Introduction

Shearer (1993) defined freshwater ascomycetes in a broad ecological sense as “all ascomycetes that occur on submerged or partially submerged substrates in aquatic habitats”. Freshwater fungi have also been defined as “fungi that the whole or part of their life cycle rely on freshwater” (Thomas 1996). The definition of Shearer (1993) has been regarded as the best current working definition for the freshwater ascomycetes (Cai et al. 2003a, 2014), and we follow the definition of Shearer (1993) in this study.

Lignicolous freshwater fungi are those that grow on submerged woody debris in freshwater streams, ponds, lakes, tree hollows, peat swamps and dams (Ho et al. 2002a; Vijaykrishna et al. 2005; Hyde et al. 2016a). They play an important role in nutrient and carbon cycling, biological diversity and ecosystem functioning of freshwater ecosystems with their ability to decompose lignocellulose in woody litter, softening the wood and releasing nutrients (Hyde et al. 2016a). The lignicolous freshwater fungi are highly diverse in the classes Dothideomycetes and Sordariomycetes (Hyde et al. 2013; Maharachchikumbura et al. 2016; Liu et al. 2017; Lu et al. 2018a). There are a few freshwater taxa that belong to Eurotiomycetes (Liu et al. 2015a, b), Orbiliomycetes (Swe et al. 2009) and rarely Basidiomycetes (Hyde and Goh 1998a, 1999; Jones et al. 2014) collected from submerged wood in freshwater habitats.

Sordariomycetes is the second largest class of Ascomycota (Kirk et al. 2008; Hyde et al. 2013; Maharachchikumbura et al. 2015, 2016). Species of Sordariomycetes are mainly characterized by non-lichenized, perithecial ascomata and inoperculate unitunicate or non-fissitunicate asci (Zhang et al. 2006; Kirk et al. 2008). The class Sordariomycetes has a cosmopolitan distribution and accommodates mostly terrestrial taxa (Tang et al. 2007; Maharachchikumbura et al. 2015, 2016; Hongsanan et al. 2017), but there are several sordariomycetous species that have been reported from aquatic habitats (Hyde and Wong 2000; Cai and Hyde 2007; Raja et al. 2009a, b; Réblová et al. 2010, 2016a; Shearer et al. 2010; Hu et al. 2013; Cai et al. 2014; Jones et al. 2015; Lu et al. 2016; Su et al. 2016; Yang et al. 2017, 2018a, b; Luo et al. 2018a, b).

Freshwater Sordariomycetes play an important role in ecosystem functioning and many of them have shown potential application value (Cai et al. 2014). Annulatascaceous, distoseptisporaceous, pleurotheciaceous and halosphaeriaceous species are the most typical and common freshwater Sordariomycetes on submerged wood (Hyde et al. 1999a; Cai et al. 2014; Li et al. 2016a; Réblová et al. 2016a; Zhang et al. 2017a, b; Luo et al. 2018a; Yang et al. 2018a). The modern classification system of the overall Kingdom of Fungi, as well as Sordariomycetes, have been significantly improved with the utilization of molecular sequence data (Hibbett et al. 2007; Maharachchikumbura et al. 2015; Spatafora et al. 2017). Cai et al. (2014) provided a phylogenetic analysis of freshwater Sordariomycetes based on LSU sequence data. Their analysis showed that freshwater Sordariomycetes are scattered in three subclasses, Sordariomycetidae, Hypocreomycecetidae and Xylariomycetidae including 13 orders. However, during the last four years, several Sordariomycetes species have been introduced comprising new genera, families, orders, subclasses and some of them were collected from freshwater habitats (Réblová et al. 2015a, b, 2016a, b; Maharachchikumbura et al. 2015, 2016; Su et al. 2016; Hongsanan et al. 2017; Wijayawardene et al. 2017, 2018; Yang et al. 2017, 2018a, b; Zhang et al. 2017a, b; Song et al. 2018a, b; Wei et al. 2018).

In this study, 129 fresh isolates of sordariomyceteous taxa were collected from freshwater habitats in China and Thailand. One new order, two new families, three new genera, 47 new species, two new combinations and 9 new records are introduced. A modified backbone tree based on phylogenetic analysis of combined LSU, SSU, RPB2 and TEF1α sequence data of the freshwater Sordariomycetes is provided. Phylogenetic trees under genus or family level are also constructed in order to confirm the classification of the new isolates.

Materials and methods

Sample collection and specimen examination

Submerged decaying wood samples were collected from freshwater habitats in Yunnan Province (China) and Chiang Rai, Chiang Mai (Thailand) from 2013 to 2018. The process of morphological studies follows Luo et al. (2018a). Single spore isolates were obtained following the method of Chomnunti et al. (2014) and cultured according to Liu et al. (2010). Herbarium materials are deposited in the Herbarium of Mae Fah Luang University (Herb. MFLU), Chiang Rai, Thailand, Herbarium of Cryptogams Kunming Institute of Botany Academia Sinica (Herb. HKAS) and Dali University (Herb. DLU), Yunnan, China. Living cultures are deposited at Mae Fah Luang University Culture Collection (MFLUCC), Thailand, Kunming Institute of Botany culture collection(KUMCC) and culture collection at Dali University (DLUCC) China. Facesoffungi and Index Fungorum numbers are provided (Jayasiri et al. 2015; Index Fungorum 2019). New taxa are established based on guidelines outlined by Jeewon and Hyde (2016). Taxa deposited in HKU(M) mostly lack sequence data and were moved to IFRD in Kunming, China. These need to be recollected so that reference specimens (sensu Ariyawansa et al. 2014) can be designated.

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from fungal mycelium grown on PDA or MEA at room temperature by using a EZ gene TM Fungal gDNA kit (GD2416) according to the manufacturer’s instructions. The gene regions of the large subunit of the nuclear ribosomal DNA (LSU), the internal transcribed spacers (ITS), the small subunit of the nuclear ribosomal DNA (SSU), the translation elongation factor (TEF1α) and RNA polymerase II subunit 2 (RPB2) were amplified using the primer pairs LR0R/LR7 (Vilgalys and Hester 1990), ITS5/ITS4, NS1/NS4 (White et al. 1990), 983F/2218R (Rehner and Buckley 2005), fRPB2-5F/fRPB2-7cR (Liu et al. 1999) respectively. The ITS, LSU, SSU, RPB2 and TEF1α amplification reactions were carried out using the method described by Luo et al. (2018a). DNA sequencing were performed with the primers mentioned above at Tsingke Biological Engineering Technology and Services Co., Ltd (Yunnan, P.R. China).

Phylogenetic analysis

Sequences generated from different primers were analyzed with other sequences obtained from GenBank. The sequences were deposited in GenBank and the accession numbers in the analyses were provided in Supplementary material 1. The sequence data were aligned using MAFFT v.7.110 online program (http://mafft.cbrc.jp/alignment/server/) (Katoh and Standley 2013) and manually adjusted via BioEdit v.7.2.3 (Hall 1999).

“ALTER” (Glez-Peña et al. 2010) was used to format the aligned fasta file for RAxML analysis. Maximum likelihood (ML) analysis was performed at the CIPRES Science Gateway v.3.3 (http://www.phylo.org/portal2/; Miller et al. 2010) using RAxML v.8.2.10 as part of the “RAxML-HPC2 on XSEDE” tool (Stamatakis et al. 2008; Stamatakis 2014). All free model parameters were estimated by RAxML with ML estimates of 25 per site rate categories. The final ML search was conducted using the GTRGAMMA + I model. The best scoring tree was selected with a final likelihood value of − 194313.319877. Phylogenetic tree was visualized using FigTree v1.4.0 (http://tree.bio.ed.ac.uk/software/figtree/, Rambaut 2012).

Results

Phylogenetic analysis of combined LSU, SSU, RPB2 and TEF1α sequence data

The combined LSU, SSU, RPB2 and TEF1α sequence dataset comprised 857 taxa with Dothidea sambuci (AFTOL ID 274), Leotia lubrica (AFTOL ID 1) and Microglossum rufum (AFTOL ID 1292) as the outgroup taxa. The combined aligned sequence matrix comprises LSU (768 bp), SSU (884 bp), RPB2 (850 bp) and TEF1α (815 bp) sequence data for 857 taxa with a total of 3317 characters (Including the gaps), of which 1046 characters were constant, 391 variable characters were parsimony-uninformative and 1880 characters were parsimony informative. The RAxML tree is shown in Fig. 1.

Fig. 1
figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1figure 1

Phylogram generated from maximum likelihood analysis based on combined LSU, SSU, RPB2 and TEF1α sequence data for the species of Sordariomycetes. RAxML bootstrap support values equal to or greater than 75% are given above the nodes. Newly generated sequences are in red. Strains of known species collected from freshwater habitats are marked as “green circle” followed by strain number. Ex-type strains are in bold

In the phylogenetic tree (Fig. 1), the 854 strains (including 129 fresh isolates) of Sordariomycetes included in the analysis cluster into six subclasses: Sordariomycetidae, Hypocreomycetidae and Xylariomycetidae as in the previous treatments of Lumbsch and Huhndorf (2010), Diaporthomycetidae and Lulworthiomycetidae as suggested by Maharachchikumbura et al. (2015), as well as Savoryellomycetidae as suggested by Hongsanan et al. (2017). Of these included strains, 356 were isolated from freshwater habitats and distributed in 47 clades as follows:

Clade 1 represents the family Junewangiaceae with ten species which belong to three genera, viz. Dictyosporella, Junewangia and Sporidesmiella. In this study, we provide sequence data for Sporidesmiella novae-zelandiae and S. hyalosperma for the first time, and introduce a new species S. aquatica.

