2020)
Hannaella pagnoccae Phytotelmata of bromeliad tanks, rock pools Brazil Morais etal. (2020)
Hannaella sinensis Freshwater Everglades USA Fell etal. (2011)
Hannaella zeae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Vishniacozyma carnescens Water (lake) Brazil Brandão etal. (2011)
Vishniacozyma dimennae Glacial meltwater (river) Italy Branda etal. (2010)
Vishniacozyma victoriae Water (lake), submerged roots, tapwater, glacial melt-
water (river), acidic water (lake, river) Argentina, Brazil,
Norway, Slovenia Russo etal. (2008),
Brandão etal. (2011),
Vaz etal. (2011),
Kohout etal. (2012),
de García etal.
(2012)
Cryptococcaceae

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Cryptococcus festucosus Glacial meltwater (river), acidic water (river) Argentina, Italy de García etal. (2007),
Russo etal. (2008),
Branda etal. (2010)
Cryptococcus heveanensis Water (lake) Brazil Brandão etal. (2011)
Cryptococcus hungaricus Water (river) Brazil Medeiros etal. (2012)
Cryptococcus macerans Glacial meltwater (river) Argentina, Italy de García etal. (2007),
Branda etal. (2010)
Cryptococcus saitoi Water (lake), glacial meltwater (river) Argentina Brazil, Italy Libkind etal. (2009),
Branda etal. (2010),
Brandão etal. (2011),
Libkind etal. (2014)
Cryptococcus tephrensis Water (lake), glacial meltwater (river) Brazil, Italy Branda etal. (2010,
2011)
Cryptococcus watticus Glacial meltwater (river) Italy Branda etal. (2010)
Gelidatrema spencermartinsiae Glacial meltwater (river) Argentina de García etal. (2010)
Kwoniella dendrophila Water (lake), phytotelmata of bromeliad tanks Brazil Brandão etal. (2011),
Morais etal. (2020)
Kwoniella heveanensis Phytotelmata of bromeliad tanks, rock pools Brazil Morais etal. (2020)
Kwoniella mangrovensis Phytotelmata of bromeliad tanks, rock pools Brazil Morais etal. (2020)
Naemateliaceae
Dimennazyma cisti-albidi Stagnant water Inácio etal. (2005)
Rhynchogastremaceae
Papiliotrema flavescens Phytotelmata of bromeliad tanks, water (wetland) Brazil, USA Fell etal. (2011),
Morais etal. (2020)
Papiliotrema laurentii Tropical freshwaters, phytotelmata of bromeliad tanks,
rock holes, surfaces of freshwater turtles Podocnemis
expansa, P. unifilis, polluted water (lake, river), water
(lake, pond, river, wetland), acidic water (lake), sedi-
ment (lake)
Argentina, Brazil,
Colombia, South
Africa, Slovakia,
USA,
van Uden and Ahearn
(1963), Woollett and
Hedrick (1970), Slá-
viková etal. (1992),
Sláviková and
Vadkertiová (1995b),
Russo etal. (2008),
Morais etal. (2010),
Fell etal. (2011), Van
Wyk etal. (2012),
Silva-Bedoya etal.
(2014), Brilhante
etal. (2016), Morais
etal. (2020)
Papiliotrema nemorosa Phytotelmata of bromeliad tanks (Very common species
on this subsrate) Brazil Morais etal. (2020)
Papiliotrema pseudoalba Water (wetland) USA Fell etal. (2011)
Papiliotrema rajasthanensis Phytotelmata of bromeliad tanks, water, sediment (lake) Brazil, Colombia Silva-Bedoya etal.
(2014), Morais etal.
(2020)
Papiliotrema taeanensis Freshwater Everglades USA Fell etal. (2011)
Papiliotrema terrestris Phytotelmata of bromeliad tanks, rock hole Brazil Morais etal. (2020)
Rhynchogastrema complexa Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Tremellaceae
Tremella globispora Phytotelmata of bromeliad tanks, rock holes Brazil Morais etal. (2020)
Trimorphomycetaceae
Saitozyma flava Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Saitozyma podzolica Phytotelmata of bromeliad tanks, surfaces of freshwater
turtles Podocnemis expansa, P. unifilis, water, sedi-
ment (lake)
Brazil, Colombia Morais etal. (2010,
2020), Silva-Bedoya
etal. (2014)
Trichosporonales

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Trichosporonaceae
Apiotrichum laibachii Water (lake, river), sediment (lake) Brazil, Colombia Medeiros etal. (2012),
Silva-Bedoya etal.
(2014)
Apiotrichum montevideense Water (river), tapwater Brazil, Slovenia Medeiros etal. (2012),
Novak Babič etal.
(2016)
Apiotrichum scarabaeorum Water (river), mud (river, canal) Brazil, Egypt Medeiros etal. (2012),
Moubasher etal.
(2018)
Cutaneotrichosporon cutaneum Water (lake, pond, river) Poland, Slovakia, USA van Uden and Ahearn
(1963), Czeczuga
etal. 1990, Sláviková
and Vadkertiová
(1995b)
Cutaneotrichosporon jirovecii Water, sediment (lake) Colombia Silva-Bedoya etal.
(2014)
Cutaneotrichosporon moniliiforme Water (lake) Antarctica Guého etal. (1992)
Cutaneotrichosporon mucoides Water (river) Poland Pietryczuk etal. (2018)
Trichosporon aculeatum Freshwater Japan Sugita (2011)
Trichosporon asahii Mud (river, canal), surfaces of freshwater turtles Podoc-
nemis unifilis, polluted water (river) Brazil, USA, Egypt Woollett and Hedrick
(1970), Morais etal.
(2010, 2020), Mou-
basher etal. (2018)
Trichosporon coremiiforme Water (lake), groundwater Brazil, Slovenia Brandão etal. (2011),
Novak Babič etal.
(2016)
Tremellomycetidae
Cystofilobasidiales
Cystofilobasidiaceae
Cystofilobasidium bisporidii Water (wetland) USA Fell etal. (2011)
Cystofilobasidium capitatum Water (lake), glacial meltwater (river), acidic water
(river) Argentina Brazil,
Slovenia Russo etal. (2008),
Branda etal. (2010),
Brandão etal. (2011),
Libkind etal. (2014)
Cystofilobasidium infirmominia-
tum Water (lake) Argentina, Brazil Libkind etal. (2003),
Brandão etal. (2011)
Cystofilobasidium lacus-mascardii Water (lake) Argentina Libkind etal. (2009)
Cystofilobasidium macerans Glacial melt water (river), water (lake), acidic water
(lake, river) Argentina de García etal. (2007),
Russo etal. (2008),
Libkind etal. (2009,
2014)
Mrakiaceae
Itersonilia pannonica Glacial melt water (river) Argentina de García etal. (2007)
Krasilnikovozyma huempii Water (river) Brazil Medeiros etal. (2012)
Mrakia aquatica Water (lake), Equisetum culms, foam (river, rivulet),
submerged decaying leaves (stream), stemflow and
throughfall of tree canopies, glacial meltwater (river)
Argentina, Austria,
Canada, India, Ire-
land, Poland, UK
Jones and Slooff
(1966), Bärlocher
(1987), Voglmayr
(1996), Harrington
(1997), Sridhar and
Karamchand (2009),
de García etal.
(2012)
Mrakia cryoconiti Alpine glacier cryoconite Austria Margesin and Fell
(2008)

Fungal Diversity
1 3
Table 13 (continued)
Taxon Substrate Distribution References
Mrakia frigida Glacial meltwater (river) Argentina de García etal. (2007)
Mrakia robertii Glacial meltwater (river) Argentina de García etal. (2012)
Tausonia pullulans Water (lake), polluted water (river), glacial meltwater
(river) Argentina¸ USA Woollett and Hedrick
(1970), Brandão etal.
(2011), de García
etal. (2012)
Udeniomyces megalosporus Glacial meltwater (river) Argentina de García etal. (2012)
Udeniomyces pyricola Freshwater Everglades, glacial meltwater (river), water
(pond) Argentina, Slovakia,
USA Sláviková and Vadk-
ertiová (1993), Fell
etal. (2011), de
García etal. (2012)
USTILAGINOMYCETES
Ustilaginomycetidae
Ustilaginales
Anthracoideaceae
Gymnocintractia samanensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Ustilaginaceae
Anthracocystis elionuri Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Anthracocystis everhartii Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Macalpinomyces spermophorus Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Moesziomyces aphidis Phytotelmata of bromeliad tanks, surfaces of freshwater
turtles Podocnemis expansa, water (wetland) Brazil, USA Morais etal. (2010,
2020), Fell etal.
(2011)
Moesziomyces parantarcticus Freshwater Everglades USA Fell etal. (2011)
Pseudozyma crassa Tapwater Slovenia Novak Babič etal.
(2016)
Moesziomyces rugulosus Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Pseudozyma flocculosa Freshwater Everglades USA Fell etal. (2011)
Pseudozyma hubeiensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Pseudozyma prolifica Surfaces of freshwater turtles Podocnemis expansa Brazil Morais etal. (2010)
Sporisorium andropogonis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Sporisorium graminicola Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Sporisorium trachypogonis-
plumosi Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Ustilago sparsa Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Ustilaginales genera incertae sedis
Anomalomyces panici Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Anomalomyces yakirrae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Kalmanozyma brasiliensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Kalmanozyma vetiveriae Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)
Langdonia jejuensis Phytotelmata of bromeliad tanks Brazil Morais etal. (2020)

Fungal Diversity
1 3
Microsporidia, now classified as fungi, are obligate,
intracellular, spore-forming parasites of a variety of hosts,
including insects, birds, fish, mollusks, platyhelminths,
while 17 species are known to infect human (Feng and Li
2019). Microsporidia are rarely lethal, generally causing
chronic debilitating diseases, resulting in reduced longev-
ity, fertility, weight and general vigour of their hosts. Micro-
sporidians may also be hyperparasites of other microsporidia
(Haine etal. 2004). Microsporidia evolutionary have lost
many of their mitochondrial and Golgi genes, with reduced
ribosomal RNAs size and genomes 2.5 to 11.6Mb in size,
encoding 1,848 to 3,266 protein (Corradi and Selman
2013). Microsporidial classification remains embryonic
but currently determined by sequence data (Vossbrinck and
Debrunner-Vossbrinck 2005; Alimov 2007; Tedersoo etal.
2018; Turland etal. 2018). Species identification is based
on spore morphology, their ultrastructure (especially their
unique resistant spore) and sequence data. The life cycle
commences with a polar filament injecting its spore con-
tents into a host cell (Cali etal. 2017). Fish microsporidia
can be broadly classified into two groups: those associ-
ated with xenoma (cyst-like) formation (e.g., Glugea) and
those associated with non‐xenoma forming microsporidia
(e.g., Nucleospora) (Shaw and Kent 1999; Kent etal.
2014). Microsporidia are transmitted both horizontally and
vertically.
The life cycle of a microsporidian is initiated by a spore
which on detecting a host cell swell resulting in the break-
down of the sporoplasmic membrane and the rupture of the
anchoring disk followed by the release of the polar tube and
the transfer of the sporoplasm into the host cell. Inside the
host cell more spores are produced and released into the
environment.
Hosts for freshwater microsporidia include bryozoans,
arthropods and fish, with nine genera and approximately
50 species described from copepods (Sprague and Vavra
1977; Bronnvall and Larsson 2001; Vávra etal. 2005). The
small planktonic crustacean Daphnia is the host of some 17
microsporidia (Table16). Unusual hosts for microsporidia
includes Alloglugea bufonis on metamorphosing tadpoles
of the toad Bufo marinus and Amblyospora camposi on
Culex renatoi, a bromeliad-inhabiting mosquito (Paperna
and Lainson 1995; Micieli etal. 2000). Hosts may harbour
more than one microsporidians e.g., the freshwater amphi-
pod Gammarus roeseli was infected by three species (Haine
etal. 2004). Fish are common hosts for microsporidians,
especially those reared under high stocking densities with
many recorded from salmon species.
Fig. 7 Conidia of some Ingoldian species collected from fresh-
water habitats in Thailand. A Flabellospora multiradiata. B Den-
drosporium lobatum. C Campylospora chaetocladia. D Anguillos-
pora longissima. E Lemonniera sp. F Brachiosphaera tropicalis. G
Tetracladium setigerum, H Sigmoidea sp. I Triscelophorus acumi-
natus. J Mycocentrospora sp. Scale bars: A, B, E, F = 10 μm, C,
I = 20μm, D = 30μm G = 25μm, H = 50μm, J = 30μm

Fungal Diversity
1 3
Table 14 Taxonomic outline of Ingoldian fungi recorded from freshwater habitats
Order Family Genus Species
ASCOMYCOTA
Dothideomycetes
Capnodiales Neodevriesiaceae Tripospermum T. myrti
Jahnulales Aliquandostipitaceae Brachiosphaera B. jamaicensis
B. tropicalis
Pleosporales Amniculicolaceae Anguillospora A. crassa
A. filiformis
A. furtiva
A. gigantea
A. mediocris
A. pseudolongissima
A. pulchella
A. rosea
A. rubescens
A. virginiana
Amniculicola A. aquatica
A. guttulata
A. immersa
A. lignicola
A. longissima
A. parva
Dothidotthiaceae Mycocentrospora M. acerina
M. angulata
M. aquatica
M. clavata
M. varians
Melanommataceae Tumularia T. aquatica
T. tuberculata
Tetraplosphaeriaceae Tetraploa T. abortiva
T. aquatica
T. aristata
T. ellisii
T. puzheheiensis
T. scheueri
T. thailandica
T. yunnanensis
Tubeufiales Tubeufiaceae Helicomyces H. ambiguus
H. bellus
H. chiayiensis
H. colligatus
H. geniculatus
H. hyalosporus
H. macrofilamentosus
H. roseus
H. scandens
H. torquatus
Helicosporium H. aquaticum
H. flavidum
H. flavisporum
H. flavum
H. gracile
H. lumbricopsis
H. murinum
H. panachaeum
H. setiferum
H. thailandense
H. vesicarium
Wiesneriomycetaceae Speiropsis S. hyalospora
S. pedatospora
S. scopiformis
Wiesneriomyces W. conjunctosporus
W. laurinus

Fungal Diversity
1 3
Table 14 (continued)
Order Family Genus Species
Dothideomycetes genera incertae sedis Clavariopsis C. aquatica
C. azlanii
C. brachycladia
C. bulbosa
Leotimycetes
Helotiales Discinellaceae Cladochasiella C. divergens
Fontanospora F. alternibrachiata
F. eccentrica
F. fusiramosa
Gyoerffyella G. craginiformis
G. entomobryoides
G. gemellipara
G. rotula
G. speciosa
G. tatrica
G. tricapillata
G. myrmecophagiformis
Lemonniera L. alabamensis
L. aquatica
L. centrosphaera
L. cornuta
L. filiformis
L. pseudofloscula
L. terrestris
L. yulongensis
Margaritispora M. aquatica
M. monticola
Tetrachaetum T. elegans
Varicosporium V. aquaticum
V. delicatum
V. elodeae
V. giganteum
V. helicosporum
V. macrosporum
V. scoparium
V. tricladiiforme
V. trimosum
Helotiaceae Articulospora A. atra
A. moniliformis
A. proliferata
A. tetracladia
Dimorphospora D. foliicola
Geniculospora G. grandis
G. inflata
Hyaloscyphaceae Arbusculina A. fragmentans
A. irregularis
Ploettnerulaceae Volucrispora V. aurantiaca
V. graminea
Tricladiaceae Mycofalcella M. calcarata
M. iqbalii

Fungal Diversity
1 3
Table 14 (continued)
Order Family Genus Species
Tricladium T. aciculum
T. alaskense
T. angulatum
T. anomalum
T. attenuatum
T. biappendiculatum
T. brunneum
T. castaneicola
T. caudatum
T. chaetocladium
T. curvisporum
T. fallax
T. fuscum
T. gracile
T. indicum
T. intermedium
T. kelleri
T. marylandicum
T. minutum
T. obesum
T. patulum
T. procerum
T. robustum
T. splendens
T. terrestre
T. varicosporioides
T. varium
Vibrisseaceae Vibrissea V. decolorans
V. dura
V. filisporia
V. flavovirens
V. guernisacii
V. leptospora
V. norvegica
V. sporogyra
V. truncorum
V. vibrisseoides
Helotiales genera incertae sedis Filosporella F. annelidica
F. aquatica
F. exilis
F. fistucella
F. pinquis
F. versimorpha
Tetracladium T. apiense
T. breve
T. furcatum
T. globosum
T. maxilliforme
T. nainitalense
T. palmatum
T. setigerum
T. marchalianum
Leotiales Leotiales genera incertae sedis Alatospora A. acuminata
A. constricta
A. crassipes
A. flagellata
A. pulchella
Flagellospora F. curvula
F. fusarioides
F. leucorhynchos
F. minuta
F. saccata
F. stricta
Leotiomycetes genera incertae sedis Gorgomyces G. honrubiae
Orbiliomycetes