Clade 2 represents the family Pseudoproboscisporaceae with four species from four genera, viz. Aquaticola hyalomura, Cateractispora recepticuli, Diluvicola aquatica, Pseudoproboscispora thailandensis.

Clade 3 represents the family Atractosporaceae. Presently, Atractosporales comprises the single family Atractosporaceae which includes two genera. All taxa of Atractosporales are collected from freshwater habitats (Réblová et al. 2016b; Zhang et al. 2017a, b). Two freshwater fungal strains obtained from this study cluster together with other Atractospora species and represent an independent lineage with strong support (100% ML). We therefore introduce a new species Atractospora aquatica sp. nov.

Clade 4 represents the genus Cancellidium with two strains of Cancellidium applanatum. These two strains were collected from freshwater habitats (Zelski et al. 2014).

Clade 5 represents the family Conlariaceae including three freshwater species of Conlarium, and one Riomyces species collected from freshwater.

Clade 6 represents the order Xenospadicoidales which was introduced by Réblová et al. (2018). Five freshwater fungal strains obtained in this study cluster together and represent an independent lineage with strong support (100% ML) within Xenospadicoidaceae. A new genus Neospadicoides is introduce herein based on phylogeny and morphology.

Clade 7 represents the order Sporidesmiales with a single family Sporidesmiaceae. Most of the species in Sporidesmiaceae are reported from freshwater habitats. For the first time, Zhang et al. (2017a, b) provided the sexual morph for the genus Sporidesmium. In this study, we introduce two new species, Sporidesmium lageniforme and Sporidesmium lignicola, based on both morphological characters and phylogenetic evidence, while S. lignicola is reported with both of asexual and sexual morphs.

Clade 8 represents the family Rhamphoriaceae established by Réblová and Štěpánek (2018) which includes four genera. We introduce a new Rhodoveronaea species which is the first species described from freshwater in this family.

Clade 9 represents the family Acrodictyaceae established by Xia et al. (2017) with a single genus Acrodictys. A previously described species, Barbatosphaeria aquatica MFLUCC 18–0356 (Hyde et al. 2018), is synonymized under Acrodictys aquatica. We introduce a new species, Acrodictys fluminicola.

Clade 10 represents the monotypic genus Pseudostanjehughesia introduced by Yang et al. (2018a) with P. aquitropica as the type species and it was collected from freshwater habitats. We introduce the second species, P. lignicola sp. nov.

Clade 11 represents the family Papulosaceae established by Winka and Eriksson (2000). Papulosaceae is typified by the monotypic marine genus Papulosa (Winka and Eriksson 2000).

Clade 12 represents the species Sporidesmium tropicale (HKUCC 10838, MFLUCC 16–0185). The strain MFLUCC 16–0185 was obtained from the specimen collected from a freshwater habitat (Yang et al. 2018a).

Clade 13 represents the typical freshwater genus Bullimyces introduced by Ferrer et al. (2012).

Clade 14 represents the family Barbatosphaeriaceae which was introduced by Zhang et al. (2017a, b). We introduce a new species, Barbatosphaeria lignicola based on morphology and phylogeny.

Clade 15 represents the new order Distoseptisporales introduced herein. Su et al. (2016) introduced the family Distoseptisporaceae to accommodate Sporidesmium-like taxa based on morphology and phylogeny. Phylogenetic analysis based on combined LSU, SSU, RPB2 and TEF1α sequence data show that the species of Distoseptisporales cluster together with strong support and form a distinct, strongly supported clade within Diaporthomycetidae.

Clade 16 represents two strains of the genus Cyanoannulus. Zhang et al. (2017a, b) established the family Woswasiaceae to accommodate Woswasia, Xylochrysis and Cyanoannulus in Diaporthomycetidae families incertae sedis with weak support. In our phylogenetic analysis, species of Cyanoannulus formed a distinct clade basal to the order Distoseptisporales and therefore we suggest to place Cyanoannulus in Diaporthomycetidae genera incertae sedis.

Clade 17 represents the order Annulatascales introduced by Maharachchikumbura et al. (2015). The family Annulatascaceae is accepted in this order. Some species of Annulatascaceae are lacking sequence data and therefore it is not possible to resolve their phylogenetic affinities.

Clade 18 represents the order Myrmecridiales established by Crous et al. (2015) with a single family, Myrmecridiaceae. We introduce a new species Myrmecridium aquaticum. Presently, three species (M. aquaticum, M. fluviae, M. montsegurinum) are collected from freshwater habitats.

Clade 19 represents the family Ophioceraceae, with eight species which were reported from freshwater habitats. We introduce a new species, Ophioceras submersum.

Clade 20 represents the new family Ceratosphaeriaceae introduced herein. The order Magnaporthales was introduced by Thongkantha et al. (2009) to accommodate Magnaporthaceae based on morphological characters of both asexual and sexual morphs together with phylogenetic analyses of combined LSU and SSU sequence data. There are four families accepted in Magnaporthales, viz. Magnaporthaceae, Ophioceraceae, Pseudohalonectriaceae, Pyriculariaceae. We introduce the new family Ceratosphaeriaceae to accommodate Ceratosphaeria species.

Clade 21 represents the family Pseudohalonectriaceae established by Hongsanan et al. (2017) based on the genus Pseudohalonectria. Five strains collected from freshwater habitats are included in our phylogentic tree.

Clade 22 represents the family Magnaporthaceae introduced by Cannon (1994). We introduce a new monotypic genus Aquafiliformis.

Clade 23 represents the family Tirisporellaceae with a freshwater species, Thailandiomyces bisetulosus (BCC 00018, BCC 00200) which was introduced by Pinruan et al. (2008).

Clade 24 represents the family Jobellisiaceae with a freshwater species, Jobellisia guangdongensis (GD14–4) collected from southern China (Liu et al. 2011a, b).

Clade 25 represents the freshwater genus Hyalorostratum established by Raja et al. (2010) with Hyalorostratum brunneisporum as the type species.

Clade 26 represents the family Chaetosphaeriaceae established by Réblová et al. (1999) based on Chaetosphaeria (Tulasne and Tulasne 1863). We introduce 11 new species in this family, viz. Chaetosphaeria aquatica, C. catenulata, C. guttulata, C. submersa, Codinaea yunnanensis, Dictyochaeta cangshanensis, D. ellipsoidea, D. lignicola, D. submersa, Tainosphaeria lunata and T. obclavata. Chloridium aseptatum (MFLUCC 11–0216) (Wei et al. 2018) is synonymized under Chloridium gonytrichii.

Clade 27 represents the family Phyllachoraceae with five strains. Phyllachoraceae sp. WF33A was collected from freshwater habitats.

Clade 28 represents the single isolate of Clohiesia. The species Clohiesia corticola (HKUCC 3712) which was collected from freshwater, clusters with species of Phyllachoraceae with weak support.

Clade 29 represents the order Sordariales established by Hawksworth and Eriksson (1986) and comprises three families, viz. Chaetomiaceae, Sordariaceae and Lasiosphaeriaceae sensu lato. Six freshwater species are represented in this clade and most of them are in Lasiosphaeriaceae.

Clade 30 represents the family Cordanaceae. We introduce two new species herein, Cordana aquatica and C. lignicola, based on morphology and phylogeny. We also provide descriptions, illustrations and molecular sequence data for Cordana abramovii and C. terrestris.

Clade 31 represents the family Coniochaetaceae with five Coniochaeta species and Coniochaeta gigantospora (ILLS 60816) which was originally collected from a freshwater habitat (Raja et al. 2012).

Clade 32 represents the family Sporocadaceae and we introduce a new species, Seiridium aquaticum.

Clade 33 represents the family Amphisphaeriaceae established by Winter (1885). We introduce a new species, Lepteutypa aquatica.

Clade 34 represents the family Apiosporaceae established by Hyde et al. (1998a). A new species, Arthrinium aquaticum is introduced.

Clade 35 represents a new species Peroneutypa lignicola. The genus Peroneutypa has been accomodated in Diatrypaceae (Shang et al. 2018). In our multi-gene phylogenetic analysis, the new species forms a distinct subclade basal to Lopadostomataceae but close to Diatrypaceae, although this relationship is not supported.

Clade 36 represents the species Sporidesmium gyrinomorphum (MFLUCC 16–0186) introduced by Yang et al. (2018a, b).

Clade 37 represents the family Hypoxylaceae with four Hypoxylon strains. We introduce a new species Hypoxylon lignicola based on morphology and phylogeny. Wendt et al. (2018) emended Hypoxylaceae and accepted 14 genera in this family.

Clade 38 represents the family Nectriaceae including 66 strains. Lombard et al. (2015) accepted 47 genera in this family based on molecular sequence data. We introduce a new species Cosmospora aquatica, based on morphology and phylogeny. DNA sequence data for Aquanectria jacinthicolor, A. penicillioides, Chaetopsina beijingensis, Mariannaea samuelsii, M. superimposita and Paracremonium binnewijzendii are provided.

Clade 39 represents the family Stachybotriaceae established by Crous et al. (2014a) in the order Hypocreales to accommodate the genera Myrothecium, Peethambara and Stachybotrys. In this study, morphological characterization and DNA sequence data for Stachybotrys chartarum and S. chlorohalonata are provided.

Clade 40 represents the family Halosphaeriaceae. In this study, we introduce a new Ascosacculus species, A. fusiformis, based on morphology and phylogeny.

Clade 41 represents the new family Triadelphiaceae that we introduce herein based on its distinct morphology and molecular phylogeny.

Clade 42 represents the family Reticulascaceae. We introduced a new species herein, Cylindrotrichum submersum sp. nov. and synonymise Blastophorum aquaticum (Hyde et al. 2016) under Cylindrotrichum aquaticum.