Fungal Diversity
1 3
Table 14 (continued)
Order Family Genus Species
Orbiliales Orbiliaceae Dactylella T. microaquatica
T. submersa
T. yunnanensis
Dicranidion D. fragile
D. gracile
Trinacrium T. incurvum
T. indicum
T. robustum
T. subtile
Sordariomycetes
Chaetosphaeriales Chaetosphaeriales genera incertae sedis Nawawia N. filiformis
N. oviformis
N. quadrisetulata
Glomerellales Reticulascaceae Cylindrotrichum C. aquaticum
C. clavatum
C. gorii
C. helisciforme
C. oligospermum
C. submersum
C. zignoellae
Hypocreales Nectriaceae Aquanectria A. devians
A. jacinthicolor
A. penicillioides
A. submersa
A. tenuispora
Heliscus H. tentaculus
Neonectria N. coccinea
N. lugdunensis
Microascales Halosphaeriaceae Clavatospora C. filiformis
C. longibrachiata
C. tentacula
Sigmoidea S. aurantiaca
S. contorta
S. praelonga
S. prolifera
Sordariales Sordariales genera incertae sedis Lunulospora L. curvula
L. cymbiformis
Hypocreomycetidae genera incertae sedis Campylospora C. brasiliensis
C. chaetocladia
C. filicladia
C. parvula
Ascomycota incertae sedis Alatosessilispora A. bibrachiata
Arborispora A. dolichovirga
A. palma
A. paupera
Blodgettia B. indica
Calcarispora C. hiemalis
Casaresia C. sphagnorum
Cladoconidium C. articulatum
Clavariana C. aquatica
Condylospora C. flexuosa
C. gigantea
C. spumigena
Culicidospora C. aquatica
C. gravida
Curucispora C. ombrogena
C. ponapensis
Colispora C. cavincola
C. curvata
C. elongata

Fungal Diversity
1 3
Table 14 (continued)
Order Family Genus Species
Dendrospora D. erecta
D. fastuosa
D. fusca
D. juncicola
D. moniliformis
D. nana
D. polymorpha
D. tenella
D. torulosa
D. yessemreddyi
Dendrosporium D. lobatum
Diplocladiella D. appendiculata
D. aquatica
D. scalaroides
D. tricladioides
Dwayaangam D. colodena
D. cornuta
D. dichotoma
Flabellocladia F. gigantea
Flabellospora F. acuminata
F. amphibia
F. crassa
F. multiradiata
F. tetracladia
F. verticillata
Gemmulina G. botryosa
Heliscella H. stellata
H. stellatacula
Isthmolongispora I. asymmetrica
I. geniculata
I. lanceata
I. minima
I. rotundata
Isthmotricladia I. britannica
I. gombakiensis
I. laeensis
Lateriramulosa L. quadriradiata
L. uniinflata
Mirandina M. corticola
M. taiwanensis
M. uncinata
Polycladium P. equiseti
Pseudoanguillospora P. gracilis
P. prolifera
P. stricta
Pyramidospora P. casuarinae
P. constricta
P. densa
P. fluminea
P. herculiformis
P. quadricellularis
P. ramificata
P. robusta
P. stellata
Scutisporus S. brunneus
Subulispora S. biappendiculata
S. longirostrata
S. malaysiana
S. procurvata
Sympodiocladium S. frondosum
Tetrabrachium T. elegans
Tetrabrunneospora T. ellisii

Fungal Diversity
1 3
Table 14 (continued)
Order Family Genus Species
Titaea T. callispora
T. clarkeae
Tricladiopsis T. flagelliformis
T. foliosa
Tricellula T. aquatica
T. inaequalis
T. ornata
Trifurcospora T. irregularis
Triglyphium T. alabamense
Triramulispora T. gracilis
T. obclavata
Triscelophorus T. acuminatus
T. curviramifer
T. konajensis
T. magnificus
T. monosporus
T. ponapensis
T. septatus
Trisulcosporium T. acerinum
BASIDIOMYCOTA
Agaricomycetes
Atheliales Atheliaceae Taeniospora T. gracilis
T. gracilis var. gracilis
T. nasifera
Cantharellales Hydnaceae Ingoldiella I. fibulata
I. nutans
Agaricomycetes genera incertae sedis Dendrosporomyces D. prolifer
D. splendens
Fibulotaeniella F. canadensis
Titaeella T. capnophila
Tricladiomyces T. geniculatus
T. malaysianus
Classiculomycetes
Classiculales Classiculaceae Classicula C. fluitans
Jaculispora J. submersa
Microbotryomycetes
Kriegeriales Camptobasidiaceae Crucella C. subtilis
Ustilaginomycetes
Ustilaginales Ustilaginaceae Pseudozyma P. crassa
P. flocculosa
P. hubeiensis
Basidiomycota genera incertae sedis Arcispora A. bisagittaria

Fungal Diversity
1 3
Table 15 Classification of fungal basal clades recorded from freshwater environment including information on habitats, substrates/hosts, and
geographical distribution
Taxon Substrate/Host Country References
APHELIDIOMYCOTA
APHELIDIOMYCETES
Aphelidiales
Aphelidiaceae
Amoeboaphelidium achnanthis Parasitoid of Achnanthes lanceolata Hungary Letcher and Powell (2019)
Amoeboaphelidium chlorellavorum Parasitoid of Chlorella Russia Letcher and Powell (2019)
Amoeboaphelidium occidentale Scenedesmus dimorphus (pond) USA Letcher etal. (2015b)
Amoeboaphelidium protococcarum Ponds USA Letcher etal. (2013)
Amoeboaphelidium radiatum Parasitoid of Kirchniriella and Ankistrodesmus Russia Letcher and Powell (2019)
Aphelidium arduennense Roadside ditch Belgium Tcvetkova etal. (2019)
Aphelidium chaetophorae Parasitoid of Chaetophora elegans Hungary Letcher and Powell (2019)
Aphelidium chlorococcorum Parasite of representatives of various genera of chloro-
coccus algae – Gromov and Mamkaeva
(1975)
Aphelidium chlorococcorum f. chlo-
rococcorum Parasite of representatives of various genera of chloro-
coccus algae – Schnepf etal. (1971),
Letcher and Powell
(2019)
Aphelidium chlorococcorum f. majus Parasitoid of Kirchneriella obesa Letcher and Powell (2019)
Aphelidium collabens Parasitoid of Coccomyxa sp. (pond) Japan Seto etal. (2020a)
Aphelidium desmodesmi Parasitoid of green alga Desmodesmus armatus USA Letcher etal. (2017)
Aphelidium deformans Parasitoid of Coleochaeta Germany Zopf (1885)
Aphelidium melosirae Parasitoid of Melosira varians: diatom alga Melosira
varians Hungary Letcher and Powell (2019)
Aphelidium tribonematis Parasitoid of Tribonema Hungary Letcher and Powell (2019)
Paraphelidium tribonemae Roadside ditches Russia Karpov etal. (2017b)
Paraphelidium letcheri Dried stream Russia Karpov etal. (2017c)
BLASTOCLADIOMYCOTA
BLASTOCLADIOMYCETES
Blastocladiales
Blastocladiaceae
Allomyces anomalus Rozella allomycis isolated from roadside ditch Hattiesburg Powell etal. (2017)
Allomyces arbusculus Water on dead Diptera India Butler (1911)
Allomyces catenoides Tropical fish aquarium USA Sparrow (1964)
Allomyces javanicus Water, mud from lake Indonesia, Egypt Ji and Dayal (1971), Ali
and Abdel-Raheem
(2003)
Allomyces neomoniliformis Lake India Dubey etal. (2016)
Allomyces macrogynus Mud Egypt Ali and Abdel-Raheem
(2003)
Allomyces moniliformis Mud Egypt Ali and Abdel-Raheem
(2003)
Blastocladia arborata Styles of submerged flowers of Caryopteris and myrta-
ceous fruit India, Egypt Dasgupta and John (1988),
El-Hissy etal. (1996)
Blastocladia angusta Weeping white birch twigs, rose hips, lake Denmark, Egypt El-Hissy etal. (1996)
Blastocladia aspergilloides Lake Australia Nagai and Kobayashi
(1995)
Blastocladia bonaerensis Eutrophic water litter Argentina Steciow and Marano
(2006)
Blastocladia caduca Stamens of submerged flowers of Crateva religiosa India Dasgupta and John (1988)
Blastocladia coronata Submerged fruit of Eugenia jambolana India Dasgupta and John (1988)
Blastocladia didyma Submerged fruit of Physalis peruviana India Dasgupta and John (1988)
Blastocladia elegans Submerged fruit of Mangifera indica India Dasgupta and John (1988)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Blastocladia excelsa Ripe guava fruit bait, tanks India Dasgupta and John (1988)
Blastocladia filamentosa Eugenia jambolana fruit India Dasgupta and John (1988)
Blastocladia fruticosa Submerged fruits of Eugenia jambolana India Dasgupta and John (1988)
Blastocladia fusiformis Submerged stamens and pistils of flower of Crateva
religiosa India Dasgupta and John (1988)
Blastocladia globosa Pond water India Manoharachary etal.
(1983)
Blastocladia glomerata Sunken twigs of Aesculus UK Khulbe (2001)
Blastocladia gracilis River, lake, pond, pool Denmark, UK Waterhouse (1942)
Blastocladia heterosporangia Submerged fruit on Vitis vinifera India Dasgupta and John (1988)
Blastocladia incrassata Water Japan, USA Hsiao (1969)
Blastocladia mammillata Submerged fruit of Pyrus India Dasgupta and John (1988)
Blastocladia pileota Submerged fruit of Pyrus India Dasgupta and John (1988)
Blastocladia pringsheimii Water on rotten fruits of Malus, tomato skin (river) Germany, USA,
UK Waterhouse 1942, Hsiao
(1969), Rooney and
McKnight (1972)
Blastocladia pusilla Submerged fruit of Mangifera indica India Dasgupta and John (1988)
Blastocladia ramosa Submerged twigs, water (river) Poland, USA, UK Waterhouse (1942),
Rooney and McKnight
(1972), Czeczuga etal.
(1990)
Blastocladia rostrata Lake USA Hsiao (1969)
Blastocladia sessilis Various submerged fruits India Dasgupta and John (1988)
Blastocladia sparrowii –USA Hsiao (1969)
Blastocladia spiciformis Psidium India Dasgupta and John (1988)
Blastocladia strangulata Aphidoidea USA Barrett (1912)
Blastocladia truncata Submerged fruit of Malus, submerged plant (river) North America,
UK Sparrow (1932), Water-
house (1942)
Blastocladiopsis parva Water reservoir, lake, water (stream) Poland, USA Hsiao (1969), Czeczuga
etal. (1990), Kiziewicz
and Kurzątkowska
(2004)
Catenariaceae
Catenophlyctis peltata Snake skin in water Cuba Karling (1968a)
Catenophlyctis variabilis Water and sediment (reservoir), lake Brazil, Egypt,
Poland, UK Godlewska etal. (2009)
Nematoceromyces ramosus Parasitic in eggs of Dicrotendipes modestus (stream) USA Martin (1978)
Paraphysodermataceae
Paraphysoderma sedebokerense Haematococcus pluvialis Israel James etal. (2012)
Catenomycetales
Catenomycetaceae
Catenomyces persicinus Grasses in water samples USA Hanson (1944b)
Coelomomycetaceae
Coelomomyces quadrangulatus Anopheles quadrimaculatus larvae, lake USA Porter etal. (2011)
Coelomomyces punctatus Anopheles quadrimaculatus larvae, lake USA Porter etal. (2011)
Coelomomyces lativittatus Anopheles spp. larvae, pond USA Porter etal. (2011)
Coelomycidium ephemerae Cloeon rufulum larvae Former Czechoslo-
vakia Weiser (1984)
Coelomycidium simulii Simulium larvae, stream France Debaisieux (1919)
Blastocladiomycetes genus incertae
sedis