Clade 43 represents one of most common and typical freshwater family Pleurotheciellaceae established by Réblová et al. (2016a). We introduce a new species, Phaeoisaria filiformis.

Clade 44 represents another typical freshwater fungal family Savoryellaceae. We introduce a new genus Dematiosporium in this family.

Clade 45 represents the order Conioscyphales established by Réblová et al. (2016a) with a single family Conioscyphaceae. We introduce two new species, Conioscypha aquatica and C. submersa.

Clade 46 represents the monotypic order Fuscosporellales established by Yang et al. (2016a) based on the family Fuscosporellaceae. Most of the species in this order are reported from freshwater habitats.

Clade 47 represents the genus Flammispora established by Pinruan et al. (2004a).

Taxonomy

Based on the outline and multigene phylogeny of freshwater Sordariomycetes provided in this study, freshwater Sordariomycetes are scattered in six sub-classes, viz. Diaporthomycetidae, Hypocreomycetidae, Lulworthiomycetidae, Savoryellomycetidae, Sordariomycetidae and Xylariomycetidae. In this section, we provide the notes for each freshwater genera and species including information on known distribution, herbarium and sequence data. The outline of freshwater Sordariomycetes based on literature up to November 2018 was provided in Supplementary material 2. Descriptions, illustrations and sequence data for the fungi collected from freshwater habitats in China and Thailand from 2013 to 2018 are also provided.


Detailed information of freshwater Sordariomycetes are as follows:

Diaporthomycetidae Senan. et al.

Annulatascales D’souza et al.

Annulatascaceae Wong et al.

Annulatascus K.D. Hyde, Aust. Syst. Bot. 5(1): 118 (1992)

Asexual morph Undetermined. Sexual morph Updated generic description and illustrations see Maharachchikumbura et al. (2016).

Type species: Annulatascus velatisporus K.D. Hyde, Aust. Syst. Bot. 5(1): 118 (1992)

Notes: Hyde (1992a) introduced the genus Annulatascus to accommodate two ascomycete species, A. velatisporus and A. bipolaris, with the former as the type species and both of them were collected from submerged decaying wood in Australia. There are 19 epithets of Annulatascus listed in Index Fungorum (December, 2018), however Hyde et al. (1999a, b) transferred Annulatascus bipolaris to the genus Cataractispora as C. bipolaris based on the morphological characters. Subsequently, Campbell and Shearer (2004) established a new genus Annulusmagnus for Annulatascus triseptatus based on LSU sequence data, and Luo et al. (2015) combined Annulatascus biatriisporus as Pseudoannulatascus biatriisporus. Réblová et al. (2018) synonymised Annulatascus biatriisporus and Pseudoannulatascus biatriisporus under Torrentispora biatriispora, based on phylogenetic analysis. Currently, 16 species are accepted in Annulatascus and most of them are reported from freshwater habitats in tropical areas (Barbosa et al. 2008; Shearer et al. 2010; Boonyuen et al. 2012; Hu et al. 2012a) and only two species (A. citrisporus and A. licualae) are known from terrestrial habitats on palm rachides (Fröhlich and Hyde 2000).

Annulatascus apiculatus F.R. Barbosa & Gusmão

Distribution: Brazil, on submerged decaying wood in a stream (Barbosa et al. 2008)

Asexual morph: Undetermined

Notes: Holotype HUEFS 134723. Sequence data is not available.

Annulatascus aquaticus Ho et al.

Distribution: China, Hong Kong, Tai Po Kau Forest stream, on submerged decaying wood (Ho et al. 1999a).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 4526 (now in IFRD). Sequence data is not available.

Annulatascus aquatorba Boonyuen & Sri-indrasutdhi

Distribution: Thailand, Narathiwat Province, Sirindhorn Peat Swamp Forest, on submerged wood test block of Erythrophleum teysmannii (Boonyuen et al. 2012).

Asexual morph: Undetermined

Notes: Holotype BBH 29936. LSU and SSU sequence data are available.

Annulatascus fusiformis K.D. Hyde & S.W. Wong

Distribution: Australia, Queensland, on submerged wood (Vijaykrishna and Hyde 2006); China, Yunnan Province, Yiliang county, Xiao Bai Long Mountain, on submerged culms of Phyllostachys bambusoides in a stream (Cai et al. 2006a); Philippines, Mindawao, Bukidnon, Impalutao, Natigbasan creek, on submerged wood (Hyde and Wong 2000).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 3102 (now in IFRD). Sequence data is not available.

Annulatascus hongkongensis Ho et al.

Distribution: China, Hong Kong, Plover Cove Reservoir, on submerged wood (Ho et al. 1999b).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 4702 (now in IFRD). LSU sequence data is available.

Annulatascus joannae Tsui et al.

Distribution: China, Hong Kong, Tai Po, Lam Tsuen River, on submerged wood (Tsui et al. 2002).

Asexual morph: Undetermined

Notes: Holotype IFRD 8648. Sequence data is not available.

Annulatascus lacteus Tsui et al.

Distribution: China, Hong Kong, Tai Po, Lam Tsuen River, on submerged wood (Tsui et al. 2002).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 4623 (now in IFRD). Sequence data is not available.

Annulatascus liputii L. Cai & K.D. Hyde

Distribution: Philippines, on submerged bamboo in Liput River (Cai et al. 2003b).

Asexual morph: Undetermined

Notes: Holotype PDD 75038. Sequence data is not available.

Annulatascus menglensis Hu et al.

Distribution: China, Yunnan Province, Mengla, Wudaoban Stream, on submerged wood (Hu et al. 2012a).

Asexual morph: Undetermined

Notes: Holotype IFRDSC 002–017. Sequence data is not available.

Annulatascus nilensis Abdel-Wahab & Abdel-Aziz

Distribution: Egypt, Sohag, Nile River, on decayed submerged stems of Phragmites australis (Abdel-Wahab et al. 2011).

Asexual morph: Undetermined

Notes: Holotype IMI 397966. LSU sequence data is available.

Annulatascus palmietensis Goh et al.

Distribution: Brunei, Tutong River, on submerged wood (Fryar et al. 2004; Hyde and Sarma 2006); South Africa, Durban, Palmiet River, on submerged wood (Hyde et al. 1998b).

Asexual morph: Undetermined

Notes: Holotype IFRD 8649; Sequence data is not available.

Annulatascus saprophyticus Z.L. Luo & K.D. Hyde

Distribution: Thailand, Chiang Mai Province, on submerged wood in a stream (Luo et al. 2015).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–0070, isotype HKAS 86445. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available and obtained from ex-type culture.

Annulatascus tropicalis Ranghoo & K.D. Hyde

Distribution: China, Hong Kong, Tai Po, Plover Cove Reservoir, on submerged wood (Tsui et al. 2002).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 5253 (now in IFRD). Sequence data is not available.

Annulatascus velatisporus K.D. Hyde

Distribution: Australia, Queensland, on submerged wood (Hyde 1992a; Dayarathne et al. 2016); Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Anak, on submerged wood, China, Hong Kong, New Territories, Tai Po Kau Country Park, on twigs submerged in a stream; Philippines, Mindanao, Bukidnon, Impalutao, Natigbasan Creek, on submerged wood (Wong et al. 1999a); Japan, Koito River, on submerged wood (Tsui et al. 2001a); India, Western Ghats, Kali River, submerged woody litter (Sudheep and Sridhar 2011); Malaysia, Lipur Lentang Nature Reserve, on submerged wood in a stream (Ho et al. 2001); South Africa, Durban, Palmiet River, on submerged wood (Hyde et al. 1998b); Thailand, Chiang Mai Province, on submerged wood (Hu et al. 2012b).

Asexual morph: Undetermined

Notes: Holotype BRIP 17373, epitype MFLU 16–2204. LSU and SSU sequence data are available.


Annulusmagnus J. Campb. & Shearer, Mycologia 96(4): 826 (2004)

Asexual morph Undetermined. Sexual morph Description and illustrations see Campbell and Shearer (2004).

Type species: Annulusmagnus triseptatus (Wong et al.) J. Campb. & Shearer, Mycologia 96(4): 826 (2004)

Notes: Campbell and Shearer (2004) introduced the genus Annulusmagnus for Annulatascus triseptatus based on LSU sequence data. Annulusmagnus triseptatus was first collected from submerged wood in Brunei (Wong et al. 1999a) and subsequently reported from Australia, Canada and the USA, on submerged corticated or decorticated wood in both lotic and lentic habitats (Campbell and Shearer 2004). There is only one species reported in this genus (Campbell and Shearer 2004).

Annulusmagnus triseptatus (Wong et al.) J. Campb. & Shearer

Annulatascus triseptatus Wong et al., Mycol. Res. 103(5): 563 (1999)

Distribution: Australia, Koah, Clohiesy River, on submerged wood (Campbell and Shearer 2004); Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Anak, on submerged wood (Wong et al. 1999a); Canada, Ontario and Manitoba, on submerged wood; USA, on submerged wood; Venezuela, on submerged wood (Campbell and Shearer 2004).

Asexual morph: Undetermined

Notes: Holotype IFRD 8650. LSU, SSU and RPB2 sequence data are available.


Aqualignicola Ranghoo et al., Mycol. Res. 105(5): 628 (2001)

Asexual morph Undetermined. Sexual morph Description and illustrations see Ranghoo et al. (2001) and Hu et al. (2012a).

Type species: Aqualignicola hyalina Ranghoo et al., Mycol. Res. 105(5): 628 (2001)

Notes: Aqualignicola was introduced by Ranghoo et al. (2001) with Aqualignicola hyalina as type species. Currently, two species are accepted in this genus (Ranghoo et al. 2001; Hu et al. 2012a), and both species were collected from freshwater habitats in Asia (Ranghoo et al. 2001; Sudheep and Sridhar 2011; Hu et al. 2012a).

Aqualignicola hyalina Ranghoo et al.

Distribution: China, Hong Kong, New Territories, Tai Po, the Lam Tsuen River and Plover Cove Reservoir, on submerged wood (Ranghoo et al. 2001); India, Western Ghats, Kali River, submerged woody litter (Sudheep and Sridhar 2011).

Asexual morph: Undetermined

Notes: Holotype IFRD 8680. Sequence data is not available.

Aqualignicola vaginata Hu et al.

Distribution: China, Yunnan Province, Mengla, Wudaoban Stream, on submerged wood (Hu et al. 2012a).

Asexual morph: Undetermined

Notes: Holotype IFRDC 021–043. Sequence data is not available. Aqualignicola vaginata was introduced by Hu et al. (2012a) and it is so far only known from the type locality.


Ascitendus J. Campb. & Shearer, Mycologia 96(4): 829 (2004)

Asexual morph Undetermined. Sexual morph Description and illustrations see Campbell and Shearer (2004).

Type species: Ascitendus austriacus (Réblová et al.) J. Campb. & Shearer, Mycologia 96(4): 829 (2004)

Notes: Réblová and Winka (2001) introduced a new species, Ascolacicola austriaca from a stream in Australia. Campbell and Shearer (2004) examined 27 collections of Ascolacicola austriaca and found its morphology to be different from the type species of the genus. Based on the morphological differences between Ascolacicola austriaca and A. aquatica and phylogenetic analysis, a new genus Ascitendus was proposed for Ascolacicola austriaca (Campbell and Shearer 2004). Hyde et al. (2018) introduced the second species for this genus. Currently, two species of Ascitendus are accepted and both were collected from freshwater habitats (Réblová and Winka 2001; Campbell and Shearer 2004; Hyde et al. 2018).

Ascitendus aquaticus Dayarathne et al.

Distribution: Australia, North Queensland, Mulgrave River, on decaying wood submerged in a River (Hyde et al. 2018).

Asexual morph: Undetermined

Notes: Holotype MFLU 18–0143. LSU and SSU sequence data are available.


Ascitendus austriacus (Réblová et al.) J. Campb. & Shearer

Ascolacicola austriaca Réblová et al., Mycologia 93: 486 (2001)

Distribution: Australia, Wien 19, Herman-nskogel, on wood of Fagus sylvatica submerged in a stream (Réblová and Winka 2001), Canada, Ontario and Manitoba, on submerged wood; USA, on submerged wood; Venezuela, on submerged wood (Campbell and Shearer 2004).

Asexual morph: Undetermined

Notes: Holotype PRM 842991. LSU sequence data is available.


Ayria Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 248 (2004)

Asexual morph Undetermined. Sexual morph Description and illustrations see Fryar and Hyde (2004), Raja et al. (2009a).

Type species: Ayria appendiculata Fryar & K.D. Hyde, Cryptog. Mycol. 25(3): 248 (2004)

Notes: Fryar and Hyde (2004) established the genus Ayria with A. appendiculata as type species, the specimen was collected from rotting wood submerged in brackish and sea water, in Brunei. Raja et al. (2009a) introduced the second species A. nubispora in this genus.

Ayria appendiculata Fryar & K.D. Hyde

Distribution: Distribution: USA, Florida, on submerged decaying wood (Raja et al. 2009b).

Asexual morph: Undetermined

Notes: Holotype IFRD 8706. The type specimen was collected from brackish, sea water. Raja et al. (2009b) reported this species from freshwater habitats but without description, illustration and information for specimens. Sequence data is not available.

Ayria nubispora Raja, Ferrer & Shearer

Distribution: Costa Rica, Heredia, La Selva stream, on submerged decorticated woody debris and Limon, Las Palmas Stream, on submerged decorticated wood; USA, Florida, Marion County, Ocala National Forest, Fore Lake,on submerged decorticated woody debris (Raja et al. 2009a).

Asexual morph: Undetermined

Notes: Holotype ILL 40594. Sequence data is not available.

Cataractispora Hyde et al., Mycol. Res. 103(8): 1019 (1999)

Asexual morph Undetermined. Sexual morph Description and illustrations see Hyde (1992a) and Hyde et al. (1999).

Type species: Cataractispora aquatica Hyde et al., Mycol. Res. 103(8): 1020 (1999)

Notes: The genus Cataractispora was introduced by Hyde et al. (1999) with three new species and one new combination and all the species were collected from freshwater habitats. Ho et al. (2004) introduced one new species C. receptaculorum which was collected from freshwater habitats in Hong Kong, China. Five species are accepted in this genus and all species are reported from freshwater habitats around the world (Hyde 1992a; Hyde et al. 1999b; Ho et al. 2004; Raja et al. 2009b).

Cataractispora appendiculata Hyde et al.

Distribution: Australia, north Queensland, Cow Bay, freshwater stream, on submerged wood; Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Esu, on submerged wood; Seychelles, Mahe, River St Marie Louise, on submerged wood (Hyde et al. 1999b).

Asexual morph: Undetermined

Notes: Holotype IFRD 8724. Sequence data is not available.

Cataractispora aquatica Hyde et al.

Distribution: Australia, north Queensland, Cow Bay, freshwater stream, on submerged wood; Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Esu, on submerged wood; Seychelles, Mahe, River St Marie Louise, on submerged wood (Hyde et al. 1999b).

Asexual morph: Undetermined

Notes: Holotype IFRD 8725. Sequence data is not available.

Cataractispora bipolaris (K.D. Hyde) Hyde et al.

Annulatascus bipolaris K.D. Hyde, Aust. Syst. Bot. 5(1): 120 (1992)

Distribution: Australia, north Queensland, Clohesy River, on submerged wood (Hyde 1992a); USA, Florida, on submerged decaying wood (Raja et al. 2009b).

Asexual morph: Undetermined

Notes: Holotype BRIP 17374. Sequence data is not available.

Cataractispora receptaculorum Ho et al.

Distribution: China, Hong Kong, Plover Cove Reservoir, on bamboo submerged in freshwater (Ho et al. 2004)

Asexual morph: Undetermined

Notes: Holotype HKU (M) 5239 (now in IFRD). Sequence data is not available.

Cataractispora viscosa Hyde et al.

Distribution: Australia, north Queensland, Cow Bay, freshwater stream, on submerged wood; China, Hong Kong, New Territories, Tai Po Kau Country Park, on twigs submerged in a stream; Mauritius, Black River National Park, Black River (Hyde et al. 1999b).

Asexual morph: Undetermined

Notes: Holotype IFRD 8726. Sequence data is not available.


Chaetorostrum Zelski et al., Mycosphere 2(5): 594 (2011)

Asexual morphTaeniolella-like. Sexual morph Description and illustrations see Zelski et al. (2011a)

Type species: Chaetorostrum quincemilense Zelski et al., Mycosphere 2(5): 595 (2011)

Notes: The genus Chaetorostrum was erected by Zelski et al. (2011) to accommodate a holomorph species collected from freshwater stream in Peru. There is only one species in this genus and known only from the type locality.

Chaetorostrum quincemilense Zelski et al.

Distribution: Peru, Camanti, Stream at Quincemil Trail 1, on submerged woody debris (Zelski et al. 2011).

Asexual morph: Taeniolella-like, see Zelski et al. (2011)

Notes: Holotype ILL 40822. Sequence data is not available.


Longicollum Zelski et al., Mycosphere 2(5): 540 (2011)

Asexual morph: Undetermined. Sexual morph: Description and illustrations see Zelski et al. (2011).

Type species: Longicollum biappendiculatum Zelski et al., Mycosphere 2(5): 540 (2011)

Notes: Zelski et al. (2011) introduced the genus Longicollum with single species L. biappendiculatum, which was collected from both lentic and lotic freshwater habitats in America.

Longicollum biappendiculatum Zelski et al.

Distribution: Brazil, Bahia, stream at Serra da Jibóia, on submerged bark debris; Costa Rica, Heredia, La Selva Biological Station, La Selva stream, on submerged bark debris; Peru, Camanti, stream at Quincemil Trail 1, on submerged woody debris; USA, Florida, Wildcat Lake, Ocala National Forest, on submerged woody debris (Zelski et al. 2011).

Asexual morph: Undetermined

Notes: Holotype ILL 40794. Sequence data is not available.


Submersisphaeria K.D. Hyde, Nova Hedwigia 62(1–2): 172 (1996)

Asexual morph Undetermined. Sexual morph Description and illustrations see Hyde (1996) and Campbell et al. (2003a, b)

Type species: Submersisphaeria aquatica K.D. Hyde, Nova Hedwigia 62(1–2): 172 (1996)

Notes: The genus Submersisphaeria was introduced by Hyde (1996) with Submersisphaeria aquatica as type species. Presently, five species were accepted in this genus and only one species has been reported from freshwater habitats. Submersisphaeria aquatica was originally described from Queensland, Australia (Hyde 1996), and Campbell et al. (2003a, b) reported it from six states in the USA. Fournier et al. (2016) found S. aquatica on submerged wood of Pseudotsuga menziesii in France.

Submersisphaeria aquatica K.D. Hyde

Distribution: Australia, Queensland, on submerged wood (Hyde 1996); France, Nièvre: Arleuf, Préperny, on submerged wood of Pseudotsuga menziesii (Fournier et al. 2016); USA, Georgia, Louisiana, Mississippi, New Hampshire, North Carolina, Tennessee, Wisconsin, on submerged wood (Campbell et al. 2003a).