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Microallomyces dendroideus Pond sediment Costa Rica Emerson and Robertson
(1974)
CHYTRIDIOMYCOTA
CHYTRIDIOMYCETES
Chytridiales
Asterophlyctaceae
Asterophlyctis michiganensis Lake USA Letcher etal. (2018)
Asterophlyctis sarcoptoides Pond USA Letcher etal. (2018)
Wheelerophlyctis interior Aquatic sample USA Letcher etal. (2018)
Wheelerophlyctis interiexterior Pond USA Letcher etal. (2018)
Chytridiaceae
Chytridium acuminatum Lake (on cellophane, baby hairs, pine pollen, Ulothrix
sp) Oogonia of Oedogonium rothii, possibly also on
O. echinospermum, zygospores of Mougeotia sp.
USA, Germany,
Russia Cornu (1872), Sorokin
(1883), Rooney and
McKnight (1972)
Chytridium adpressum Ditch India Dasgupta and John (1990)
Chytridium aggregatum Saprophytic on Spirogyra crassa, Oedogonium sp., and
Cladophora sp. USA Karling (1938)
Chytridium apophysatum Ditch India Dasgupta and John (1990)
Chytridium amphoroideum Hydrodictyon UK Playfair (1917)
Chytridium closterii Water India Dasgupta and John (1990)
Chytridium cocconeidis Living cells of Cocconeis pediculus, epiphytic on UK Canter (1947a)
Cladophora (lake)
Chytridium coleochaetes Parasitic on oogonia of Coleochaete pulvinata (tarn) Germany, UK Nowakowski (1876),
Canter (1960a)
Chytridium corniculatum Pine pollen, freshwater sand Greenland Playfair (1917)
Chytridium cornutum Heterocysts of Aphanizomenon flos-aquae (pond) Czechoslovakia Canter (1963a)
Chytridium cresentum Ditch India Dasgupta (1982)
Chytridium deltanum Parasitic,Oocystisspp., phytoplankton, lake Canada Masters (1971a)
Chytridium hyperparasiticum Parasitic, sporangium ofRhizophlyctis rosea, soil in
water Greenland Letcher and Powell (2012)
Chytridium isthmiophilum Parasitic on Staurodesmus curoatus (lake) UK Canter (1960b)
Chytridium lagenula Parasitic on Tribonema sp. (pool) UK Sparrow (1957)
Chytridium lecythi Parasitic on rhizopod Lecythium Byalinum (stream) UK Ingold (1941)
Chytridium marylandicum Saprobic, matrix of Botryococcus, Blue Pond USA Masters (1971b)
Chytridium neopapillatum Saprobic, pine pollen, sand in water Greenland, USA Masters (1971b)
Chytridium oedogonii Oedogonium (lake) UK, USA Couch (1938) Canter
(1950)
Chytridium olla Oedogonium Germany Powell and Letcher (2014)
Chytridium oocystidis Oocystis lacustris Switzerland Huber-Pestalozzi (1944)
Chytridium ottariense Saprobic, Volvox, chitin, snake skin, impoundment USA Roane (1973)
Chytridium rhizophydii Chitin and pollen baits (pond) USA Davis etal. (2018)
Chytridium reniforme Water, Spirogyra, ditch India Dasgupta and John (1988)
Chytridium schenkii Water Dasgupta (1982)
Chytridium sparrowii Water Dasgupta and John (1988)
Chytridium telmatoskenae Parasitic, Telmatoskene mucosa, pond, Volary, central Bohemia Fott (1957)
Chytridium versatile Nitzschia sigmoidea (lake) UK Canter (1947a)
Chytridium xylophilum Water (river) Poland Czeczuga etal. (1990)
Dendrochytridium crassum Tree-canopy detritus flooded with water Australia Letcher etal. (2014)
Dinochytrium kinnereticum Lake, freshwater springs, standing water pools Israel, Oman Leshem etal. (2016), Has-
sett etal. (2020)
Irineochytrium annulatum Water (lake) USA Dogma (1969a)
Polyphlyctis willoughbyi Permanently submerged river mud Australia Letcher etal. (2018)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Polyphlyctis unispina Lake, bog USA Letcher etal. (2018)
Zopfochytrium polystomum Submerged shoreline vegetation, mud, and water, Lake USA Powell etal. (2018)
Chytriomycetaceae
Avachytrium platense Lake Argentina Vélez etal. (2013)
Chytriomyces appendiculatus Saprophytic on chitinous substrata in freshwater and
muck soil, water column and sediment (reservoir),
mud (lake)
Brazil, UK, USA Willoughby (1961),
Letcher and Powell
(2002), Davis etal.
(2019), Jerônimo and
Pires-Zottarelli (2020)
Chytriomyces aureus Unidentified insects in fresh water, mud (lake) Brazil, UK, USA Karling (1945a), Wil-
loughby (1961)
Chytriomyces closterii Weakly parasitic on Closterium rostratum, ditch USA Letcher and Powell (2002)
Chytriomyces confervae Parasitic on Tribonema Sweden, Hungary,
USA, Germany,
UK, Canada,
Poland
Letcher and Powell (2002)
Chytriomyces cosmaridis Water Poland Letcher and Powell (2002)
Chytriomyces fructicosus Saprophytic in cellulosic substrata in soil and water USA Letcher and Powell (2002)
Chytriomyces heliozoicola Lake plankton, parasitic on helizoan Raphidiocystis
lemani, Esthwaite Water and windermere UK Letcher and Powell (2002)
Chytriomyces hyalinus Water, saprobes on chitin, pollen Brazil, China, UK,
USA Letcher and Powell
(2002), Davis etal.
(2019)
Chytriomyces laevis Soil–water culture, parasitic on Pythium sp, growing on
snake skin Panama Letcher and Powell (2002)
Chytriomyces lucidus Saprophytic in cellulosic substrata in soil and water USA Letcher and Powell (2002)
Chytriomyces nagatoroensis Weakly parasitic on Spirogyra sp. Japan Letcher and Powell (2002)
Chytriomyces rotoruaensis In watered mud samples, saprophytic on chitin New Zealand Letcher and Powell (2002)
Chytriomyces stellatus Saprophytic on chitinous substrata in freshwater and
muck soil USA, Brazil Letcher and Powell (2002)
Chytriomyces tabellariae Water and mud, parasitic on Tabellaria species Switzerland, UK Letcher and Powell (2002)
Entophlyctis apiculata Agricultural pond, parasitized cells of Korea Shin etal. (2001)
Chlamydomonas sp.
Entophlyctis aurea Saprobic on various Poaceae, Triticum aestivum, Avena
sativa, and Zea mays leaves, regenerated cellulose
film, filter paper, and lens paper submerged in water
Canada, UK Haskins (1946)
Entophlyctis bulbigera Ditch, cells of Spirogyra filaments (pool) India, UK Richards (1956), Dasgupta
and John (1990)
Entophlyctis caudiformis Closterium, tank India Dasgupta and John (1990)
Entophlyctis confervae-glomeratae Aquatic, cells of filaments of Oedogonium and Spiro-
gyra (pond) Germany, UK Sparrow (1943), Richards
(1956)
Entophlyctis crenata On Vallisneria: North freshwater New Zealand Karling (1968b)
Entophlyctis lobata Shrimp chitin bait, littoral of Derwentwater UK Willoughby and Townley
(1961)
Entophlyctis luteolus Onion skin and Utricularia, bogs and acid lake, mud
pond USA Longcore (1995)
Entophlyctis mammilliformis Oospores of Spirogyra, pond India Dasgupta and John (1990)
Entophlyctis molesta Stylosphaeridium stipitatum, phytoplankton, English
lakes UK Barr (1971)
Entophlyctis reticulospora Closterium moniliferum, swampy area USA Cook (1966)
Entophlyctis sphaerioides Reproductive body of Spirogyra, ditch India Dasgupta and John (1990)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Entophlyctis texana Moribund cells of the aquatic plantEriocaulon USA Karling (1941)
Fayochytriomyces spinosus Onion skin placed in a muck sample taken from a
temporary pond USA Davis etal. (2015)
Obelidium hamatum Chironomidae larvae, pond USA Blackwell etal. (2012)
Obelidium megarhizum From submerged mud: river UK, USA Blackwell etal. (2012)
Obelidium mucronatum On the empty skin of a mosquito larva, pond, strips of
purified shrimp chitin (water on outdoor tub) USA, New Zea-
land Karling (1967), Blackwell
etal. (2012)
Odontochytrium milleri Pollen and chitin baits (pond), bog Argentina, USA Vélez etal. (2013), Davis
etal. (2019)
Pendulichytrium sphaericum Aulacoseira granulata, lake Japan Seto and Degawa (2018)
Physocladia obscura Growing on onion skin bait placed in water from Bry-
ant’s Bog USA Davis etal. (2019)
Podochytrium chitinophilum From submerged mud, wastewater UK Willoughby and Townley
(1961)
Podochytrium clavatum Parasitic on diatoms Poland Kadłubowska (2014)
Podochytrium cornutum Planktonic diatom Stephanadiscus niagarae, lake USA Sparrow (1951), Canter
(1969)
Podochytrium dentatum Saprobic on chitin, lake, pond USA Longcore (1992)
Podochytrium ellerbeckense Water lily leaf stalks, garden pond UK Longcore (1992)
Podochytrium emmanuelense Melosira varians and on Nitzschia UK Paterson (1958a)
Podochytrium lanceolatum Melosira varians UK Canter (1953)
Rhizidium brasiliense Insect remnants and exuviae in water (stream) Brazil Karling (1944b)
Rhizidium elongatum Pond water, chitinous substrata (river) India, USA Karling (1949b), Dayal
and Kiran (1980)
Rhizidium variabile Spirogyra (pond) UK Canter (1947a)
Rhizidium varians Pond water, chitinous material in water (river) India, USA Karling (1949b), Dayal
and Kiran (1980)
Rhizidium verrucosum Bits of insects and exuviae, pieces of onion skin and
grass leaves in a water sample Brazil Karling (1944b)
Rhizoclosmatium aurantiacum Saprotrophic in water on insect exuviae, chitin, cel-
lulose, and pollen Denmark, USA Paterson (1967), Powell
etal. (2019)
Rhizoclosmatium globosum Saprotrophic on insect exuviae, chitin, cellulose, pol-
len, keratin, and depleted filaments of Aphanomyces,
purified shrimp chitin from a rainwater culture in an
outdoor tub with Nitella
Denmark, New
Zealand, USA Karling (1967), Davis
etal. (2019), Powell
etal. 2019)
Rhizoclosmatium hyalinum Saprotrophic on insect exuviae and chitin, water in
outdoor tub with Elodea New Zealand Karling (1967), Powell
etal. (2019)
Rhizoclosmatium sparsum Saprotrophic on chitin and pollen, spring water, river USA Powell etal. (2019)
Rhizoclosmatium umbonatum Saprotrophic on chitin and pollen, water edge USA Powell etal. (2019)
Rhizoclosmatium persicum Saprotrophic on chitin and pollen, dam spillway, lake Powell etal. (2019)
Rhizoclosmatium pessaminum Saprotrophic on chitin and pollen, edge of lake, pond USA Powell etal. (2019)
Siphonaria petersenii Saprophytic inside the exuviae of mayflies and other
insects, lake, Salmon Pond USA, Brazil Karling (1945b), Simmons
etal. (2020)
Siphonaria sparrowii Mud (pond), purified shrimp chitin in water (lake) USA, New Zea-
land Karling (1967), Simmons
etal. (2020)
Siphonaria variabilis Purified shrimp chitin in water (lake) New Zealand Karling (1967)
Phlyctochytriaceae
Phlyctochytrium aequale Melosira sp. from a roadside pool Iceland Johnson (1968)
Phlyctochytrium aureliae Bog USA Ajello (1945), Davis etal.
(2019)
Phlyctochytrium apophysatum Phlyctochytrium dissolutum, Closterium, tank, garden India Dasgupta and John (1990)
Phlyctochytrium bullatum Pond, on Cladophora USA, Iceland Johnson (1968)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Phlyctochytrium circulidentatum Saprobic on pollen and moribund Closterium, bottom
debris, lake Germany Umphlett and Koch (1969)
Phlyctochytrium dentiferum Dead Cladophora, lake USA Sparrow (1937a)
Phlyctochytrium dissolutum Closterium, tank, garden India Dasgupta and John (1990)
Phlyctochytrium furcatum Test of a testate amoeba in water collected from Bry-
ant’s Bog USA Sparrow (1966), Davis
etal. (2019)
Phlyctochytrium incrustans Bog USA Sparrow and Lange (1977)
Phlyctochytrium irregulare Mud (lake) UK Willoughby (1961)
Phlyctochytrium lippsii Parasites of Ascobolus, Ascophanus, Lasiobolus and
Sordaria, reservoir USA Lohman (1942)
Phlyctochytrium longicollum Parasites of Euglena spp., bog USA Sparrow (1952)
Phlyctochytrium macrosporum Lake USA Sparrow (1978)
Phlyctochytrium multidentatum Saprobic on pollen, water USA Umphlett and Koch (1969)
Phlyctochytrium mucronatum Parasitic on Closterium pritchardianum, bog, pollen
bait (pond) UK, USA Canter (1949a), Davis
etal. (2018)
Phlyctochytrium parasitans Parasites of eggs of Achlya flagellata, damp soil Hispaniola
(Republic of
Haiti and the
Dominican
Republic)
Sparrow and Dogma
(1973)
Phlyctochytrium planicorne Growing on a filamentous conjugating green alga, bog,
pond, mud (lake) UK, USA Willoughby (1961),
Letcher and Powell
(2005a)
Phlyctochytrium powhatanense Impoundment USA Johnson (1969)
Phlyctochytrium proliferum Parasitic on Chlamydomonas (pond) UK Ingold (1941)
Phlyctochytrium urceolare Dead Cladophora, lake Sparrow (1938)
Phlyctorhizaceae
Phlyctorhiza endogena Remains of mosquitoes, mayflies, dragonflies and gnats
found on streams, Peep Pond USA Hanson (1946)
Pseudorhizidiaceae
Pseudorhizidium endosporangia-
tum Pine pollen in water (bog) USA Davis etal. (2019)
Scherffeliomycetaceae
Scherffeliomyces parasitans Freshwater UK Sparrow (1936)
Chytridiales genus incertae sedis
Delfinachytrium mesopotamicum Marsh in semipermanent stream Argentina Vélez etal. (2013)
Zygorhizidiales
Zygorhizidiaceae
Zygorhizidium cystogenum Dinobryon divergens and Uroglena americana cysts
(lake) UK Canter (1963b)
Zygorhizidium parallelosede Parasitic on Elakatothrix gelatinosa and Ankistrodesmus
sp. (lake) UK Canter (1954), Canter and
Lund (1968)
Zygorhizidium planktonicum Asterionella formosa and Synedra ascus var. angustis-
sima UK Canter and Lund (1953),
Paterson 1958b)
Zygorhizidium willei Cylindrocystis brebissoni, surface of a moist rock, lake Japan Seto etal. (2020b)
Zygophlyctidales
Zygophlyctidaceae
Zygophlyctis asterionellae Asterionella formosa, lake Japan Seto etal. (2020b)
Zygophlyctis melosirae Aulacoseira subarctica, lake, parasitic on Melosira
italica subsp. subarctica UK Paterson (1958b), Seto
etal. (2020b)
Zygophlyctis planktonica Ulnaria sp, lake Germany Seto etal. (2020b)
Polyphagales
Polyphagaceae

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Polyphagus arnaudovii Hyponeustonic Bulgaria Doweld (2014)
Polyphagus asymmetricus Hyponeustonic, on Botrydiopsis Bulgaria Doweld (2014)
Polyphagus elegans Parasitizing Ulothrix mucosum UK Canter (1963b)
Polyphagus euglenae Ponds Germany Johns (1964)
Polyphagus laevis Cysts of Euglena viridis and of Chlamydomonas Northern hemi-
sphere Bartsch (1945)
Polyphagus ramosus Sphaerocystis schroeteri Switzerland Johns (1964)
Polyphagus serpentinus Lakes: parasitizing Spondylosium planum United Kingdom Canter (1963b)
Polyphagus starrii Parasitic, Eudorina, Pandorina, pond USA Johns (1964)
Polyphagus sinicus Draparnaldia China Doweld (2014)
Chytridiomycetes family incertae
sedis
Sparrowiaceae
Sparrowia subcruciformis Sporangia of Rhizophydium coronum, Chytriomyces
poculatus, Chytriomyces annularus and Rhizophlyctis
rosea, lake
USA Papa and Cruz-Papa
(2020)
Chytridiomycetes genera incertae sedis
Bertramia marionensis Parasite of Chironomidae (Diptera) larvae, lake Canada Weiser and McCauley
(1974)
Blyttiomyces conicus Stream Norway Johnson (1977)
Blyttiomyces helicus Lake, bog USA Sparrow and Barr (1955),
Davis etal. (2019)
Blyttiomyces laevis Parasitic on filaments of Zygnema sp., bog USA Sparrow (1952)
Blyttiomyces lenis Parasitic on zygospore of Spirogyra, ditch India Dasgupta and John (1990)
Blyttiomyces spinosus Spirogyra, aquatic habitat India Dasgupta and John (1990)
Blyttiomyces spinulosus Zygospores of Sirogonium, ditch China, Japan,
Philippines,
Cuba,
USA,
Argentina
Blackwell etal. (2011)
Canteria apophysata Clay pond, on Mougeotia UK Karling (1971)
Dangeardia echinulata Vitreochlamys aulata, pool Poland Batko (1970)
Dangeardia laevis Parasitic in Gloeodinium sp. USA Sparrow and Barr (1955)
Dangeardia mamillata Parasite on Pandorina morum, small pond, on Yamag-
ishiella unicocca, oligo-mesotrophic lake Poland, Germany,
UK Canter (1946), Van den
Wyngaert etal. (2018)
Dangeardia ovata Parasitic in Sphaerocystis schroeteri, lake USA Paterson (1958a)
Dangeardia sporapiculata Chlamydomonas, lake Austria Geitler (1963)
Dangeardiana eudorinae Parasitic on oospores of Eudorina elegans, water Bulgaria Valkanov (1964)
Dangeardiana leptorrhiza Zygnema, swampy pool USA Johns (1956)
Dictyomorpha dioica Parasitic of oomycetes, lake USA Mullins (1961)
Dictyomorpha pythiensis Parasitic on Pythium aphanidermatum, pond India Sarkar and Dayal (1988)
Ichthyochytrium vulgare Parasite of freshwater fishes Cyprinus Germany Korting (1983)
Loborhiza metzneri River USA Hanson (1944a)
Macrochytrium botrydioides Debris and skeletonized leaves found in a stream USA Crasemann (1954)
Mitochytridium ramosum Bog, swampy ground France Dangeard (1911)
Mitochytridium regale Pond Poland Hassan (1982)
Mucophilus cyprini Cyprinus: freshwater fish Germany Červinka etal. (1974)
Perolpidium utriculiforme Parasitic on Closterium lunula, C. costatum, C. dianeae UK Canter (1949b)
Pseudopileum unum Parasitizing cysts of Mallomonas, Lake UK Canter (1963b)
Rhizosiphon akinetovorax Parasitic on Anabaena affinis var. intermedia UK Gleason etal. (2014)
Rhizosiphon akinetum Parasitic on Anabaena affinis var. intermedia UK Gleason etal. (2014)
Rhizosiphon anabaenae Parasitic on Anabaena affinis var. intermedia Sweden Gleason etal. (2014)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Rhizosiphon crassum Parasitic on Cyanophyceae, parasitic on Anabaena
solitaria (tarn) Hungary, UK Canter and Lund (1968),
Gleason etal. (2014)
Rhopalophlyctis sarcoptoides Insect skeletons from freshwater Brazil, USA Karling (1945a), Davis
etal. (2019)
Saccomyces dangeardii Small water bodies Ukraine Demchenko (2019)
Septosperma anomalum Parasites of freshwater chytrids, swamp soils, parasitic
on Asterionella (tarn) USA, UK, Hun-
gary Canter and Lund (1953),
Seymour (1971)
Septosperma rhizophidii Parasites of freshwater chytrids, swamp soils USA, UK, New
Zealand Seymour (1971)
Septosperma irregulare Water New Zealand Dogma (1974)
Septosperma multiforme Parasites of freshwater chytrids, swamp soils, quiescent
cells and immature cysts of Amphichrysis compressa
(pool), immature cysts of Mallomonas sp. (lake)
USA, UK, New
Zealand Canter (1963b), Seymour
(1971)
Sorokinocystis mirabilis Submerged decaying wood Italy Saccardo (1888)
Sporophlyctidium africanum Parasitic on Protoderma sp, from water cultures of
vegetable matter Morocco Sparrow (1933b)
Sporophlyctidium neustonicum Lake water USA Sparrow (1978)
Sporophlyctis chinensis Draparnaldia China Sparrow (1960)
Sporophlyctis rostrata Draparnaldia and Chaetophora Russia Serbinow (1900)
Truittella setifera Soil and water cultures USA Karling (1949a)
Volvorax ingoldii Coenobium of Eudorina, reservoir Germany, UK Doweld (2013), Van den
Wyngaert etal. (2018)
Zygochytrium aurantiacum Dead insect, water bodies Poland Czeczuga and Godlewska
(2014)
CLADOCHYTRIOMYCETES
Cladochytriales
Catenochytridiaceae
Catenochytridium carolinianum Mud (lake) UK Willoughby (1961)
Catenochytridium hemicysti Moribund desmid West Virginia Barr etal. (1987)
Cladochytriaceae
Cladochytrium aneurae Spring Poland Czeczuga etal. (2005)
Cladochytrium aureum Pond Poland Czeczuga etal. (2005)
Cladochytrium aurantiacum Pond, fragments of leaves, stems and roots of Avena and
Triticum (canal) Poland, UK Czeczuga etal. (2005)
Cladochytrium cornutum Submerged plants France de Wildeman (1896)
Cladochytrium crassum Decaying vegetation, lake USA Hillegas (1941)
Cladochytrium granulatum Decaying vegetation Brazil Karling (1944a)
Cladochytrium hyalinum Dead grass, stream USA Berdan (1941)
Cladochytrium irregulare Submerged plants France de Wildeman (1895)
Cladochytrium menyanthis Parasitic on leaves of Menyanthes trifoliata (bog, ditch) USA, UK Sparrow (1952, 1957)
Cladochytrium taianum Spring Poland Czeczuga etal. (2005)
Cladochytrium replicatum Pond, lake, spring, water column and sediment (reser-
voir), mud (lake), plant debris (lake) Brazil, Egypt, UK,
USA Karling (1931), Richards
(1956), Willoughby
(1961), Jerônimo and
Pires-Zottarelli (2020)
Cladochytrium salsuginosum Blackish water Poland Batko and Hassan (1986)
Cladochytrium setigerum Sediment (reservoir) Brazil Jerônimo and Pires-Zotta-
relli (2020)
Cladochytrium tenue Acorus calamus and Iris pseudacorus, water Brazil, Poland Nowakowski (1877), Jesus
etal. (2021)
Endochytriaceae
Diplophlyctis amazonensis Submerged decaying plant debris, mud (lake) Brazil, UK Karling (1944c), Wil-
loughby (1961)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Diplophlyctis asteroidea Mud-acid bogs USA Dogma (1976)
Diplophlyctis buttermerensis Submerged mud, ditch UK, USA Dogma (1969b)
Diplophlyctis chitinophila Submerged mud UK Willoughby (1962)
Diplophlyctis complicata Submerged mud lakes - Papa and Cruz-Papa
(2020)
Diplophlyctis intestina Fresh-blackish water Characea, cortical cells of Chara
sp. (lake), sediment (reservoir), moribund and dead
internodal cells of Nitella sp. (bog)
Brazil, Germany,
India, USA Engler etal. (1887), Spar-
row (1952), Jerônimo
and Pires-Zottarelli
(2020)
Diplophlyctis laevis Cladophora from Crooked Lake, dead internodal cells
of Nitella (bog) USA Sparrow (1937b, 1952)
Diplophlyctis nephrochytrioides Saprophytic or weakly parasitic in moribund internodes
of New Zealand Karling (1968b)
Nitella sp, lake
Diplophlyctis sarcoptoides Insects’ substrate in aquatic, Nitella and Chara (lake)
water (bog), pond, water column and sediment (res-
ervoir)
Brazil, New Zea-
land, UK, USA Sparrow (1957), Karling
(1967), Letcher etal.
(2018), Papa and Cruz-
Papa (2020)
Diplophlyctis sexualis Submerged decaying plant debris UK, Canada, USA Haskins (1950)
Diplophlyctis verrucosa Chara fragilis cells, moor and ponds Japan Czeczuga etal. (2005)
Diplophlyctis versiformis Bog water USA Papa and Cruz-Papa
(2020)
Endochytrium digitatum Saprophytic in internodes of Chara coronata, Nitella
flexilis, and other algae, mud (lake) UK, USA Karling (1938), Wil-
loughby (1961)
Endochytrium multiguttulatum Bogs USA Dogma (1969b)
Endochytrium pseudodistomum Parasite in Stigeoclonium tenue, lake USA Karling (1941)
Endochytrium oophilum Parasitic on rotifers eggs USA Sparrow (1933a)
Endochytrium ramosum Cladophora sp. USA Sparrow (1933a)
Nowakowskiellaceae
Nowakowskiella crenulata Sediment surface Brazil Jerônimo etal. (2019)
Nowakowskiella delica Mud (lake) UK Willoughby (1961)
Nowakowskiella elegans Grass leaf blade and mud (lake) UK, USA Willoughby (1961), Pater-
son (1967)
Nowakowskiella hemisphaerospora Sediment surface, water (reservoir), mud (lake) Brazil, UK, USA Shanor (1942), Wil-
loughby (1961),
Jerônimo etal. (2019)
Nowakowskiella macrospora Water (river) Poland Czeczuga etal. (1990)
Nowakowskiella multispora Water (stream) Brazil Jerônimo etal. (2019)
Nowakowskiella ramosa Water column (reservoir) Brazil Jerônimo etal. (2019)
Nowakowskiella crassa Debris in water USA Karling (1949c)
Nowakowskiella methistemichroma Onion-skin bait from stagnant water Poland Batko and Hassan (1982)
Nowakowskiella moubasherana Sphagnum bog water Poland Hassan (2014)
Septochytriaceae
Septochytrium macrosporum Water USA Karling (1942a)
Septochytrium plurilobulum Water in decaying leaves of Acer pensylvanicum USA Johanson (1943)
Cladochytriales genera incertae
sedis
Nephrochytrium appendiculatum Pond Poland Czeczuga etal. (2005)
Nephrochytrium aurantium Mud (lake) UK, USA Whiffen (1941), Wil-
loughby (1961)
Nephrochytrium bipes Pond Poland Hassan (1983)
Nephrochytrium stellatum Pond Poland Czeczuga etal. (2005)
LOBULOMYCETES