Asexual morph: Undetermined

Notes: Holotype BRIP 22743, other specimens collected from freshwater habitats: ILL 40186, ILL 40260. LSU sequence data is available.


Vertexicola Hyde et al., Mycologia 92(5): 1019 (2000)

Asexual morph Undetermined. Sexual morph Description and illustrations see Ranghoo et al. (2000) and Barbosa et al. (2013).

Type species: Vertexicola caudatus Hyde et al., Mycologia 92(5): 1020 (2000)

Notes: The genus Vertexicola is characterized by asci with a refractive apical ring and a tail-like pedicel and distoseptate ascospores with relatively thick walls and lacking appendages or a sheath (Ranghoo et al. 2000). There are two species accepted in this genus and both were collected from freshwater habitats.

Vertexicola ascoliberatus Shearer & F.R. Barbosa

Distribution: Costa Rica, Heredia, La Selva Biological Station, Salto 30, on submergd wood (Barbosa et al. 2013).

Asexual morph: Undetermined

Notes: Holotype ILL 40815. Sequence data is not available.

Vertexicola caudatus Hyde et al.

Distribution: China, Hong Kong, Tai Po, Plover Cove Reservoir, submerged wood (Ranghoo et al. 2000).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 3108 (now in IFRD). Sequence data is not available.


Atractosporales Zhang et al.

Atractosporaceae Zhang et al.

Atractospora Réblová & J. Fourn., Mycol Prog 15: 8 (2016)

Asexual morph Undetermined. Sexual morph Description and illustrations see Réblová et al. (2016a).

Type species: Atractospora reticulata Réblová & J. Fourn., Mycol Prog 15(21): 10 (2016)

Notes: The genus Atractospora was described for perithecial ascomycetes occurring on decaying wood submerged in freshwater habitats (Réblová et al. 2016b). Five species are accepted in this genus, of which Atractospora ellipsoidea and A. thailandensis are reported from Asia and USA (Ho et al. 1999c; Campbell and Shearer 2004; Fryar et al. 2004), while other three species were collected from Europe (Réblová et al. 2016b).

Atractospora aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF555639, Facesoffungi number: FoF 05411, Fig. 2

Fig. 2
figure 2

Atractospora aquatica (MFLU 18–2322, holotype) a Ascomata on decaying wood. b Section of the ascoma. c Structure of peridium. d Paraphyses. eh Asci. il Ascospores. m Germinating ascospore. Scale bars: b 100 μm, c, d 25 μm, ef 50 μm, im 10 μm

Etymology: Referring to the aquatic habitat of this fungus.

Holotype: MFLU 18–2322

Saprobic on decaying wood submerged in freshwater habitats. Asexual morph Undetermined. Sexual morphAscomata 157–251 μm high, 154–234 μm diam., dark brown to black, solitary, semi-immersed to superficial, globose to subglobose, unilocular. Ostiole periphysate. Peridium 20–34 μm thick, consisting two-layers, Outer layer consisting of thick-walled, mid brown cells of textura prismatica, towards the interior grading into several layers of thin-walled subhyaline, flattened cells. Paraphyses hyaline, unbranched, cylindrical. Asci 110–168 × 9–13 µm (\( {\bar{x}} \) = 139 × 11 µm, n = 20), 8-spored, unitunicate, cylindrical, with a slender, tapering stipe, apex broadly rounded, with a conspicuous, non-amyloid apical ring. Ascospores 15–19 × 7–9 µm (\( {\bar{x}} \) = 17 × 8 µm, n = 20), uniseriate or obliquely uniseriate, fusiform, guttulate, aseptate, hyaline, smooth-walled.

Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2017, H.W. Shen, S-1398 (MFLU 18–2322, holotype).

Notes: Atractospora aquatica resembles A. thailandensis in having globose to subglobose, unilocular, dark brown to black ascomata, hyaline, unbranched paraphyses, cylindrical asci with a slender, tapering stipe and conspicuous, non-amyloid, apical ring and fusiform, hyaline, aseptate, guttulate ascospores (Zhang et al. 2017a, b). However, Atractospora aquatica differs from A. thailandensis in having shorter asci (110–168 vs. 175–215 μm) and smaller ascospores (15–19 vs. 20–26 μm) with smaller guttules. Atractospora aquatica also resembles A. decumbens and A. reticulata in having solitary, dark brown ascomata, unitunicate, 8-spored, cylindrical asci and fusiform, hyaline ascospores uniseriate or obliquely uniseriate in the ascus (Réblová et al. 2016b). However, Atractospora aquatica differs from A. decumbens and A. reticulata in having shorter asci and aseptate ascospores, while A. decumbens and A. reticulata have 3-septate ascospores. Phylogenetic results show that Atractospora aquatica is distinct from other species of Atractospora (Fig. 1, clade 3).

Atractospora decumbens Réblová & J. Fourn

Distribution: France, Midi-Pyrénées: Ariège, Rimont, L’ Estanque brook, on submerged wood of Sambucus nigra in a stream (Réblová et al. 2016b).

Asexual morph: Undetermined

Notes: Holotype PRM 934676. ITS, LSU, SSU and RPB2 sequence data are available.

Atractospora ellipsoidea (Ho et al.) Réblová & J. Fourn

Aquaticola ellipsoidea Ho et al., Fungal Divers 3: 90 (1999)

Distribution: Brunei, Tutong River, on submerged wood (Fryar et al. 2004); China, Hong Kong, Tai Po, Tai Po Kau Forest Stream, on submerged wood of Machilus velutina (Ho et al. 1999c); USA, North Carolina (Campbell and Shearer 2004).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 6033 (now in IFRD). LSU sequence data is available.

Atractospora reticulata Réblová & J. Fourn

Distribution: France, Midi-Pyrénées: Ariège, Rimont, Peyrau brook, on submerged wood of Sambucus nigra in a stream (Réblová et al. 2016b).

Asexual morph: Undetermined

Notes: Holotype PRM 934677. ITS, LSU and RPB2 sequence data are available.

Atractospora thailandensis Dong et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged wood in a small River (Zhang et al. 2017).

Asexual morph: Undetermined

Notes: Holotype HKAS 96226. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Atractospora verruculosa Réblová & J. Fourn

Distribution: France, Pyrénées-Atlantiques, Lescun, Borde Bayé, Lescun stream, on submerged wood of Alnus glutinosa (Réblová et al. 2016b).

Asexual morph: Undetermined

Notes: Holotype PRM 934679. ITS, LSU, SSU and RPB2 sequence data are available.


Rubellisphaeria Réblová & J. Fourn., Mycol Prog 15: 13 (2016)

Asexual morph Undetermined. Sexual morph Description and illustrations see Réblová et al. (2016a).

Type species: Rubellisphaeria abscondita Réblová & J.Fourn., Mycol Prog 15: 13 (2016)

Notes: Réblová et al. (2016a) introduced the genus Rubellisphaeria with single species Rubellisphaeria abscondita, collected from freshwater in France. This is a monotypic genus and known only from the type locality.

Rubellisphaeria abscondita Réblová & J. Fourn

Distribution: France, Loire: Saint Jean la Vêtre, Les Yverneaux, on submerged twigs of Abies alba in a peat bog (Réblová et al. 2016b).

Asexual morph: Undetermined

Notes: Holotype PRM 934681. ITS, LSU, SSU and RPB2 sequence data are available.


Jobellisiales M.J. D’souza & K.D. Hyde

Jobellisiaceae Réblová

Jobellisia M.E. Barr, Mycotaxon 46: 60 (1993)

Asexual morph Undetermined. Sexual morphAscomata superficial or basally immersed, nonstromatic, globose to subglobose to obpyriform to lageniform, brown or externally with yellowish pigments, glabrous or slightly rugose, with short to long papilla or with long upright neck. Peridium comprising two or three layers. Paraphyses numerous, septate, hyaline. Asci 8-spored, unitunicate, cylindrical-clavate, with nonamyloid apex and distinct refractive apical annulus, with basally swollen stipe, stipe remnants attached to the ascogenous hyphae after dehiscence. Ascospores ellipsoidal to reniform to navicular, aseptate or transversely 1-septate with one or two polar germ pores, brown.

Type species: Jobellisia luteola (Ellis & Everh.) M.E. Barr, Mycotaxon 46: 61 (1993)

Notes: The genus Jobellisia currently comprises nine species and they are found in tropical and temperate zones of the northern hemisphere (Réblová 2008; Maharachchikumbura et al. 2016). Three species have been found in freshwater habitats.

Jobellisia guangdongensis F. Liu & L. Cai

Distribution: China, Guangdong Province, on submerged decaying wood (Liu et al. 2011).

Asexual morph: Undetermined

Notes: Holotype HMAS 251240. ITS and LSU sequence data are available.

Jobellisia luteola (Ellis & Everh.) M.E. Barr

Letendraea luteola Ellis & Everh., Proc. Acad. nat. Sci. Philad. 47: 415 (1895)

Distribution: USA, on submerged decaying wood (Raja et al. 2009b).

Asexual morph: Undetermined

Notes: Holotype Morgan 1109. LSU sequence data is available. Jobellisia luteola was originally collected from decayed wood on terrestrial habitats. Raja et al. (2009b) reported this species from freshwater habitats during their studies on latitudinal, habitat and substrate distribution patterns of freshwater ascomycetes in Florida, USA.

Jobellisia viridifusca K.M. Tsui & K.D. Hyde

Distribution: China, Hong Kong, Lantau Island, Tung Chung River, on submerged decaying wood (Ranghoo et al. 2001)

Asexual morph: Undetermined

Notes: Holotype HKU (M) 80455 (now in IFRD). Sequence data is unavailable.


Diaporthales Nannf.