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Lobulomycetales
Alogomycetaceae
Algomyces stechlinensis Eudorina elegans (lake) Germany Van den Wyngaert etal.
(2018)
Lobulomycetaceae
Clydaea vesicula Creek bank mud USA Simmons etal. (2009)
Lobulomyces angularis Sphagnum from acidic lake, mud pond, pollen bait
(pond) USA Simmons etal. (2009),
Davis etal. (2018)
MESOCHYTRIOMYCETES
Gromochytriales
Gromochytriaceae
Gromochytrium mamkaevae Parasite of Tribonema gayanum, ditch Russia Karpov etal. (2014)
Mesochytriales
Mesochytriaceae
Mesochytrium penetrans Lake, pond Russia Gromov etal. (2000), Kar-
pov etal. (2010)
POLYCHYTRIOMYCETES
Polychytriales
Arkayaceae
Arkaya lepida River USA Longcore and Simmons
(2012)
Polychytriaceae
Karlingiomyces asterocystis Shrimp exoskeleton: water and soil USA Blackwell etal. (2004)
Karlingiomyces curvispinosus River USA Blackwell etal. (2004)
Karlingiomyces dubius River USA Blackwell etal. (2004)
Karlingiomyces marylandicus Water (ditch), mud (lake) UK, USA Willoughby (1961),
Blackwell etal. (2004)
Lacustromyces hiemalis Lake and pond USA Longcore (1993)
Neokarlingia chitinophila Chitinous substrata, water and soil USA Longcore and Simmons
(2012)
Polychytrium aggregatum Decaying plant material in bog: USA Ajello (1945)
RHIZOPHYDIOMYCETES
Rhizophydiales
Alphamycetaceae
Alphamyces chaetifer Stream, pollen bait (pond), Cosmarium contractum var.
ellipsoideum, Staurastrum cingulum, S. planktonicum,
S. lunatum, S. anatinum, S. pseudopelagicum, Xan-
thidium subhastiferum, Asterococcus sp., Spirogyra,
Spondylosium planum
Argentina, USA,
UK Canter (1961), Letcher
etal. (2008b), Davis
etal. (2018)
Betamyces americae-meridionalis Roadside pond onto pollen Argentina Letcher etal. (2012)
Angulomycetaceae
Angulomyces argentinensis Isolated on pollen from stream Argentina Letcher etal. (2008b)
Aquamycetaceae
Aquamyces chlorogonii Semi-permanent roadside pond Argentina Letcher etal. (2008b)
Collimycetaceae
Collimyces mutans Microglena coccifera, pond Japan Seto and Degawa (2018)
Coralloidiomycetaceae
Coralloidiomycesdigitatus Submersed mud in lake: Chubut Argentina Letcher etal. (2008a)
Globomycetaceae
Globomyces pollinis-pini Lake, pine pollen (bog) Argentina, USA Sparrow (1952), Letcher
etal. (2008b)
Urceomyces sphaerocarpum Lake Argentina Letcher etal. (2008b)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Gorgonomycetaceae
Gorgonomyces haynaldii Stream Argentina Letcher etal. (2008b)
Halomycetaceae
Halomyces littoreus River USA Letcher etal. (2015a)
Paranamyces uniporus Pond USA Letcher etal. (2015a)
Ulkenomyces aestuarii River, submersed estuary mud Germany Letcher etal. (2015a)
Kappamycetaceae
Kappamyces laurelensis Water cultures of moss-covered soil from along a stream USA Letcher and Powell
(2005b)
Pateramycetaceae
Pateramyces corrientinensis Small lake Argentina Letcher etal. (2008b)
Protrudomycetaceae
Protrudomyces laterale Small lake Argentina Letcher etal. (2008b)
Rhizophydiaceae
Rhizophydium algavorum Parasitic, Chlorococum minutum, pond Russia Gromov etal. (1999)
Rhizophydium androdioctes Parasitic on Dictyosphaerium pulchellum, Loch Leven Scotland Canter (1971)
Rhizophydium apiculatum Pond water India Dayal and Kiran (1980)
Rhizophydium blastocladianum Parasitic on Blastocladia on apple, tank sediment India Dasgupta and John (1990)
Rhizophydium blyttiomycerum Parasitic on Blyttiomyces spinosus and zygospore of
Spirogyra, ditch India Dasgupta and John (1990)
Rhizophydium clavatum Parasitic on Closterium, tank India Dasgupta and John (1990)
Rhizophydium columnare Parasitic on Spirotaenia condensata (pond) UK Canter (1947b)
Rhizophydium conchiforme Ditch India Dasgupta and John (1990)
Rhizophydium constantineanui Dead cells of Oedogonium sp., and Cosmarium sp
(pool) UK Richards (1956)
Rhizophydium contractophilum Parasitic on Eudorina elegans and Eudorina sp, lake UK Canter (1959)
Rhizophydium coronum Organic debris in water (bog), cellulose bait (pond) USA Davis etal. (2019) Davis
etal. (2018)
Rhizophydium difficile Parasitic on Staurastrum jaculiferum (lake) UK Canter (1954)
Rhizophydium dubium Saprobic on decaying leaves of Eichornia, pond India de Wildeman (1895)
Rhizophydium echinocystoides Saprobic on pine pollen, bog USA Letcher and Powell (2012)
Rhizophydium elyense Sediment (reservoir), mud (lake) Brazil, UK Willoughby (1961),
Jerônimo and Pires-
Zottarelli (2020)
Rhizophydium fugax Parasitic on Cryptomonas sp., plankton of lakes UK Canter and Lund (1968)
Rhizophydium fusus Parasitic on Melosira granulata UK Canter and Lund (1953)
Rhizophydium gibbosum Staurastrum anatinum, S. pseudopelagicum, Staurodes-
mus curvatus, Staurodesmus sp. UK Canter (1961)
Rhizophydium gonapodyanum Parasitic on sporangium of Gonapodya polymorpha,
tank sediment India Dasgupta and John (1990)
Rhizophydium granulosporum Parasitic on Tribonema sp. (ditch) UK Sparrow (1957)
Rhizophydium jobii Benthic detritus Oman Hassett etal. (2020)
Rhizophydium keratinophilum Human hair bait, algae debris (bog) USA Sparrow (1952)
Rhizophydium koreanum Pond South Korea Hyde etal. (2019)
Rhizophydium lagenaria Parasitic on sporangium of Blastocladia on fruit bait,
tank sediment India Dasgupta and John (1990)
Rhizophydium lenelangeae Saprobic on pine pollen bait, riverbank soil Guatemala Sparrow and Lange (1977)
Rhizophydium mammillatum Weak parasite or saprophyte on cells of Cladophora
(ditch) UK Richards (1956)
Rhizophydium manoense Parasitic on diatom (Cocconeis), stream USA Sparrow (1965a)
Rhizophydium mougeotiae Parasitic on Mougeotia gracillima, Lake Switzerland Letcher and Powell (2012)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Rhizophydium nobile Parasitic on the resting spores of Ceratium hirundinella,
lakes UK Canter (1968)
Rhizophydium oblongum Parasitic on Dinobryon spp. (Lake) UK Canter (1954)
Rhizophydium obpyriformis Parasitic on chytrids (pond) USA Davis etal. (2018)
Rhizophydium planktonicum Parasitic on Asterionella formosa, lakes UK Canter and Lund (1948)
Rhizophydium poculiforme Parasitic on Closterium, tank India Dasgupta and John (1990)
Rhizophydium porosum Bog USA Sparrow and Lange (1977)
Rhizophydium punctatum Lake Russia Letcher and Powell (2012)
Rhizophydium reflexum Parasitic on the sporangium of Blastocladia on Pyrus
fruit, tank sediment India Dasgupta and John (1990)
Rhizophydium rhizinum Parasitic on the sporangium of Blastocladia on Pyrus
fruit, tank sediment India Dasgupta and John (1990)
Rhizophydium rotundum Parasitic on Closterium, Oedogonium, and Zygnema,
tank India Dasgupta and John (1990)
Rhizophydium sphaerotheca Pool and bog UK Sparrow (1957)
Rhizophydium sparrowii “Saproparasitic” on Ulothrix sp., ditch India Dasgupta and John (1990)
Rhizophydium spinosum Parasitic on Blastocladia on fruit bait, tank sediment India Dasgupta and John (1990)
Rhizophydium squamosum Lake Russia Letcher and Powell (2012)
Rhizophydium stellatum Parasitic on the sporangium of Gonapodya polymorpha,
tank sediment India Dasgupta and John (1990)
Rhizophydium stipitatum Sediment (reservoir) Brazil Jerônimo and Pires-Zotta-
relli (2020)
Rhizophydium tetragenum Parasitic on Asterionella formosa, Lake Switzerland Letcher and Powell (2012)
Rhizophydium tubulatum Parasitic on Closterium India Dasgupta and John (1990)
Rhizophydium ubiquetum Saprobic on Cosmarium and other desmids, plankton
of lakes UK Canter (1961)
Rhizophydium uniguttulum Parasitic on Gemellicystis neglecta (lake) UK Canter (1954)
Rhizophydium undatum Bog, lake Russia, USA Sparrow and Lange
(1977), Letcher and
Powell (2012)
Rhizophydium urceolatum Parasitic on Blastocladia on fruit bait, tank sediment India Dasgupta and John (1990)
Rhizophydium venustum Parasitic on Uroglena americana, lakes UK Canter (1963b)
Staurastromycetaceae
Staurastromyces oculus Lake Germany Van den Wyngaert etal.
(2017)
Homolaphlyctis polyrhiza Acidic oligotrophic lake USA Longcore etal. (2012)
Terramycetaceae
Boothiomyces angulosus Sediment (reservoir) Brazil Jerônimo and Pires-Zotta-
relli (2020)
Boothiomyces macroporosus Pollen bait (pond) USA Davis etal. (2016)
RHIZOPHLYCTIDOMYCETES
Rhizophlyctidales
Borealophlyctidaceae
Borealophlyctis nickersoniae Pond USA Davis etal. (2016)
Rhizophlyctidaceae
Karlingia polonica Water (pond, river) Poland Hassan (1983), Czeczuga
etal. (1990)
Rhizophlyctis columellae Zygospore of Spirogyra India Das-Gupta and John
(1990)
Rhizophlyctis petersenii Cellophane "bait," in water culture with soil USA Sparrow (1952)
Rhizophlyctis rosea Mud (lake) UK Willoughby (1961)
SPIZELLOMYCETES

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Spizellomycetales
Spizellomycetaceae
Spizellomyces palustris Peat bog ditch Germany Wakefield etal. (2010)
SYNCHYTRIOMYCETES
Synchytriales
Synchytriaceae
Endodesmidium formosum Bog, Parasitic in Netrium oblongum, Cylindrocystis
crassa and Cylindrocystis brebissonii UK Canter (1949c)
Synchytrium laeve Mougeotia sp. (pond) UK Canter (1949c)
Synchytrium microbalum Mud Pond, acidic lake USA Longcore etal. (2016)
Synchytrium petersenii Mougeotia sp. (bog) UK Canter (1949c)
Synchytriales genus incertae sedis
Micromyces intermedius Zygnema (bog) UK Canter (1949c)
Micromyces mirabilis Growing in Closterium cells, Bog USA Davis etal. (2019)
Micromyces oedogonii Oedogonium (lake) UK Roberts (1953)
Micromyces zygogonii Growing in Mougeotia filaments (pond), pasitic in
vegetative filaments of Mougeotia genuflexa (bog),
Spirogyra (pond)
UK, USA Canter (1949c), Spar-
row (1952), Davis etal.
(2019)
Chytridiomycota genera incertae sedis
Achlyella flahaultii River Poland Czeczuga and Muszynska
(2001)
Achlyogeton entophytum River Poland Czeczuga and Muszyńska
(1993)
Achlyogeton rostratus Water France Sorokine (1896)
ENTOMOPHTHOROMYCOTA
Entomophthoromycetes
Entomophthorales
Ancylistaceae
Ancylistes closterii Closterium sp. (bog) USA Davis etal. (2019)
Conidiobolus villosus Saprophytic on cast skin of snake used as "bait" in water
culture of soil and rotten wood (bog) USA Sparrow (1952)
Entomophthoraceae
Erynia aquatica Aedes pupal cadaver (woodland pool) USA Molloy and Wraight
(1982), Sridhar and
Kaveriappa (1992),
Voglmayr (1996)
Erynia rhizospora Foam (stream, rivulet, water) Austria, India,
Poland Sridhar and Kaveriappa
(1992), Voglmayr
(1996), Nadeau etal.
(1996), Davis etal.
(2019)
Erynia conica Foam (stream, rivulet), Simulium verecundum complex
(lake) Austria, Canada,
India Czeczuga (1996)
MONOBLEPHAROMYCOTA
HYALORAPHIDIOMYCETES
Hyaloraphidiales
Hyaloraphidiaceae
Hyaloraphidium curvatum Freshwater South Africa Ustinova etal. (2000)
MONOBLEPHARIDOMYCETES
Monoblepharidales
Gonapodyaceae
Gonapodya bohemica Forest pools Czech Republic Cejp (1947)

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Gonapodya polymorpha Submerged twigs USA Thaxter (1895b)
Gonapodya prolifera Decaying apples in water, tomato bait (river) USA, UK Thaxter (1895b), Water-
house (1942)
Gonapodya siliquiformis Decaying apples in water USA Thaxter (1895b)
Harpochytriaceae
Harpochytrium adpressum Parasitic on freshwater algae Hungary Schumacher and Whitford
(1961)
Harpochytrium apiculatum Freshwater algae USA Schumacher and Whitford
(1961)
Harpochytrium botryococci Freshwater UK Jane (1946)
Harpochytrium hedenii Freshwater algae USA, France,
Germany, Rus-
sia, UK
Jane (1946)
Harpochytrium hyalothecae Freshwater USA, Germany,
UK Jane (1946)
Harpochytrium intermedium Freshwater Jane (1946)
Harpochytrium monae Zygogonium ericetorum in a temporary pool UK Jane (1946)
Harpochytrium natrophilum Trisetum cernuum, freshwater USA Jane (1946)
Harpochytrium ornithocephalum Freshwater Switzerland Jane (1946)
Harpochytrium tenuissimum Tubs in the ground, freshwater UK Jane (1942)
Harpochytrium vermiforme In cells of freshwater algae Germany Jane (1946)
Harpochytrium viride Parasitic on freshwater algae Hungary Scherffel (1926)
Monoblepharidaceae
Monoblepharis bullata Submerged twigs of Quercus robur and of Fraxinus
excelsior UK Perrott (1955)
Monoblepharis fasciculata Submerged sticks USA Thaxter (1895a)
Monoblepharis hypogyna Submerged twigs of Quercus robur, water (river) Poland, UK Perrott (1955), Czeczuga
etal. (1990)
Monoblepharis insignis Submerged sticks, pool and ditch USA Thaxter (1895a)
Monoblepharis laevis Fruit of Rosa, immersed in water USA Sparrow (1960)
Monoblepharis macrandra Submerged plants (ponds, stream) Sweden, USA Lagerheim (1900), Spar-
row (1952)
Monoblepharis polymorpha Saprobic on Quercus robur L. and Fraxinus excelsior L France? Perrott (1957)
Monoblepharis macrandra Submerged twigs, swamps USA Sparrow (1965b)
Monoblepharis ovigera Submerged fallen branches of Abies picea Sweden Lagerheim (1900)
Monoblepharis sphaerica Submerged twigs of Quercus robur France Cornu (1872)
Monoblepharis regignens Submerged fallen branches of Pinus sylvestris Sweden Lagerheim (1900)
Oedogoniomycetaceae
Oedogoniomyces lymnaeae shells of living Limnaea ollula and Linaea reticulata Japan Kobayasi and Ôkubo
(1954)
Telasphaerulaceae
Telasphaerula gracilis Lake water, associated with Chara USA Karpov etal. (2017a)
SANCHYTRIOMYCOTA
SANCHYTRIOMYCETES
Sanchytriales
Sanchytriaceae
Amoeboradix gromovi Parasitic on Tribonema gayanum, Tribonema vulgare
and Ulothrix tenerrima, water of a sand-pit Russia Karpov etal. (2018)
Sanchytrium tribonematis Pond Russia Karpov etal. (2017a)
MORTIERELLOMYCOTA
MORTIERELLOMYCETES
Mortierellales

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Mortierellaceae
Aquamortierella elegans Midge larvae in a freshwater stream New Zealand,
Japan Vandepol etal. (2020)
Gryganskiella fimbricystis Moss cushion in a very wet bog Argentina Vandepol etal. (2020)
Mortierella elongata Water (river) South Korea Nguyen etal. (2019b)
Mortierella fluviae Water (river) South Korea Hyde etal. (2016b)
Mortierella minutissima Water (river) South Korea Nguyen and Lee (2016)
MUCOROMYCOTA
ENDOGONOMYCETES
Endogonales
Endogonaceae
Endogone pisiformis Sphagnum (bog) USA Sparrow (1952)
MUCOROMYCETES
Mucorales
Choanephoraceae
Gilbertella persicaria Water (river) South Korea Lee etal. (2018)
Cunninghamellaceae
Absidia glauca Mud (river canal) Egypt Moubasher etal. (2018)
Cunninghamella bertholletiae Water (river) South Korea Nguyen etal. (2017a)
Gongronella namwonensis Freshwater South Korea Crous etal. (2020)
Lichtheimiaceae
Rhizomucor pusillus Water (tank) USA Schell etal. (2011)
Mucoraceae
Actinomucor elegans Water (river) South Korea Nguyen etal. (2017b)
Mucor abundans Water (stream) South Korea Nguyen etal. (2020a)
Mucor aligarensis Water (stream) South Korea Nguyen etal. (2020a)
Mucor amphibiorum Pathogenic to Ornithorhynchus anatinus (creek, river) Australia Connolly etal. (1998)
Mucor fluvius Freshwater South Korea Wanasinghe etal. (2018)
Mucor heterogamus Freshwater, sediment South Korea Nguyen etal. (2020a)
Mucor hiemalis Pathogens of Cyprinis carpio communis and Oncorhyn-
chus mykiss (cultured farm) India Magray etal. (2020)
Mucor moelleri Water, Sediment (stream) South Korea Nguyen etal. (2020a)
Mucor ramosissimus Freshwater South Korea Nguyen and Lee (2018)
Mucor racemosus Polluted water (canal) Egypt El-Morsy etal. (2013)
Rhizopodaceae
Rhizopus oryzae Polluted water (canal) Egypt El-Morsy etal. (2013)
Rhizopus stolonifer Tap water Portugal Gonçalves etal. (2006)
Syncephalastraceae
Syncephalastrum racemosum Polluted water (canal) Egypt El-Morsy etal. (2013)
OLPIDIOMYCOTA
OLPIDIOMYCETES
Olpidiales
Olpidiaceae
Olpidium gregarium Freshwater (river), rotifer eggs (pools) Poland, UK Sparrow (1957), Czeczuga
etal. (1990)
Olpidium hyalothecae Water UK Canter (1949b)
Olpidium pendulum Pine pollen (lake) USA Sparrow (1952)
Olpidium rotiferum Freshwater (river) Poland Czeczuga etal. (1990)
ZOOPAGOMYCOTA
ZOOPAGOMYCETES

Fungal Diversity
1 3
Table 15 (continued)
Taxon Substrate/Host Country References
Zoopagales
Zoopagaceae
Acaulopage dichotoma Foam (stream) Austria Voglmayr (1996)
Zoophagus cornus Parasitic on rotifers (lake), mud (paddy field) India, Japan Karling (1966), Glockling
(1997)
Zoophagus insidians Cadophora (freshwater), rotifers in water (fen, bog,
ditch) Austria, UK, USA Sommerstorff (1911),
Sparrow (1957), Davis
etal. (2019)