Diaporthaceae Hohn. ex Wehm.

Diaporthe Nitschke, Pyrenomycetes Germanici 2: 240 (1870)

Asexual morph Coelomycetous, pycnidial, ostiolate, brown to black, scattered or aggregated, globose to subglobose. Conidiophores cylindrical, sometimes filiform, aseptate or septate, cylindrical, sometimes branched. Conidiogenous cells phialidic, cylindrical, terminal. Conidia dimorphic, hyaline, smooth, with usually fusiform and biguttulate alpha conidia and usually filiform, hamate, non-guttulate beta conidia. Sexual morphAscomata globose to subglobose, coriaceous, immersed to semi-immersed, single to clustered, brown to black. Neck cylindrical, black. Peridium comprising compressed cells of textura angularis. Paraphyses cylindrical, longer than asci, septate. Asci 8-spored, unitunicate, thin-walled, apedicellate, broad cylindrical to obclavate, with a minute apical ring. Ascospores overlapping biseriate, ellipsoidal to fusiform, septate, hyaline, thin-walled, smooth-walled.

Type species: Diaporthe eres Nitschke, Pyrenomyc. Germ. 2: 245 (1870)

Notes: Species of Diaporthe have broad host ranges and are widely distributed (Udayanga et al. 2012). Currently, MycoBank (accessed December 2018) lists more than 900 names in the genus Diaporthe, but this was reduced to 171 species (Dissanayake et al. 2017). Hu et al. (2012c) introduced a new Diaporthe species D. aquatica, collected from submerged wood in stream in Yunnan Province, China.

Diaporthe aquatica Hu et al.

Distribution: China, Guizhou Province, Guiyang city, on submerged wood in a small ditch (Hu et al. 2012c)

Asexual morph: undetermined

Notes: Holotype IFRD 021–018; ITS sequence data are available, ex-type strain IFRDCC 3051. D. aquatica is the only Diaporthe species restricted to freshwater habitats (Hu et al. 2012c).


Gnomoniaceae G. Winter

Ambarignomonia Sogonov, Stud. Mycol. 62: 35 (2008)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Senanayake et al. (2018).

Type species: Ambarignomonia petiolorum (Schwein.) Sogonov, Stud Mycol 62: 36 (2008)

Notes: The genus Ambarignomonia was introduced by Sogonov et al. (2008) to accommodate Sphaeria petiolorum. Presently, there is only one species accepted in this genus. Sogonov et al. (2008) mentioned the monotypic Ambarignomonia with A. petiolorum and is restricted to one plant host, Liquidambar styraciflua, whereas other genera of the Gnomoniaceae do not show such consistency in host associations.

Ambarignomonia petiolorum (Schwein.: Fr.) Sogonov

Sphaeria petiolorum Schwein.: Fr., Schr. Naturf. Ges. Leipzig 1: 41. 1822: Syst. Mycol. 2: 517 (1823)

Gnomonia petiolorum (Schwein.: Fr.) Cooke, Grevillea 7: 54 (1878)

Gnomoniella amoena var. petiolorum (Schwein.: Fr.) Sacc., Syll. Fung. 1: 414 (1882)

Distribution: USA, Wisconsin, Trout lake/Big Muskellunge lake, on submerged partially decomposed Acer rubrum leaves (Fallah and Shearer 2001).

Asexual morph: Undetermined

Notes: Lectotype BPI 800519, Epitype BPI 844274, specimens collected from freshwater habitats: ILLS 54015, ILLS 54016. ITS, LSU, RPB2, TEF1α and β-tubulin sequence data are available. Fallah and Shearer (2001) collected two specimens (ILLS 54015 and ILLS 54016) from freshwater habitats and provided descriptions and illustrations and identified the collections as Gnomonia petiolorum. Sogonov et al. (2008) combined this species as Ambarignomonia petiolorum.


Gnomonia Ces. & De Not., Comm. Soc. crittog. Ital. 1(4): 231 (1863)

Asexual morph see Sivanesan and Shaw (1977). Sexual morph Description see Maharachchikumbura et al. (2016).

Type species: Gnomonia vulgaris Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 232 (1863)

Notes: The genus Gnomonia was introduced by Cesati and De Notaris (1863) and typified by Gnomonia gnomon. Gnomonia comprises 273 species and only one species has been reported from freshwater habitats (Sivanesan and Shaw 1977; Fallah and Shearer 2001; Senanayake et al. 2018).

Gnomonia papuana Sivan. & D.E. Shaw

Distribution: Papua New Guinea, on submerged leaves (Sivanesan and Shaw 1977).

Asexual morph: Sesquicillium-like, see Sivanesan and Shaw (1977)

Notes: Holotype IMI 197503. Sequence data is not available.


Gnomoniella Sacc., Michelia 2 (7): 312 (1881)

Asexual morph Undetermined. Sexual morphAscomata globose to subglobose, immersed. Paraphyses deliquescence. Asci cylindrical, subsessiles. Ascospores fusiform, ellipse, hyaline, septate.

Type species: Gnomoniella tubaeformis (Tode) Sacc., Syll. fung. (Abellini) 1: 413 (1882)

Notes: Kirk et al. (2008) accepted 13 species in the genus Gnomoniella. Two species have been found in freshwater habitats and both were collected from Asia (Ho et al. 2001).

Gnomoniella microspora M. Monod

Distribution: Malaysia, on submerged wood (Ho et al. 2001).

Asexual morph: Undetermined

Notes: ITS sequence data are available. Gnomoniella microspora was originally collected from terrestrial habitats (Monod 1983). Ho et al. (2001) reported Gnomoniella microspora from freshwater habitats and we consider this species as freshwater fungus.

Gnomoniella rubicola Pass.

Distribution: Brunei, on submerged wood (Ho et al. 2001).

Asexual morph: Undetermined

Notes: Sequence data is not available. Same as Gnomoniella microspora, the original collection of G. rubicola was not collected from freshwater habitats. However, we consider this species as freshwater fungus as Ho et al. (2001) found it on submerged wood in Brunei.


Diaporthales genera incertae sedis

Phruensis Pinruan, Mycologia 96 (5): 1165 (2004)

Asexual morph Hyphomycetous. Colonies on PDA effuse, brown mycelium party immersed, nonstromatic. Conidiophores semimacronematous, mononematous, branched, straight or slightly flexuous, pale brown to brown, smooth. Conidiogenous cells monophialidic, determinate, with small collarettes. Conidia straight or curved, oblong, hyaline, smooth. Sexual morphAscomata immersed, subglobose, black, coriaceous, ostiolate, with long central cylindrical neck. Peridium composed of 2 layers, with outer layer parenchymatous, intensely brown and merging with the host cells, with inner layer, cells elongate and hyaline. Paraphyses hyaline, broad, septate. Asci unitunicate, cylindrical to fusiform, apedicellate, apically rounded, J-, subapical ring. Ascospores cylindrical, straight or curved, versicolorous, transseptate, brown with hyaline or pale brown end cells.

Type species: Phruensis brunneispora Pinruan, Mycologia 96(5): 1165 (2004)

Notes: Pinruan et al. (2004b) introduced the genus Phruensis with a single species P. brunneispora, which was collected from a submerged palm in Sirindhorn Peat Swamp Forest in Thailand. No more species reported for this genus since it was introduced.

Phruensis brunneispora Pinruan

Distribution: Thailand, Narathiwat Province, Sirindhorn peat swamp forest, on submerged palm in freshwater stream (Pinruan et al. 2004b).

Asexual morph: Phialophora-like, see Pinruan et al. (2004b)

Notes: Holotype BBH, Pinruan Wah 113.1. SSU sequence data is available.

Distoseptisporales Z.L. Luo, H.Y. Su & K.D. Hyde, ord. nov

Index Fungorum number: IF 555640, Facesoffungi number: FoF 05412

Asexual morphMycelium mostly immersed, composed of branched, septate, smooth, pale brown hyphae. Conidiophores macronematous, mononematous, septate, unbranched, erect, straight or flexuous, smooth. Conidiogenous cells monoblastic, integrated, determinate, terminal, cylindrical. Conidia acrogenous, solitary, euseptate or distoseptate, obclavate or cylindrical with rounded apex. Conidial secession schizolytic. Sexual morph Undetermined.

Type family: Distoseptisporaceae K.D. Hyde & McKenzie, Fungal Divers 80:402 (2016)

Notes: Distoseptisporaceae was established by Su et al. (2016) with a single genus Distoseptispora based on morphology and phylogeny. Phylogenetic analysis based on combined LSU, SSU, RPB2 and TEF1α sequence data here shows that species of Distoseptispora and Aquapteridospora cluster together with strong support and form a distinct clade within subclass Diaporthomycetidae. We therefore raise Distoseptisporaceae to order Distoseptisporales.


Distoseptisporaceae K.D. Hyde & McKenzie

Distoseptispora Hyde et al., Fungal Divers 80: 402 (2016)

Asexual morph Description and illustration see Su et al. (2016) and Yang et al. (2018). Sexual morph Undetermined.

Type species: Distoseptispora aquatica Luo et al., Fungal Divers 80: 402 (2016)

Notes: Su et al. (2016) introduced the genus Distoseptispora to accommodate twoSporidesmium-like species. Yang et al. (2017) emended the description of the genus Distoseptispora. Currently, there are 13 species in Distoseptispora with ten species collected from freshwater habitats (Su et al. 2016; Hyde et al. 2016b; Luo et al. 2018a; Yang et al. 2018a).

Distoseptispora appendiculata D.F. Bao, Z.L. Luo & H.Y. Su, sp. nov.

Index Fungorum number: IF 556690, Facesoffungi number: FoF 06302, Fig. 3

Fig. 3
figure 3

Distoseptispora appendiculata (DLU B95, holotype) a Colonies on wood. be Conidiophores with conidia. f Conidiogenous cells with conidia. g Conidiogenous cells. hk Conidia. l Germinating conidium. Scale bars: bf 30 μm, gl 20 μm

Fig. 4
figure 4

Distoseptispora guttulata (B-43) a Colonies on wood. bd Conidiophores with conidia. eg Conidia. h Germinating conidium. Culture on PDA from above (i) and reverse (j). Scale bars: bh 30 μm

Fig. 5
figure 5

Distoseptispora lignicola (MFLU 18–1458, holotype). a, b Colonies on wood. cd Conidiophores with conidia. ef Conidiogenous cells and conidia. gj Conidia. k Germinating conidium. Culture on PDA from surface (l) and reverse (m). Scale bars: d 30 μm, c, ek 20 μm

Fig. 6
figure 6

Distoseptispora multiseptata (B-37) a, b Colonies on wood. cf Conidiophores with conidia. gk Conidia. l Conidiophores with conidiogenous cells. m Germinating conidium. Culture on PDA from above (n) and reverse (o). Scale bars: ck, m 30 μm, l 10 μm

Fig. 7
figure 7

Distoseptispora neorostrata (DLU B103, holotype) a, b Colonies on wood. c, d Conidiophores with conidia. e, f Conidiogenous cells with conidia. gj Conidia. k Germinating conidium. Culture on MEA from above (l) and reverse (m). Scale bars: ck 30 μm

Fig. 8
figure 8

Distoseptispora obclavata (DLU B71, holotype) a, b Colonies on wood. cg Conidiophores with conidia. h, i Conidiogenous cells with conidia. jm Conidia. n Germinating conidium. Culture on PDA from above (o) and reverse (p). Scale bars: ch 30 μm, in 20 μm

Fig. 9
figure 9

Aquapteridospora fusiformis (MFLU 18–1601, holotype). a Colonies on substrate. bd Conidiophore with conidia. e, f Conidiogenous cells with conidia gj Conidia, k Germinating conidium. Culture on PDA from surface (l) and reverse (m). Scale bars: bd 50 μm, e, f 20 μm, gk 10 μm

Etymology: Referring to its gelatinous conidia appendage

Holotype: DLU B95

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, olivaceous or mid-brown, hairy, velvety. Mycelium mostly immersed, consisting of branched, septate, smooth, subhyaline to pale brown hyphae. Conidiophores 62–86 μm long, 4.5–5.5 μm wide (\( {\bar{x}} \) = 74 × 5 μm, n = 10), macronematous, mononematous, solitary, erect, straight or flexuous, olivaceous or brown, 5–6-septate, smooth. Conidiogenous cells monoblastic, holoblastic, terminal, dark brown. Conidia 67–89 μm long, 10–16 μm wide (\( {\bar{x}} \) = 78 × 13 μm, n = 20), acrogenous, solitary, obpyriform or obclavate, 13–17-distoseptate, thick-walled, olivaceous or dark brown below, hyaline towards apex, truncate at base, slender and rounded at apex, smooth, with a conspicuous, gelatinous, hyaline sheath around tip. Sexual morph Undetermined.

Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon, saprobic on decaying wood submerged in a freshwater stream, 1st October 2017, Z.L. Luo, B-95 (DLU B95, holotype), ex-type living culture MFLUCC 18–0259.

Notes: Distoseptispora appendiculata resembles D. obpyriformis in having effuse, olivaceous or mid-brown, hairy, velvety colonies, macronematous, straight or flexuous, olivaceous or brown, septate conidiophores and obpyriform, distoseptate, olivaceous or dark brown conidia (Luo et al. 2018a). However, Distoseptispora appendiculata is easily distinguished from D. obpyriformis and other species in Distoseptispora by its conspicuous, gelatinous, hyaline appendage sheath. Phylogenetic results show that Distoseptispora appendiculata is distinct from other species of Distoseptispora (Fig. 10).

Fig. 10
figure 10

Phylogram generated from maximum likelihood analysis based on ITS, LSU, RPB2 and TEF1α sequence data for species of Distoseptisporales (with Sordaria fimicola as outgroup). The best scoring RAxML tree with a final likelihood value of − 30852.86243 is presented. RAxML bootstrap support values equal to or greater than 75% are given before the forward slash (black). Bayesian posterior probability equal to or higher than 0.95 are given after the forward slash (red). Hyphen (‘-’) indicates a value lower than 75% for RAxML and Bayesian posterior probability lower than 0.95. Newly generated sequences are in red. Ex-type or ex-epitype strains are in bold

Fig. 11
figure 11

Ceratosphaeria aquatica (MFLU 18–2323, holotype) a Appearance of necks on substrate. b Section through ascomata. c, d Section through neck. e Structure of peridium. f Paraphyses. gj Asci. k Apical ring. lo Ascospores. Scale bars: b, c 100 μm, d 50 μm, ej, lo 20 μm, k 10 μm

Fig. 12
figure 12

Ceratosphaeria lignicola (MFLU 18–1457, holotype). a, b Appearance of neck on substrate. c, d Section of ascoma. e Structure of peridium. f Paraphyses. gk Asci. ln Ascospores. o Germinating ascospore. culture on PDA from surface (p) and reverse (q). Scale bars: c 100 μm, d, g 50 μm, ho 30 μm, e, f 20 μm

Fig. 13
figure 13

Aquafiliformis lignicola (MFLU 18–2325, holotype) a Appearance of necks on substrate. b Section through ascoma. c Structure of peridium. d Paraphyses. e, f Asci. gj Ascospores. Culture on PDA from surface (k) and reverse (l). Scale bars: b 150 μm, c 50 μm, df 30 μm, gj 20 μm

Fig. 14
figure 14

Ophioceras submersum (MFLU 18–1459, holotype). a Appearance of neck on substrate. bc Section of ascoma. d Structure of peridium. e Paraphyses. fh Asci. il Ascospores. Scale bars: b 150 μm, c 50 μm, dl 20 μm

Distoseptispora aquatica Luo et al.

Distribution: China, Yunnan Province, Dali, Cangshan Mountain, on submerged wood in stream (Su et al. 2016).

Sexual morph: Undetermined

Notes: Holotype HKAS 83991. ITS and LSU sequence data are available.

Distoseptispora cangshanensis Luo et al.

Distribution: China, Yunnan Province, Dali, Cangshan Mountain, on submerged wood (Luo et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 18–0474. ITS, LSU and TEF1α sequence data are available.

Distoseptispora fluminicola McKenzie et al.

Distribution: China, Yunnan Province, Dali, on submerged wood (Su et al. 2016).

Sexual morph: Undetermined

Notes: Holotype HKAS 84006. ITS, LSU and TEF1α sequence data are available.

Distoseptispora guttulata J. Yang & K.D. Hyde

Facesoffungi number: FoF 03357, Fig. 4

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, brown to black, hairy or velvety. Mycelium partly superficial, partly immersed, consisting of branched, septate, smooth, subhyaline to pale brown hyphae. Conidiophores 28–84 μm long, 4–5 μm wide (\( {\bar{x}} \) = 56 × 4.5 μm, n = 20), macronematous, mononematous, mid or dark brown, 3–4(–7)-septate, solitary or caespitose, smooth, straight or slightly flexuous, cylindrical, rounded at the apex. Conidiogenous cells monoblastic, integrated, terminal, determinate, mid to dark brown, cylindrical, sometimes proliferating percurrently. Conidia 70–190(–520) μm long, 8.5–10.5 μm wide (\( {\bar{x}} \) = 130 × 9.5 μm, n = 20), acrogenous, solitary, holoblastic, obclavate or lanceolate, rostrate, curved, 9–14(–27)-euseptate, mid to dark brown, or olivaceous, smooth, truncate at the base, tapering to the apex. Sexual morph Undetermined.

Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon, saprobic on decaying wood submerged in a freshwater stream, 1st October 2017, Z.L. Luo, B-43, living culture DLUCC B43.

Distribution: Thailand, Prachuap Khiri Khan Province, on decaying wood submerged in a freshwater stream (Yang et al. 2018a)

Sexual morph: Undetermined

Notes: Holotype MFLU 17–0852, isotype GZAAS 17–0005. ITS, LSU, SSU and TEF1α sequence data are available. Distoseptispora guttulata was introduced by Yang et al. (2018a) based on a collection obtained from a freshwater stream in Thailand. Morphologically, our isolate fits well with the characters of D. guttulata (Yang et al. 2018a). Phylogenetic analysis also shows that our isolate clusters with ex-type of D. guttulata with good support (Fig. 10).

Distoseptispora lignicola D.F. Bao, Z.L. Luo, H.Y. Su & K.D. Hyde, sp. nov.

Index Fungorum number: IF 555641, Facesoffungi number: FoF 05413, Fig. 5

Etymology: Referring to this taxon dwelling on wood

Holotype: MFLU 18–1458

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, scattered, hairy, pale brown to brown. Mycelium mostly immersed, composed of branched, septate, brown, smooth hyphae. Conidiophores 84–124 μm long, 4–5 μm wide (\( {\bar{x}} \) = 104 × 4.5 μm, n = 20), macronematous, mononematous, solitary or in groups, erect, straight or slightly flexuous, 6–10-septate, unbranched, cylindrical, brown, smooth. Conidiogenous cells monoblastic, integrated, terminal, determinate, brown, cylindrical. Conidia 60–108 μm long, 7–9 μm wide (\( {\bar{x}} \) = 84 × 8 μm, n = 20), acrogenous, solitary or catenate, obclavate, truncate at base, tapering towards the apex, straight or slightly curved, 5–9-euseptate, slightly constricted at septa, guttulate, brown, smooth. Sexual morph Undetermined.