Fungal Diversity
1 3
Table 16 Classification of Microsporidia and Rozellomycota genera incertae sedis recorded from freshwater environment including information
on habitats, substrates/hosts, and geographical distribution
Taxon Host Distribution References
ROZELLOMYCOTA
Microsporidea
Amblyosporida
Amblyosporidae
Amblyospora bakcharia Ochlerotatus excrucians larva (temporary
vernal pool) Russia Andreadis etal. (2012)
Amblyospora baritia Ochlerotatus excrucians larva (temporary
vernal pool) Russia Andreadis etal. (2012)
Amblyospora bogashova Ochlerotatus excrucians larva (temporary
vernal pool) Russia Andreadis etal. (2012)
Amblyospora bracteata Simulium damnosum larva (river), S. venustum,
S. verecundum, and S. vittatum (pond, river) Canada, Nigeria Vávra and Undeen (1981), Weiser and
Prasertphon (1982)
Amblyospora burlaki Culex pipiens larva (floodplain water body) Russia Pankova etal. (2000)
Amblyospora camposi Culex renatoi, a bromeliad-inhabiting mosquito Argentina Micieli etal. (2000)
Amblyospora canadensis Aedes canadensis, A. cinereus larva (bog
swamp) USA Andreadis (1993)
Amblyospora caspius Aedes caspius, A. communis larva (reservoir
from melted snow) Russia Pankova etal. (2000)
Amblyospora chulymia Ochlerotatus caspius larva (temporary vernal
pool) Russia Andreadis etal. (2012)
Amblyospora dyxenoides Culex annulirostris adult and larvae and Meso-
cyclops albicans Australia Sweeney etal. (1988)
Amblyospora flavescens Ochlerotatus dianaetus, Aedes flavescens larva
(sphagnum bog) Russia Simakova and Pankova (2005),
Andreadis etal. (2012)
Amblyospora hristinia Ochlerotatus communis larva (temporary vernal
pool) Russia Andreadis etal. (2012)
Amblyospora jurginia Ochlerotatus excrucians larva (temporary
vernal pool) Russia Andreadis etal. (2012)
Amblyospora kazankia Ochlerotatus dianaetus larva (temporary vernal
pool) Russia Andreadis etal. (2012)
Amblyospora khaliulini Aedes communis and Acanthocyclops vernalis
(vernal pool) USA Vossbrinck etal. (2004), Andreadis etal.
(2018)
Amblyospora kolarovi Ochlerotatus punctor, Aedes communis (sphag-
num bog) Russia (micro-
sporidium), USA
(host)
Simakova and Pankova (2005),
Andreadis etal. (2012)
Amblyospora mavlukevia Aedes cinereus larva (riverside pond) Russia Andreadis etal. (2012)
Amblyospora mocrushinia Ochlerotatus punctor larva (temporary vernal
pool) Russia Andreadis etal. (2012)
Amblyospora modestium Aedes cinereus larva (pond) Russia Andreadis etal. (2012)
Amblyospora orbiculata Aedes diantaeus larva (sphagnum bog) Russia Simakova and Pankova (2005)
Amblyospora rugosa Ochlerotatus cataphylla, O. cantans, Aedes
euedes (temporary reservoirs after snow
melting)
Russia Simakova and Pankova (2005), Simakova
etal. (2011), Andreadis etal. (2012)
Amblyospora salairia Culex modestus larva (temporary vernal pool) Russia Andreadis etal. (2012)
Amblyospora severinia Ochlerotatus excrucians larva (temporary
vernal pool) Russia Andreadis etal. (2012)
Amblyospora shegaria Aedes cinereus larva (temporary vernal pool) Russia Andreadis etal. (2012)
Amblyospora timirasia Aedes cinereus larva (temporary vernal pool) Russia Andreadis etal. (2012)
Amblyospora undata Culex pipiens larva (continental reservoir) Russia Simakova and Pankova (2005)
Amblyospora urski Aedes communis larva (temporary reservoirs
after snow melting) Russia Simakova and Pankova (2005)
Amblyospora weiseri Aedes cantans larvae (pool) Bohemia Lukeš and Vávra (1990)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Andreanna caspii Ochlerotatus caspius instar (temporary vernal
pool) Russia Simakova etal. (2008)
Becnelia sigarae Testes of the water boatmen, Sigara lateralis Czech Republic Tonka and Weiser (2000)
Crepidulospora beklemishevi Anopheles beklemishevi larva (water body river
floodplain) Russia Simakova etal. (2003, 2004)
Culicospora damnosi Simulium damnosum larva (river) Nigeria Weiser and Prasertphon (1982)
Culicospora magna Culex restuans larvae (river) USA Becnel etal. (1987)
Culicosporella lunata Culex pilosus larvae (ditch) USA Hazard etal. (1984)
Dimeiospora palustris Aedes punctor larvae (sphagnum swamp) Russia Simakova etal. (2003)
Hyalinocysta chapmani Culiseta melanura larva, Orthocyclops modes-
tus (sphagnum bog) USA Andreadis and Vossbrinck (2002)
Hyalinocysta expilatoria Bogfly Odagmia ornata larva (stream) Sweden Larsson (1983)
Intrapredatorus barri mosquito Culex fuscanus larva (artificial tank) China Chen etal. (1998)
Intrapredatorus trinus Culex halifaxi larvae(stream) Australia Becnel andSweeney (1990)
Novothelohania ovalae Ochlerotatus caspius larva (temporary vernal
pool) Russia Andreadis etal. (2012)
Parathelohania anopheles Anopheles quadrimaculatus larva, Microcy-
clops varicans (wading pool) USA Avery and Undeen (1990)
Parathelohania divulgata Anopheles messeae, A. beklemishevi larva
(reservoir of river floodplain) Kazakhstan, Russia Simakova and Pankova (2004),
Andreadis etal. (2012)
Parathelohania formosa Anopheles messeae larva (reservoir of river
floodplain) Russia Simakova and Pankova (2004)
Parathelohania iranica Anopheles superpictus larva (leakage pool) Iran Omrani etal. (2017)
Parathelohania sibirika Anopheles messeae larva (reservoir of river
floodplain) Russia Simakova and Pankova (2004)
Parathelohania teguldeti Anopheles beklemishevi larva (water body of
river floodplain) Russia Simakova and Pankova (2004)
Parathelohania tomski Anopheles messeae (reservoir of river flood-
plain) Russia Simakova and Pankova (2004),
Andreadis etal. (2012)
Trichoctosporea pygopellita Acanthocyclops reductus, Ochlerotatus cyprius,
C. excrucians Russia Simakova etal. (2011), Andreadis etal.
(2012)
Tricornia muhezae Mosquito Mansonia africana larva (pond) Tanzania Pell and Canning (1992)
Caudosporidae
Binucleospora elongata Ostracod Candona (temporary water bodies on
a meadow_ Sweden Bronnvall and Larsson (1995)
Caudospora simulii Blackfly Prosimulium mixtum larva (stream) Canada Vávra and Undeen (1981)
Flabelliforma ostracodae Ostracods (Candona sp.), collected on flooded
meadows Sweden Bronnvall and Larsson (2001)
Neoflabelliforma aurantiae Sludge work Tubifex tubifex (pond) UK Morris and Freeman (2010)
Scipionospora tetraspora Midge (Tanytarsus, Endochironomus) larva
(pond, river) France, Sweden Bylén and Larsson (1996)
Gurleyidae
Agglomerata cladocera Daphnia magna (lake) Russia Sokolova etal. (2016)
Agglomerata daphniae Daphnia magna China Weng etal. (2020)
Agglomerata volgensae Daphnia magna (pond) Russia Larsson and Voronin (2000)
Binucleata daphniae Daphnia magna (pond) Belgium Refardt etal. (2008)
Conglomerata obtusa Daphnia species Czech Republic,
Poland Vávra etal. (2018)
Episeptum anaboliae Anabolia furcate larva (pond) Czech Republic Hyliš etal. (2007)
Episeptum circumscriptum caddisfly Hydropsyche siltalai, H. incognita
(river) Czech Republic,
Sweden Larsson (1996), Hyliš etal. (2007)
Episeptum invadens Limnephilus fuscicornis larva (small river) Sweden Larsson (1996)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Episeptum inversum Caddis fly Holocentropus picicornis larvae
(pond) Sweden Hyliš etal. (2007)
Episeptum pseudoinversum Sericostoma personatum larva (stream) Czech Republic Hyliš etal. (2007)
Episeptum trichoinvadens Potamophylax cingulatus larva (stream) Czech Republic Hyliš etal. (2007)
Gurleya daphniae Daphnia pulex in ponds Austria Friedrich etal. (1996)
Gurleya vavrai Daphnia pulex and D. longispina (sewage
lagoon) UK Green (1974)
Lanatospora costata Copepod Megacyclops viridis (woodland pool) Czech Republic Vávra etal. (2016b)
Lanatospora macrocyclopis Copepod Macrocyclops albidus (lake) Russia Voronin (1989)
Larssonia obtusa Daphnia pulex in pond Lithuania Vidtmann (1993)
Marssoniella elegans Copepod Cyclops vicinus Czech Republic Vávra etal. (2005)
Norlevinea daphniae Daphnia longispina in carp pond China, Czechoslo-
vakia Vávra (1984)
Paraepiseptum hydropsy-
cheae Hydropsyche siltalai larva (river) UK Xie and Canning (1986)
Paraepiseptum plectrocne-
miae Caddis flies Plectrocnemia conspersa larva
(stream) Czech Republic Hyliš etal. (2007)
Paraepiseptum polycentropi Caddis flies Polycentropus flavomaculatus,
Hydropsyche fulvipes (stream) Czech Republic Hyliš etal. (2007)
Pseudoberwaldia daphniae Daphnia longispina complex (drinking water
reservoir) Central Europe Vávra etal. (2019)
Senoma globulifera Mosquito Anopheles messeae (floodplain pond) Russia Simakova etal. (2005)
Zelenkaia trichopterae Halesus digitatus (stream) and Micropterna
sequax (pond), aquatic caddis flies Bulgaria Hyliš etal. (2013)
Amblyosporida genera incertae sedis
Alfvenia sibirica Cyclopid Cyclops sp. (lake) Russia Sokolova etal. (2016)
Takaokaspora nipponicus Mosqution Ochlerotatus japonicus japonicus,
O. hatorii (riverside rock pools) Japan Andreadis etal. (2013)
Trichotuzetia guttata Cyclops vicinus (pond) Czech Republic Vávra etal. (1997)
GLUGEIDA
Glugeidae
Alloglugea bufonis Bufo marinus tadpoles and metamorphosing
toads (lake) Brazil Paperna and Lainson (1995)
Amazonspora hassar Teleost fish Hassar orestis Brazil Azevedo and Matos (2003)
Cambaraspora floridanus Floridian crayfish, Procambarus paeninsulanus USA Bojko etal. (2020b)
Glugea anomala Fish Gasterosteus aculeatus, G. pungitius and
Gobius minutus (ditches, stream) Belgium, France,
Germany, Russia,
UK
Ward etal. (2005a), Minter (2019)
Glugea hertwigi Rainbow Smelt Osmerus mordax (lake) Canada Fantham etal. (1941), Dechtiar (1965),
Putz etal. (1965), Dechtiar (1972),
Nepszy etal. (1978)
Glugea mesocotti Fish Mesocottus haitej (river) Russia Voronin and Iukhimenko (2010)
Glugea plecoglossi Ayu Plecoglossus altivelis altivelis China Zhou etal. (2018)
Glugea stephani Fish France, Russia, UK,
USA Bychowskaya-Pavlovskaya etal. (1962)
Loma myrophi Fish Myrophis platyrhynchus (river) Brazil Azevedo and Matos (2002)
Loma psittaca Fish Colomesus psittacus (river) Brazil Casal etal. (2009)
Loma salmonae Fish Oncorhynchus sp. (rearing tanks) Canada Becker and Speare (2007)
Pseudoloma neurophilia Zebrafish Danio rerio (laboratroy colony) USA Matthews etal. (2001), Ramsay etal.
(2010)
Spragueidae
Apotaspora heleios Grass shrimp Palaemonetes paludosus (wet-
land) USA Sokolova and Overstreet (2018)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Microgemma hepaticus Grey mullet, Chelon labrosus – Ralphs and Matthews (1986)
Potaspora aequidens Fish Aequidens plagiozonatus (river) Brazil Videira etal. (2015)
Potaspora macrobrachium Prawn Macrobrachium nipponense (pond) China Ding etal. (2016)
Potaspora morhaphis Teleostean fish, Potamorhaphis guianensis
(river) Brazil Casal etal. (2008)
Pseudokabatana alburnus Topmouth culter (Culter alburnus) (lake) China Liu etal. (2019)
Thelohaniidae
Cucumispora dikerogam-
mari Amphipod Dikerogammarus villosus (reservoir,
river) France, Poland,
Ukraine Ovcharenko etal. (2010)
Cucumispora ornata Amphipod Dikerogammarus haemobaphes
(river) UK Bojko etal. (2015)
Cucumispora roeselii Amphipod Gammarus roeseli (stream) Poland Bojko etal. (2017)
Napamichum aequifilum Aquatic mite Limnochares aquatica Sweden Larsson (1990b)
Napamichum cellatum Midge Endochironomus sp. larva (pond) Sweden Bylén and Larsson (1994)
Nudispora biformis Dragonfly Coenagrion hastulaturn (bog) Sweden Larsson (1990a)
Thelohania contejeani Crayfish Paranephrops planifrons, white-
clawed crayfish Austropotamobius pallipes New Zealand, Italy Henneguy and Thélohan (1892), Pretto
etal. (2018)
Thelohania fibrata Simulium damnosum larva (river), Simulium
venustum/verecundum complex larva (cove) Canada, Nigeria Vávra and Undeen (1981), Weiser and
Prasertphon (1982)
Thelohania montirivulorum Crayfish, Cherax destructor Australia Jones and Lawrence (2001), Moodie
etal. (2003a)
Thelohania varians Blackfly Simulium tuberosum (pond) and S.
venustum (river) larva Canada Vávra and Undeen (1981)
Unikaryonidae
Dictyocoela diporeiae Amphipod Diporeia sp. (lake) USA Winters and Faisal (2014)
Dictyocoela muelleri Amphipod Gammarus duebeni celticus, G.
roeseli (lake) Ireland Terry etal. (2004)
Unikaryon moniezi Eucyclops macrurus (lake) Russia Voronin (1991, 1999)
Unikaryon piriformis Trematode larvae from aquatic snails Malaysia Canning etal. (1974)
Unikaryon pfeifferi Acanthocyclops viridis (lake) Russia Voronin (1977, 1999)
Unikaryon vrevi Macrocyclops distinctus (lake) Russia Voronin (1977, 1999)
Glugeida genus incertae sedis
Triwangia caridinae Shrimp, Caridina formosae China Wang etal. (2013)
NEOPEREZIIDA
Berwaldiidae
Berwaldia schaefernai Daphnia sp., drinking water Czech Republic Vávra and Larsson (1994)
Berwaldia daphniae Daphnia magna (inland pool) Siberia Simakova etal. (2018)
Berwaldia singularis Daphnia pulex (pool) Sweden Larsson (1981)
Fibrillanosema crangonycis Amphipod Crangonyx pseudogracilis (pond) UK Slothouber Galbreath etal. (2004)
Neopereziidae
Bacillidium strictum Oligochaete Limnodrilus hoffmeisteri (stream) Sweden Larsson (1992b)
Bacillidium vesiculoformis Oligochaete Nais simplex (aquarium) Scotland Morris etal. (2005b)
Bryonosema plumatellae Plumatella nitens, freshwater bryozoan USA Canning etal. (2002)
Bryonosema tuftyi Plumatella sp., freshwater bryozoans (lake) UK Canning etal. (2002)
Neoperezia chironomi Chironomus plumosus (lake) Russia Issi etal. (2012)
Neoperezia semenovaiae Chironomus plumosus (lake) Russia Issi (1986), Issi etal. (2012)
Pseudonosema cristatellae Cristatella mucedo, a freshwater bryozoan
(lake) UK Canning etal. (2002)
Schroedera airthreyi Bryozoan Plumatella sp. UK Morris etal. (2005a)
Schroedera plumatellae Bryozoan Plumatella fungosa Scotland Morris and Adams (2002)
Trichonosema algonquin-
ensis Bryozoan Pectinatella magnific (lake) Canada Desser etal. (2004)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Trichonosema pectinatellae Freshwater bryozoan, Pectinatella magnifica
(lake) USA Canning etal. (2002)
Neopereziida genera incertae sedis
Janacekia tainanus Kiefferulus tainanus China Weng etal. (2021)
Neopereziida genera incertae sedis
Janacekia adipophila Midge Ptychoptera paludosa (river) Sweden Larsson (1992a)
Janacekia tainanus Midge Kiefferulus tainanus (pond) China Weng etal. (2021)
Systenostrema alba Dragonfly Aeshna grandis larva (pond), A.
viridis (reservoir) Russia, Sweden Larsson (1988), Sokolova etal. (2006)
Systenostrema candida Dragonfly Libellula quadrimaculata larva
(pond) Sweden Larsson (1988)
NOSEMATIDA
Enterocytozoonidae
Paranucleospora theridion Salmon Louse Lepeophtheirus salmonis, Atlan-
tic Salmon (Salmo salar) (farm) Norway Nylund etal. (2010)
Mrazekiidae
Helmichia aggregata Midge Endochironomus sp. larva (lake) Russia Voronin (1998)
Helmichia anomala Midge larva Microtendipes pedellus (polluted
pond) Poland Ovcharenko and Wita (2001)
Helmichia glandulicola Midge Chironomus anthracinus larva (lake) Norway Wülker and Weiser (1991)
Helmichia lacustris Midge Chironomus plumosus larva (lake) Russia Voronin (1998), Tokarev etal. (2012)
Jirovecia sinensis Oligochaete, Branchiura sowerbyi China Liu etal. (2020b)
Mrazekia cyclopis Copepod Acantho cyclop s americanus (pond) Czechoslovakia Vávra (1962), Larsson etal. (1993)
Nosematidae
Nosema eurytremae Echinoparyphium dunni, Echinostoma hystrico-
sum and Echinostoma malayanum, trematode
larvae of ‐water snails,
Malaysia Canning (1972)
Nosema granulosis Amphipod Gammarus duebeni celticus, G.
roeseli (lake) Ireland Terry etal. (2004)
Nosema pimephales Fish Pimephales promelas (lake) Canada Fantham etal. (1941)
Nosema vasicola Echinoparyphium dunni and Echinostoma
audyi, trematode larvae snails Malaysia Canning etal. (1974)
Vairimorpha austropota-
mobii Austropotamobius pallipes complex, subspecies
Austropotamobius italicus carsicus (stream) Italy Pretto etal. (2018)
Vairimorpha cheracis Freshwater crayfish, Cherax destructor Australia Moodie etal. (2003b)
Vairimorpha austropota-
mobii White-clawed crayfish, Austropotamobius
pallipes Italy Pretto etal. (2018)
Ordosporidae
Ordospora colligata Daphnia magna (pond) Germany, UK Larsson etal. (1997)
Nosematida genera incertae sedis
Anisofilariata chironomid Midge Chironomus plumosus larva (lake) Russia Tokarev etal. (2010b)
Crispospora chironomid Midge Chironomus plumosus larva (lake) Russia Tokarev etal. (2010a)
Enterocytospora artemiae Shrimp Palaemonetes sinensis (paddy field) China Jiang etal. (2020)
Glugoides intestinalis Daphnia magna and Daphnia pulex Czech Republic,
UK Larsson etal. (1996)
MICROSPORIDEA families incertae sedis
Cougourdellidae
Cougourdella coryniformis Copepod Macrocyclops albidus (lake) Russia Voronin (1986, 1993a)
Cougourdella polycentropi Hydropsyche saxonica larva (stream) Sweden Larsson (1989)
Duboscqiidae
Duboscqia sidae Holopedium gibberum (lake) Ontario, Canada Larsson and Yan (1988)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Tardivesicula duplicata Caddis fly Limnephilus centralis (temporary
pool) Sweden Larsson and Bylén (1992)
Trichoduboscqia epeori Rhithrogena iridina nymph (stream bed) Germany Batson (1982)
Fibrillasporidae
Fibrillaspora daphniae Daphnia magna (inland pool) Russia Simakova etal. (2018)
Golbergiidae
Krishtalia pipiens Mosquito Culex pipiens pipiens, Cx. pipiens
form molestus, Cx. theileri larva (artificial
container with heavily polluted water)
Ukraine Kilochitskii (1997)
Microfilidae
Microfilum brevirostris Teleostean fish Brachyhypopomus brevirostris Brazil Matos and Azevedo (2004)
Microfilum lutjani Golden African snapper Lutjanus fulgens
(river) Senegal Faye etal. (1991)
Neonosemoidiidae
Neonosemoides tilapie Cichlid fish Sarotherodon melanotheron, Tila-
pia guineensis and T. zillii Benin, Senegal Sakiti and Bouix (1987), Faye etal.
(1991)
Pleistophoridae
Heterosporis sutherlandae Fish Perca flavescens, Sander vitreus) (lake) USA Phelps etal. (2015), Tomamichel etal.
(2018)
Ovipleistophora diplos-
tomuri Fish and trematodes, crayfish Procambarus biv-
ittatus, sunfish Lepomis macrochirus (lake) USA Lovy and Friend (2017), Bojko etal.
(2020a)
Ovipleistophora ovariae Golden Shiner Notemigonus crysoleucas, Ruffe
Gymnocephalus cernuus (lake) Finland Pekkarinen etal. (2002), Phelps and
Goodwin (2008)
Pleistophora beebei Fish Brachyhypopomus beebei (river) Brazil Casal etal. (2016)
Pleistophora gigantea Fish Crenilabrus melops – Sprague and Vernick (1968)
Pleistophora hyphessobry-
conis Neon tetra disease of aquarium fishes (Characi-
dae, Cyprinidae, Cyprinodontidae, Poecili-
dae, Cichlidae), including Danio rerio and D.
nigrofasciatus
Cosmopolitan Shaw and Kent (1999)
Pleistophora mirandellae Cyprinid fish (river) France Maurand etal. (1988)
Pleistophora muelleri Amphipod Gammarus duebeni celticus Ireland Leger and Hesse (1917), Terry etal.
(2004), Ironside etal. (2008)
Pleistophora multispora
Vavraia multispora Simulium damnosum larva (river) Nigeria Weiser and Prasertphon (1982)
Toxoglugeidae
Toxospora volgae Midge Corynoneura (reservoir) Russia Voronin (1993b)
Tuzetiidae
Pankovaia semitubulata Mayfly Cloeon dipterum nymph (floodplain
pond) Russia Simakova etal. (2009a)
Paratuzetia kupermani Procercoid of Khawia armeniaca found in oli-
gochaetes Potamothrix hammonensis (lake) Armenia Poddubnaya etal. (2006)
Tuzetia boeckella Copepod Boeckella triarticulata (artificial lake) Australia Milner and Mayer (1982)
Tuzetia ceriodaphniae Ceriodaphnia affinis, C. reticulata (lake) Russia Voronin (1986)
Tuzetia diaphanosomae Diaphanosoma brachyurum (lake) Russia Voronin (1986)
Tuzetia dualis Mayfly Cloeon dipterum nymph (floodplain
pond) Russia Simakova etal. (2009b)
Tuzetia tschernischovae Daphnia cucullata larva (lake) Russia Voronin (1986)
MICROSPORIDEA genera incertae sedis
Baculea daphniae Daphnia pulex France Loubes and Akbarieh (1978)
Caullerya mesnili Daphnia galeata × hyalina Switzerland Wolinska etal. (2004)
Evlachovaia chironomi Midge Chironomus plumosus larvae (lake) Russia Issi (1986)
Globulispora mitoportans Daphnia spp. Czech Republic Vávra etal. (2016a)
Gurleyides biformis Ceriodaphnia reticulata, C. affinis (lake) Russia Voronin (1986)

Fungal Diversity
1 3
Table 16 (continued)
Taxon Host Distribution References
Hamiltosporidium magnivora Daphnia spp. (ponds) Belgium, Russia,
UK Haag etal. (2011)
Hamiltosporidium tvaermin-
nensis Parasite of Daphnia magna (pools) Finland, Sweden,
Skerry Islands of
the Baltic Sea
Haag etal. (2011)
Holobispora thermocyclopis Thermocyclops oithonoides, T. leuckarti (lake) Russia Voronin (1986)
Kabataia rondoni Teleost Gymnorhamphichthys rondoni, Brazil Casal etal. (2010)
Kabatana takedai Freshwater fish, Pangasius sutchi Asia Lom etal. (2001)
Microsporidium brevirostris Teleostean Brachyhypopomus brevirostris Amazon, Brazil Matos and Azevedo (2004)
Microsporidium chironomid Midge Chironomus dorsalis larva (water basin) Russia Voronin (1975, 1999)
Microsporidium epithelialis Tubificid worms Tubifex sp. (commercial
aquarium) Germany Oumouna etal. (2000)
Microsporidium fluviatilis Copepod Eucyclops serrulatus (river) Russia Voronin (1994)
Microsporidium jiroveci Cyclops vicinus (lake) Russia (Voronin 1977, 1986)
Myosporidium ladogensis Burbot Lota lota from Lake Haukivesi Finland Jones etal. (2020)
Microsporidium lamprogle-
nae Freshwater copepod Lamproglena pulchella
and cyprinid fish, Leuciscus leuciscus Europe Jones etal. (2020)
Microsporidium luciopercae Freshwater pike-perch Sander lucioperca Finland Jones etal. (2017)
Microsporidium spraguei Freshwater pike-perch Sander lucioperca Finland Jones etal. (2017)
Stempellia diaptomi Copepod Diaptomus gracilis (lake) Russia Voronin (1996)
Rozellomycota genera incertae sedis
Mitosporidium daphniae Parasite of the water flea Daphnia magna UK Haag etal. (2014)
Paramicrosporidium sac-
camoebae Parasite of Saccamoeba sp, freshwater pond Germany Corsaro etal. (2014)
Rozella achlyae Achlya flagellata USA Karling (1942b)
Rozella allomycetis Allomyces arbusculus Germany, USA Karling (1942b)
Rozella apodyae Apodachlya brachynema France Karling (1942b)
Rozella blastocladiae Blastocladia pringsheimii, and B. gracilis USA, Denmark,
Germany Karling (1942b)
Rozella canterae Chytridium oedognii UK Karling (1942b)
Rozella chytriomycetis Chytriomyces UK Karling (1946)
Rozella cladochytrii Nowakowskiella profusum, N. elegans, N.
ramosum, Cladochytrium replicatum, C. cras-
sum, and C. hyalinum
USA Karling (1942b)
Rozella coleochaetes Parasitic on Coleochaete sp., bog Karling (1942b)
Rozella diplophlyctidis Diplophlyctis intestina Karling (1942b)
Rozella endochytrii Sporangia of Endochytrium operculatum,
swamp Brazil, USA Karling (1941, 1944d)
Rozella monoblepharidis Monoblepharis polymorpha, and M. macrandra Germany, France Karling (1942b)
Rozella multimorpha Parasitic on Pythium sp., pond USA Karling (1942b)
Rozella parva Parasitic on Zygorhizidium affluens, lakes UK Canter (1969)
Rozella polyphagi Polyphagus laevis UK Karling (1942b)
Rozella rhipidii Araiospora France Karling (1942b)
Rozella rhizoclosmatii Parasitic on Rhizoclosmatium, pond USA Karling (1942b)
Rozella rhizophydii Rhizophydium globosum Brazil Karling (1944c)
Rozella septigena Achyla, Saprolegnia France Karling (1942b)

Fungal Diversity
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Acknowledgements Kevin D. Hyde acknowledges Thailand Research
Fund “The future of specialist fungi in a changing climate: baseline
data for generalist and specialist fungi associated with ants, Rhodo-
dendron species and Dracaena species” (DBG6080013) and “Impact
of climate change on fungal diversity and biogeography in the Greater
Mekong Sub-region” (RDG6130001). Kevin D. Hyde thanks Chiang
Mai University for the Award of Visiting Professor. Mark S. Calabon
is grateful to Mushroom Research Foundation, Department of Sci-
ence and Technology-Science Education Institute (Philippines), and
Thailand Science Research and Innovation (TSRI) grant “Macrofungi
diversity research from the Lancang-Mekong Watershed and surround-
ing areas” (Grant No. DBG6280009). E. B. Gareth Jones acknowledges
the award of a Distinguished Scientist Fellowship (DSFP), King Saud
University, Kingdom of Saudi Arabia. Nattawut Boonyuen thanks BIO-
TEC and the Integrated Technology Platform, NSTDA-Thailand (Grant
Number P1952580) for partial support in this work.
Declarations
Conflict of interest The authors declare that they have no conlict of
interest.
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Hu D, Zhu H, Cai L, Hyde KD, Zhang K (2007) Sirothecium triseriale,
a new chirosporous anamorphic species from China. Cryptogam
Mycol 28:311–314
Hu DM, Cai L, Chen H, Bahkali AH, Hyde KD (2010a) Four new
freshwater fungi associated with submerged wood from South-
west Asia. Sydowia 62:191–203
Hu DM, Cai L, Chen H, Bahkali AH, Hyde KD (2010b) Fungal diver-
sity on submerged wood in a tropical stream and an artificial
lake. Biodivers Conserv 19:3799–3808. https:// doi. org/ 10. 1007/
s10531- 010- 9927-5
Hu DM, Cai L, Bahkali AH, Hyde KD (2012a) Two new freshwater
species of Annulatascaceae from China. Mycotaxon 120:81–88.
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Hu DM, Cai L, Hyde KD (2012b) Three new ascomycetes from fresh-
water in China. Mycologia 104:1478–1489. https:// doi. org/ 10.
3852/ 11- 430
Hu DM, Chen H, Cai L, Bahkali AH, Hyde KD (2012c) Aquapeziza:
A new genus from freshwater and its morphological and phy-
logenetic relationships to Pezizaceae. Mycologia 104:540–546.
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Hu DM, Liu F, Cai L (2013a) Biodiversity of aquatic fungi in China.
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Hu XJ, Cheng ZY, Chen Y, Lu BS (2013b) Three new Chinese records
of freshwater fungi from Shandong Province. Mycosystema
32:1079–1085
Hu DM, Wang M, Cai L (2017) Phylogenetic assessment and taxo-
nomic revision of Mariannaea. Mycol Prog 16:271–283. https://
doi. org/ 10. 1007/ s11557- 016- 1252-2
Huang JE, Song HY, Ma J, Guan GX, Hu DM (2016) Xylohyphop-
sis aquatica sp. nov., a new aquatic hyphomycete from China.
Mycotaxon 131:391–394. https:// doi. org/ 10. 5248/ 131. 391
Huang JE, Song HY, Huang XG, Ma J, Hu DM (2017) Phaeomonilia
aquatica sp. nov., an aquatic hyphomycete from China. Myco-
taxon 132:919–923. https:// doi. org/ 10. 5248/ 132. 919
Huang SK, Jeewon R, Hyde KD, Bhat DJ, Wen TC (2018a) Novel taxa
within Nectriaceae: Cosmosporella gen. nov. and Aquanectria
sp. nov. from freshwater habitats in China. Cryptogam Mycol
39:169–192. https:// doi. org/ 10. 7872/ crym/ v39. iss2. 2018. 169
Huang SK, Jeewon R, Hyde KD, Jayarama Bhat D, Chomnunti P,
Wen TC (2018b) Beta-tubulin and actin gene phylogeny sup-
ports Phaeoacremonium ovale as a new species from freshwater
habitats in China. MycoKeys 41:1–15. https:// doi. org/ 10. 3897/
mycok eys. 41. 27536
Huber-Pestalozzi G (1944) Chytridium oocystidis (spec. nova?)—ein
Parasit auf Oocystis lacustris Chodat. Zeitschrift Für Hydrol
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Hudson HJ (1961) Heliscus submersus sp.nov., an aquatic hyphomycete
from Jamaica. Trans Br Mycol Soc 44:91-IN8. https:// doi. org/ 10.
1016/ s0007- 1536(61) 80011-9
Hudson HJ, Ingold CT (1960) Aquatic hyphomycetes from Jamaica.
Trans Br Mycol Soc 43:469-IN1. https:// doi. org/ 10. 1016/ s0007-
1536(60) 80029-0
Huldén L (1983) Laboulbeniales (Ascomycetes) of Finland and adja-
cent parts of the U.S.S.R. Karstenia 23:31–136. https:// doi. org/
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Hyde KD (1992a) Tropical Australian freshwater fungi. I. some asco-
mycetes. Aust Syst Bot 5:109–116. https:// doi. org/ 10. 1071/
SB992 0109
Hyde KD (1992b) Tropical Australian freshwater fungi. IV.* Halosar-
pheia aquatica sp. nov., Garethjonesia lacunosispora gen. & sp.
nov. and Ophioceras dolichostomum (Ascomycetes). Aust Syst
Bot 5:407–414. https:// doi. org/ 10. 1071/ SB992 0407
Hyde KD (1992c) Tropical Australian freshwater fungi. Il* Annu-
latascus velatispora gen. et sp. nov., A. bipolaris sp. nov. and
Nais aquatica sp. nov. (Ascomycetes). Aust Syst Bot 5:117–124.
https:// doi. org/ 10. 1071/ SB992 0117
Hyde KD (1993a) Tropical Aaustralian freshwater fungi. VI*. Tia-
rosporella paludosa and Clohesyomyces aquaticus gen. et sp.
nov. (Coelomycetes). Aust Syst Bot 6:169–173. https:// doi. org/
10. 1071/ SB993 0169
Hyde KD (1993b) Tropical Australian freshwater fungi. V.* Bombardia
sp., Jahnula australiensis sp. nov., Savoryella aquatica sp. nov.
and S. lignicola sp. nov. Aust Syst Bot 6:161–167. https:// doi.
org/ 10. 1071/ SB993 0161
Hyde KD (1994a) Aquatic fungi on rachides of Livistona in the West-
ern Province of Papua New Guinea. Mycol Res 98:719–725.
https:// doi. org/ 10. 1016/ S0953- 7562(09) 81043-5
Hyde KD (1994b) The genus Savoryella from freshwater habitats,
including S. grandispora sp. nov. Mycoscience 35:59–61. https://
doi. org/ 10. 1007/ BF022 68529
Hyde KD (1995a) The genus Massarina, with a description of M.
eburnea and an annotated list of Massarina names. Mycol Res
99:291–296. https:// doi. org/ 10. 1016/ S0953- 7562(09) 80902-7
Hyde KD (1995b) Tropical Australian freshwater fungi. VII. New
genera and species of ascomycetes. Nov Hedwigia 61:119–140
Hyde KD (1995c) Tropical Australian freshwater fungi. IX. Vaginatis-
pora aquatica gen. et sp. nov. Nov Hedwigia 61:233–241
Hyde KD (1995d) Tropical Australasian fungi. VII. New genera and
species of ascomycetes. Nov Hedwigia 61:119–140
Hyde KD (1995e) Tropical Australian freshwater fungi. VIII. Bertia
convolutispora sp. nov. Nov Hedwigia 61:141–146
Hyde KD (1996) Tropical Australian freshwater fungi. X Submersi-
sphaeria aquatica gen. et sp. nov. Nov Hedwigia 62:171–175
Hyde KD (1997) Additions to the genus Linocarpon (Ascomycetes:
Hyponectriaceae). Bot J Linn Soc 123:109–131. https:// doi. org/
10. 1111/j. 1095- 8339. 1997. tb014 07.x
Hyde KD (2007) Saprobic microfungi. In: Jones EBG, Hyde KD, Viki-
neswary S (eds) Malaysian fungal diversity. Mushroom Research
Centre, University of Malaya, and Ministry of Natural resources
and Environment Malaysia, Kuala Lumpur, pp 141–162
Hyde KD, Aptroot A (1998a) Tropical freshwater species of the gen-
era Massarina and Lophiostoma (ascomycetes). Nov Hedwigia
66:489–502
Hyde KD, Aptroot A (1998b) A new freshwater species of Herpotri-
chia from the tropics. Nov Hedwigia 66:247–249. https:// doi. org/
10. 1127/ nova. hedwi gia/ 66/ 1998/ 247
Hyde KD, Fröhlich J (1998) Fungi from palms XXXVII. The genus
Astrosphaeriella, including ten new species. Sydowia 50:81–132
Hyde KD, Goh TK (1997) Fungi on submerged wood in a small stream
on Mt Lewis, north Queensland, Australia. Muelleria 10:145–157
Hyde KD, Goh TK (1998a) Fungi on submerged wood in Lake Barrine,
north Queensland, Australia. Mycol Res 102:739–749. https://
doi. org/ 10. 1017/ S0953 75629 70058 68
Hyde KD, Goh TK (1998b) Tropical Australian freshwater fungi XIII.
A new species of Anthostomella and its sporodochial Geniculo-
sporium anamorph. Nov Hedwigia 67:225–233. https:// doi. org/
10. 1127/ nova. hedwi gia/ 67/ 1998/ 225
Hyde KD, Goh TK (1998c) Fungi on submerged wood in the Riviere
St Marie-Louis, the Seychelles. S Afr J Bot 64:330–336. https://
doi. org/ 10. 1016/ S0254- 6299(15) 30920-0
Hyde KD, Goh TK (1999a) Fungi on submerged wood from the River
Coln, England. Mycol Res 103:1561–1574. https:// doi. org/ 10.
1017/ S0953 75629 90089 89
Hyde KD, Goh TK (1999b) Tropical Australian freshwater fungi. XVI.
Some new melanommataceous fungi from woody substrata and
a key to genera of lignicolous loculoascomycetes in freshwater.
Nov Hedwigia 68:251–272. https:// doi. org/ 10. 1127/ nova. hedwi
gia/ 68/ 1999/ 251
Hyde KD, Sarma VV (2006) Biodiversity and ecological observations
on filamentous fungi of mangrove palm Nypa fruticans Wurumb
(Liliopsida-Arecales) along the Tutong River, Brunei. Indian J
Mar Sci 35:297–307
Hyde KD, Seifert KA (1992) Tropical Australian freshwater fungi.
III. Candelosynnema ranunculosporum, a new genus and spe-
cies of synnematous Hyphomyceteshyphomycetes. Aust Syst
Bot 5:401–405
Hyde KD, Steinke TS (1996) Two new species of Delitschia from
submerged wood. Mycoscience 37:99–102. https:// doi. org/ 10.
1007/ BF024 61463
Hyde KD, Wong SW (1999a) Didymella aptrootii sp. nov. from bam-
boo submerged in freshwater. Australas Mycol 18:54–62
Hyde KD, Wong SW (1999b) Tropical Australian freshwater fungi.
XV. The ascomycete genus Jahnula, with five new species and

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one new combination. Nov Hedwigia 68:489–509. https:// doi.
org/ 10. 1127/ nova. hedwi gia/ 68/ 1999/ 489
Hyde KD, Wong SW (2000) Annulatascus fusiformis sp. nov., a new
freshwater ascomycete from the Philippines. Mycologia 92:553–
557. https:// doi. org/ 10. 2307/ 37615 15
Hyde KD, Goh TK, Steinke T (1996a) Nawawia dendroidea, a new
synnematous hyphomycete from submerged Phragmites in South
Africa. Mycol Res 100:810–814. https:// doi. org/ 10. 1016/ S0953-
7562(96) 80026-8
Hyde KD, Wong SW, Jones EBG (1996b) Tropical Australian freshwa-
ter fungi. XI 1 Mamillisphaeria dimorphospora gen. et sp. nov.
and notes on freshwater ascomycetes with dimorphic ascospores.
Nov Hedwigia 62:513–520
Hyde KD, Read SJ, Jones EBG, Moss ST (1997) Tropical Australian
freshwater fungi. XII Rivulicola incrustata gen. et sp. nov. and
notes on Ceratosphaeria lampadophora. Nov Hedwigia 64:185–
196. https:// doi. org/ 10. 1127/ nova. hedwi gia/ 64/ 1997/ 185
Hyde KD, Goh TK, Steinke TD (1998a) Fungi on submerged wood in
the Palmiet River, Durban, South Africa. S Afr J Bot 64:151–
162. https:// doi. org/ 10. 1016/ S0254- 6299(15) 30860-7
Hyde KD, Wong SW, Jones EBG (1998b) Diluviocola capensis gen.
and sp. nov., a freshwater ascomycete with unique polar caps on
the ascospores. Fungal Divers 1:133–146
Hyde KD, Ho WH, Tsui CKM (1999a) The genera Aniptodera, Halo-
sarpheia, Nais and Phaeonectriella from freshwater habitats.
Mycoscience 40:165–183. https:// doi. org/ 10. 1007/ BF024 64295
Hyde KD, Wong SW, Jones EBG (1999b) Cataractispora gen. nov.
with three new freshwater lignicolous species. Mycol Res
103:1019–1031. https:// doi. org/ 10. 1017/ S0953 75629 80080 65
Hyde KD, Ho WH, Jones EBG, Tsui CKM, Wong WSW (2000) Tor-
rentispora fibrosa gen. sp. nov. (Annulatascaceae) from freshwa-
ter habitats. Mycol Res 104:1399–1403. https:// doi. org/ 10. 1017/
S0953 75620 00027 81
Hyde KD, Fryar S, Tian Q, Bahkali AH, Xu J (2016a) Lignicolous
freshwater fungi along a north-south latitudinal gradient in the
Asian/Australian region; can we predict the impact of global
warming on biodiversity and function? Fungal Ecol 19:190–200.
https:// doi. org/ 10. 1016/j. funeco. 2015. 07. 002
Hyde KD, Hongsanan S, Jeewon R, Bhat DJ, McKenzie EHC, Jones
EBG, Phookamsak R, Ariyawansa HA, Boonmee S, Zhao Q,
Abdel-Aziz FA, Abdel-Wahab MA, Banmai S, Chomnunti P, Cui
BK, Daranagama DA, Das K, Dayarathne MC, de Silva NI, Dis-
sanayake AJ, Doilom M, Ekanayaka AH, Gibertoni TB, Góes-
Neto A, Huang SK, Jayasiri SC, Jayawardena RS, Konta S, Lee
HB, Li WJ, Lin CG, Liu JK, Lu YZ, Luo ZL, Manawasinghe
IS, Manimohan P, Mapook A, Niskanen T, Norphanphoun C,
Papizadeh M, Perera RH, Phukhamsakda C, Richter C, André
ALCM, Drechsler-Santos ER, Senanayake IC, Tanaka K, Tenna-
koon TMDS, Thambugala KM, Tian Q, Tibpromma S, Thongbai
B, Vizzini A, Wanasinghe DN, Wijayawardene NN, Wu HX,
Yang J, Zeng XY, Zhang H, Zhang JF, Bulgakov TS, Camporesi
E, Bahkali AH, Amoozegar MA, Araujo-Neta LS, Ammirati JF,
Baghela A, Bhatt RP, Bojantchev D, Buyck B, da Silva GA, de
Lima CLF, de Oliveira RJV, de Souza CAF, Dai YC, Dima B,
Duong TT, Ercole E, Mafalda-Freire F, Ghosh A, Hashimoto A,
Kamolhan S, Kang JC, Karunarathna SC, Kirk PM, Kytövuori I,
Lantieri A, Liimatainen K, Liu ZY, Liu XZ, Lücking R, Medardi
G, Mortimer PE, Nguyen TTT, Promputtha I, Raj KNA, Reck
MA, Lumyong S, Shahzadeh-Fazeli SA, Stadler M, Soudi MR,
Su HY, Takahashi T, Tangthirasunun N, Uniyal P, Wang Y, Wen
TC, Xu JC, Zhang ZK, Zhao YC, Zhou JL, Zhu L (2016b) Fungal
diversity notes 367–490: taxonomic and phylogenetic contribu-
tions to fungal taxa. Fungal Divers 80:1–270. https:// doi. org/ 10.
1007/ s13225- 016- 0373-x
Hyde KD, Maharachchikumbura SSN, Hongsanan S, Samarakoon MC,
Lücking R, Pem D, Harishchandra D, Jeewon R, Zhao RL, Xu
JC, Liu JK, Al-Sadi AM, Bahkali AH, Elgorban AM (2017a)
The ranking of fungi: a tribute to David L. Hawksworth on his
70th birthday. Fungal Divers 84:1–23. https:// doi. org/ 10. 1007/
s13225- 017- 0383-3
Hyde KD, Norphanphoun C, Abreu VP, Bazzicalupo A, Thilini
Chethana KW, Clericuzio M, Dayarathne MC, Dissanayake AJ,
Ekanayaka AH, He MQ, Hongsanan S, Huang SK, Jayasiri SC,
Jayawardena RS, Karunarathna A, Konta S, Kušan I, Lee H, Li J,
Lin CG, Liu NG, Lu YZ, Luo ZL, Manawasinghe IS, Mapook A,
Perera RH, Phookamsak R, Phukhamsakda C, Siedlecki I, Soares
AM, Tennakoon DS, Tian Q, Tibpromma S, Wanasinghe DN,
Xiao YP, Yang J, Zeng XY, Abdel-Aziz FA, Li WJ, Senanayake
IC, Shang QJ, Daranagama DA, de Silva NI, Thambugala KM,
Abdel-Wahab MA, Bahkali AH, Berbee ML, Boonmee S, Bhat
DJ, Bulgakov TS, Buyck B, Camporesi E, Castañeda-Ruiz RF,
Chomnunti P, Doilom M, Dovana F, Gibertoni TB, Jadan M,
Jeewon R, Jones EBG, Kang JC, Karunarathna SC, Lim YW, Liu
JK, Liu ZY, Plautz HL, Lumyong S, Maharachchikumbura SSN,
Matočec N, McKenzie EHC, Mešić A, Miller D, Pawłowska J,
Pereira OL, Promputtha I, Romero AI, Ryvarden L, Su HY,
Suetrong S, Tkalčec Z, Vizzini A, Wen TC, Wisitrassameewong
K, Wrzosek M, Xu JC, Zhao Q, Zhao RL, Mortimer PE (2017b)
Fungal diversity notes 603–708: taxonomic and phylogenetic
notes on genera and species. Fungal Divers 87:1–235. https://
doi. org/ 10. 1007/ s13225- 017- 0391-3
Hyde KD, Chaiwan N, Norphanphoun C, Boonmee S, Camporesi E,
Chethana KWT, Dayarathne MC, de Silva NI, Dissanayake AJ,
Ekanayaka AH, Hongsanan S, Huang SK, Jayasiri SC, Jaya-
wardena RS, Jiang HB, Karunarathna A, Lin CG, Liu JK, Liu
NG, Lu YZ, Luo ZL, Maharachchimbura SSN, Manawasinghe
IS, Pem D, Perera RH, Phukhamsakda C, Samarakoon MC,
Senwanna C, Shang QJ, Tennakoon DS, Thambugala KM, Tib-
promma S, Wanasinghe DN, Xiao YP, Yang J, Zeng XY, Zhang
JF, Zhang SN, Bulgakov TS, Bhat DJ, Cheewangkoon R, Goh
TK, Jones EBG, Kang JC, Jeewon R, Liu ZY, Lumyong S, Kuo
CH, McKenzie EHC, Wen TC, Yan JY, Zhao Q (2018) Myco-
sphere notes 169–224. Mycosphere 9:271–430. https:// doi. org/
10. 5943/ mycos phere/9/ 2/8
Hyde KD, Tennakoon DS, Jeewon R, Bhat DJ, Maharachchikumbura
SSN, Rossi W, Leonardi M, Lee HB, Mun HY, Houbraken J,
Nguyen TTT, Jeon SJ, Frisvad JC, Wanasinghe DN, Lücking
R, Aptroot A, Cáceres MES, Karunarathna SC, Hongsanan S,
Phookamsak R, de Silva NI, Thambugala KM, Jayawardena RS,
Senanayake IC, Boonmee S, Chen J, Luo ZL, Phukhamsakda
C, Pereira OL, Abreu VP, Rosado AWC, Bart B, Randrianjo-
hany E, Hofstetter V, Gibertoni TB, da Soares AMS, Plautz HL,
Sotão HMP, Xavier WKS, Bezerra JDP, de Oliveira TGL, de
Souza-Motta CM, Magalhães OMC, Bundhun D, Harishchandra
D, Manawasinghe IS, Dong W, Zhang SN, Bao DF, Samara-
koon MC, Pem D, Karunarathna A, Lin CG, Yang J, Perera RH,
Kumar V, Huang SK, Dayarathne MC, Ekanayaka AH, Jayasiri
SC, Xiao Y, Konta S, Niskanen T, Liimatainen K, Dai YC, Ji
XH, Tian XM, Mešić A, Singh SK, Phutthacharoen K, Cai L,
Sorvongxay T, Thiyagaraja V, Norphanphoun C, Chaiwan N, Lu
YZ, Jiang HB, Zhang JF, Abeywickrama PD, Aluthmuhandiram
JVS, Brahmanage RS, Zeng M, Chethana T, Wei D, Réblová
M, Fournier J, Nekvindová J, do Nascimento Barbosa R, dos
Santos JEF, de Oliveira NT, Li GJ, Ertz D, Shang QJ, Phillips
AJL, Kuo CH, Camporesi E, Bulgakov TS, Lumyong S, Jones
EBG, Chomnunti P, Gentekaki E, Bungartz F, Zeng XY, Fryar
S, Tkalčec Z, Liang J, Li G, Wen TC, Singh PN, Gafforov Y,
Promputtha I, Yasanthika E, Goonasekara ID, Zhao RL, Zhao Q,
Kirk PM, Liu JK, Yan JY, Mortimer PE, Xu J, Doilom M (2019)
Fungal diversity notes 1036–1150: taxonomic and phylogenetic
contributions on genera and species of fungal taxa. Fungal Divers
96:1–242. https:// doi. org/ 10. 1007/ s13225- 019- 00429-2

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Hyde KD, de Silva N, Jeewon R, Bhat JD, Phookamsak R, Doilom
M, Boonmee S, Jayawardena RS, Maharachchikumbura SSN,
Senanayake IC, Manawasinghe IS, Liu NG, Abeywickrama PD,
Chaiwan N, Karunarathna A, Pem D, Lin CG, Sysouphanthong
P, Luo ZL, Wei D, Wanasinghe DN, Norphanphoun C, Ten-
nakoon DS, Samarakoon MC, Jayasiri S, Jiang H, Zeng X, Li
J, Wijesinghe S, Devadatha B, Goonasekara I, Brahmanage R,
Yang E, Aluthmuhandiram J, Dayarathne M, Marasinghe D, Li
W, Dissanayake L, Dong W, Huanraluek N, Lumyong S, Liu J,
Karunarathna S, Jones EBG, Al-Sadi A, Xu J, Harishchandra
D, Sarma VV (2020a) AJOM new records and collections of
fungi: 1–100. Asian J Mycol 3:22–294. https:// doi. org/ 10. 5943/
ajom/3/ 1/3
Hyde KD, Dong Y, Phookamsak R, Jeewon R, Bhat DJ, Jones EBG,
Liu N-G, Abeywickrama PD, Mapook A, Wei D, Perera RH,
Manawasinghe IS, Pem D, Bundhun D, Karunarathna A, Ekanay-
aka AH, Bao D-F, Li J, Samarakoon MC, Chaiwan N, Lin C-G,
Phutthacharoen K, Zhang S-N, Senanayake IC, Goonasekara ID,
Thambugala KM, Phukhamsakda C, Tennakoon DS, Jiang H-B,
Yang J, Zeng M, Huanraluek N, Liu J-K, Wijesinghe SN, Tian
Q, Tibpromma S, Brahmanage RS, Boonmee S, Huang S-K, Thi-
yagaraja V, Lu Y-Z, Jayawardena RS, Dong W, Yang E-F, Singh
SK, Singh SM, Rana S, Lad SS, Anand G, Devadatha B, Niran-
jan M, Sarma VV, Liimatainen K, Aguirre-Hudson B, Niskanen
T, Overall A, Alvarenga RLM, Gibertoni TB, Pfliegler WP, Hor-
váth E, Imre A, Alves AL, da Silva Santos AC, Tiago PV, Bul-
gakov TS, Wanasinghe DN, Bahkali AH, Doilom M, Elgorban
AM, Maharachchikumbura SSN, Rajeshkumar KC, Haelewaters
D, Mortimer PE, Zhao Q, Lumyong S, Xu J, Sheng J (2020b)
Fungal diversity notes 1151–1276: taxonomic and phylogenetic
contributions on genera and species of fungal taxa. Fungal Divers
100:5–277. https:// doi. org/ 10. 1007/ s13225- 020- 00439-5
Hyde KD, Jeewon R, Chen YJ, Bhunjun CS, Calabon MS, Jiang
HB, Lin CG, Norphanphoun C, Sysouphanthong P, Pem D,
Tibpromma S, Zhang Q, Doilom M, Jayawardena RS, Liu JK,
Maharachchikumbura SSN, Phukhamsakda C, Phookamsak R,
Al-Sadi AM, Thongklang N, Wang Y, Gafforov Y, Jones EBG,
Lumyong S (2020c) The numbers of fungi: is the descriptive
curve flattening? Fungal Divers 103:219–271. https:// doi. org/ 10.
1007/ s13225- 020- 00458-2
Hyde KD, Norphanphoun C, Maharachchikumbura SSN, Bhat DJ,
Jones EBG, Bundhun D, Chen Y-J, Bao D-F, Boonmee S,
Calabon MS, Chaiwan N, Chethana K, Dai D, Dayarathne M,
Devadatha B, Dissanayake A, Dissanayake L, Doilom M, Dong
W, Fan X, Goonasekara I, Hongsanan S, Huang S, Jayawardena
R, Jeewon R, Karunarathna A, Konta S, Kumar V, Lin C, Liu
J, Liu N, Luangsa-ard J, Lumyong S, Luo Z, Marasinghe D,
McKenzie E, Niego A, Niranjan M, Perera R, Phukhamsakda
C, Rathnayaka A, Samarakoon M, Samarakoon S, Sarma VV,
Senanayake I, Shang Q, Stadler M, Tibpromma S, Wanasinghe
D, Wei D, Wijayawardene N, Xiao Y, Yang J, Zeng X, Zhang S,
Xiang M (2020d) Refined families of Sordariomycetes. Myco-
sphere 11:305–1059. https:// doi. org/ 10. 5943/ mycos phere/ 11/1/7
Hyde KD, Bao DF, Hongsanan S, Chethana KWT, Yang J, Suwan-
narach N (2021) Evolution of freshwater Diaporthomycetidae
(Sordariomycetes) provides evidence for five new orders and
six new families. Fungal Divers 107:71–105. https:// doi. org/ 10.
1007/ s13225- 021- 00469-7
Hyliš M, Oborník M, Nebesářová J, Vávra J (2007) Aquatic tetrasporo-
blastic microsporidia from caddis flies (Insecta, Trichoptera):
Characterisation, phylogeny and taxonomic reevaluation of the
genera Episeptum Larsson, 1986, Pyrotheca Hesse, 1935 and
Cougourdella Hesse, 1935. Eur J Protistol 43:205–224. https://
doi. org/ 10. 1016/j. ejop. 2007. 03. 004
Hyliš M, Oborník M, Nebesářová J, Vávra J (2013) Description
and phylogeny of Zelenkaia trichopterae gen. et sp. nov.
(Microsporidia), an aquatic microsporidian parasite of caddis-
flies (Trichoptera) forming spore doublets. J Invertebr Pathol
114:11–21. https:// doi. org/ 10. 1016/j. jip. 2013. 04. 010
Inácio J, Portugal L, Spencer-Martins I, Fonseca Á (2005) Phyllo-
plane yeasts from Portugal: Seven novel anamorphic species in
the Tremellales lineage of the Hymenomycetes (Basidiomycota)
producing orange-coloured colonies. FEMS Yeast Res. https://
doi. org/ 10. 1016/j. femsyr. 2005. 05. 007
Inderbitzin P (2000) Ceratostomella hyalocoronata, a new pyrenomy-
cete from a stream in southern China. Mycoscience 41:167–169.
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Authors and Aliations
MarkS.Calabon1,2 · KevinD.Hyde1,2,3,4 · E.B.GarethJones5 · Zong‑LongLuo6 · WeiDong4 ·
VedprakashG.Hurdeal1,2 · EleniGentekaki1,2 · WalterRossi7 · MarcoLeonardi7 ·
VinodhiniThiyagaraja1,3,8,9,10 · AnisS.Lestari1,2,11 · Hong‑WeiShen1,2,6 · Dan‑FengBao1,6,10 ·
NattawutBoonyuen12 · MingZeng1,2,13
1 Center ofExcellence inFungal Research, Mae Fah Luang
University, ChiangRai57100, Thailand
2 School ofScience, Mae Fah Luang University,
ChiangRai57100, Thailand
3 Department ofBiology, Faculty ofScience, Chiang Mai
University, ChiangMai50200, Thailand
4 Innovative Institute forPlant Health, Zhongkai University
ofAgriculture andEngineering, Guangzhou510225, China
5 Department ofBotany andMicrobiology, College ofScience,
King Saud University, P.O. Box2455, Riyadh11451,
SaudiArabia
6 College ofAgriculture andBiological Sciences, Dali
University, Dali671003, China
7 Department MeSVA, Section Environmental Sciences,
University ofL’Aquila, 67100Coppito, AQ, Italy
8 Centre forMountain Futures (CMF), CAS Key
Laboratory forPlant Biodiversity andBiogeography
ofEast Asia (KLPB), Kunming Institute ofBotany,
Chinese Academy ofScience, Kunming650201, Yunnan,
People’sRepublicofChina
9 CIFOR-ICRAF China Program, World Agroforestry
(ICRAF), Kunming650201, People’sRepublicofChina
10 Department ofEntomology andPlant Pathology, Faculty
ofAgriculture, Chiang Mai University, ChiangMai50200,
Thailand
11 Research Center forBiomaterials, Research Organization
forLife Sciences Center, National Research andInnovation
Agency (BRIN), Cibinong16911, Indonesia
12 Plant-Microbe Interaction Research Team (APMT),
Integrative Crop Biotechnology andManagement Research
Group (ACBG), National Center forGenetic Engineering
andBiotechnology (BIOTEC), PathumThani12120,
Thailand
13 Key Laboratory forPlant Diversity andBiogeography
ofEast Asia, Kunming Institute ofBotany, Chinese Academy
ofSciences, Kunming650201, Yunnan, China