Material examined: THAILAND, SaiKhu Waterfall, on submerged decaying wood, 28 August 2017, C.G. Lin, B-2 (MFLU 18–1458, holotype), ex-type living culture MFLUCC 18–0198.

Notes: Distoseptispora lignicola resembles D. guttulata in having macronematous, mononematous, unbranched, cylindrical, septate conidiophores, solitary or in groups on natural substrata, monoblastic, integrated, terminal, determinate conidiogenous cells and acrogenous, obclavate, rostrate, euseptate, guttulate conidia (Yang et al. 2018a). However D. lignicola differs from D. guttulata in having longer conidiophores (84–124 vs. 55–90 (–145) μm) and 5–9-euseptate conidia, while D. guttulata has 11–14(–20)-euseptate conidia. Phylogenetically, Distoseptispora guttulata clusters in Distoseptispora and sister to D. leonensis (HKUCC 10822), but is distinct from other Distoseptispora species (Fig. 1, clade 15).

Distoseptispora multiseptata J. Yang & K.D. Hyde

Facesoffungi number: FoF 02244, Fig. 6

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, dark olive-green, hairy or velvety. Mycelium mostly immersed, comprised of branched, septate, smooth, hyaline to pale brown hyphae. Conidiophores 29–47 μm long, 4–6 μm wide (\( {\bar{x}} \) = 38 × 5 μm, n = 20), macronematous, mononematous, solitary, brown, 2–3-septate, straight or slightly flexuous, erect, slightly tapering distally, truncate at the apex, olive-green to dark brown. Conidiogenous cells holoblastic, monoblastic, integrated, terminal, brown, determinate, cylindrical. Conidia 147–185 μm long, 12–14 μm wide (\( {\bar{x}} \) = 16 × 13 μm, n = 20), acrogenous, solitary, obclavate, rostrate, multi-distoseptate, tapering towards the apex, dark olivaceous green. Conidial secession schizolytic. Sexual morph Undetermined.

Material examined: THAILAND, Sai khu waterfall, Prachuap khiri Khan., saprobic on decaying wood submerged in a freshwater stream, 1st September 2017, Vinit Kumar, B-37, living culture MFLUCC 18–0215.

Distribution: Thailand, Prachuap Khiri Khan Province, Hua Hin, on submerged wood in a stream (Hyde et al. 2016b).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–1144, isotype HKAS 95045. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available. Distoseptispora multiseptata was introduced by Yang et al. (2018a) based on a collection obtained from a freshwater stream in Thailand. Morphologically, our isolate fits well with the characters of D. multiseptata (Yang et al. 2018a). Phylogenetic analysis also shows that our isolate clusters with ex-type of D. multiseptata (Fig. 10).


Distoseptispora neorostrata D.F. Bao, Z.L. Luo & H.Y. Su, sp. nov.

Index Fungorum number: IF 556691, Facesoffungi number: FoF 06135, Fig. 7

Etymology: Referring to its morphological similarity to D. rostrata

Holotype: DLU B103

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, dark olivaceous, hairy. Mycelium partly superficial, partly immersed in the substrate, comprised of branched, septate, hyaline to pale brown hyphae. Conidiophores 93–117 μm long, 5.5–6.5 μm wide (\( {\bar{x}} \) = 105 × 6 μm, n = 10), macronematous, mononematous, solitary, brown, 5–7-septate, straight or flexuous, tapering distally, truncate at the apex. Conidiogenous cells monoblastic, integrated, terminal, brown, determinate. Conidia 109–147 μm long, 13–15 μm wide (\( {\bar{x}} \) = 128 × 14 μm, n = 20), acrogenous, solitary, elongate, obclavate, rostrate, multi-distoseptate, tapering towards the rounded apex, dark olivaceous to mid or dark brown, pale brown towards apex, truncate at the base, guttulate, smooth-walled. Sexual morph Undetermined.

Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon, saprobic on decaying wood submerged in a freshwater stream, 1st October 2017, Z.L. Luo, B-103 (DLU B103, holotype), ex-type living culture MFLUCC 18–0376.

Notes: Distoseptispora neorostrata shares similar morphological characters with D. rostrata in the shape, colour and size of its conidiophores and conidia (Luo et al. 2018a). However, the multi-gene phylogenetic analyses showed that they are different species (Fig. 10).

Distoseptispora obclavata D.F. Bao, Z.L. Luo & H.Y. Su, sp. nov.

Index Fungorum number: IF 556689, Facesoffungi number: FoF 06296, Fig. 8

Etymology: Referring to its obclavate conidia

Holotype: DLU B71

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, olivaceous or dark brown, hairy, velvety. Mycelium mostly immersed, consisting of branched, septate, smooth, subhyaline to pale brown hyphae. Conidiophores 117.5–162.5 μm long, 5–7 μm wide (\( {\bar{x}} \) = 140 × 6 μm, n = 20), macronematous, mononematous, pale to dark brown, solitary, 5–10-septate, erect, straight or slightly flexuous, unbranched, smooth, cylindrical, rounded at the apex. Conidiogenous cells monoblastic, integrated, terminal, determinate, pale to dark brown, cylindrical. Conidia 46–66 μm long, 9–11 μm wide (\( {\bar{x}} \) = 56 × 10 μm, n = 25), acrogenous, solitary, obclavate, 9–11-distoseptate, thick-walled, olivaceous to pale or dark brown, apering towards the rounded apex, slightly curved, truncate at the base, guttulate, smooth-walled. Sexual morph Undetermined.

Material examined: THAILAND, Khwaeng Phra Khanong Nuea, Khet Watthana Krung Thep Maha Nakhon, saprobic on decaying wood submerged in a freshwater stream, 1st October 2017, Z.L. Luo, B-71 (DLU B71, holotype), ex-type living culture MFLUCC 18–0329.

Notes: Distoseptispora obclavata resembles D. appendiculata in having effuse, olivaceous or mid-brown, hairy, velvety colonies, macronematous, straight or flexuous, septate conidiophores and obclavate, distoseptate, olivaceous or dark brown conidia. However, Distoseptispora obclavata is easily distinguished from D. appendiculata by its shorter conidia (46–66 vs. 67–89 μm) without appendage sheath. Phylogenetic results show that Distoseptispora appendiculata is distinct from other species of Distoseptispora (Fig. 10).

Distoseptispora obpyriformis Z.L. Luo & H.Y. Su

Distribution: China, Yunnan Province, Nujiang River, on submerged wood (Luo et al. 2018).

Sexual morph: Undetermined

Notes: Holotype MFLU 18–0476, paratype MFLU 18–0477. ITS, LSU, RPB2 and TEF1α sequence data are available.

Distoseptispora phangngaensis Yang et al.

Distribution: Thailand, Phang Nga Province, Bann Tom Thong Khang, on decaying wood submerged in a freshwater stream (Yang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–0855, isotype GZAAS 17–0008. ITS, LSU and TEF1α sequence data are available.

Distoseptispora rostrata Luo et al.

Distribution: China, Yunnan Province, Nujiang River, on submerged wood (Luo et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 18–0479, paratype MFLU 18–0475. ITS, LSU, RPB2 and TEF1α sequence data are available.

Distoseptispora submersa Luo et al.

Distribution: China, Yunnan Province, Nujiang River, on submerged wood (Luo et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 18–0478, isotype HKAS 92806. ITS, LSU, RPB2 and TEF1α sequence data are available.

Distoseptispora suoluoensis Yang et al.

Distribution: China, Guizhou Province, Anshun city, Gaodang village, on decaying wood submerged in Suoluo River (Yang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–0853, isotype GZAAS 17–0006, paratype MFLU 17–0854. ITS, LSU and TEF1α sequence data are available.


Distoseptisporales genera incertae sedis

Aquapteridospora Yang et al., Cryptog. Mycol. 36(4): 474 (2015)

Asexual morph Descriptions and illustration see Yang et al. (2015). Sexual morph Undetermined.

Type species: Aquapteridospora lignicola J. Yang, K.D. Hyde & Maharachch, Cryptog Mycol 36: 474 (2015)

Notes: Yang et al. (2015) introduced the genus Aquapteridospora with single asexual species, A lignicola, which was collected from freshwater stream in northern Thailand. In this study, we introduce the second species collected from freshwater. Aquapteridospora was placed as Diaporthomycetidae genera incertae sedis by Yang et al. (2015). In our phylogenetic analysis, Aquapteridospora species form a distinct clade within Distoseptisporales and basal to Distoseptisporaceae, and we therefore treat this genus as Distoseptisporales genera incertae sedis, and its familial placement needs further studies.

Aquapteridospora lignicola Yang et al.

Distribution: Thailand, Chiang Rai Province, stream flowing in Tham Luang Nang Non Cave, on submerged wood (Yang et al. 2015).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–1172. LSU sequence data is available.

Aquapteridospora fusiformis Z.L. Luo, D.F. Bao, H.Y. Su & K.D. Hyde, sp. nov.

Index Fungorum number: IF 555642, Facesoffungi number: FoF 05414, Fig. 9

Etymology: Referring to the fusiform conidia of this fungus.

Holotype: MFLU 18–1601

Saprobic on decaying wood submerged in freshwater. Asexual morphColonies on the natural substrate effuse, hairy, pale brown to brown. Mycelium superficial or partly immersed, composed of branched, septate, pale brown to brown, smooth, thin-walled hyphae. Conidiophores (88–)134–188 μm long, 5–7 μm wide (\( {\bar{x}} \) = 161 × 6 μm, n = 20), macronematous, mononematous, solitary, erect, straight or slightly flexuous, unbranched, cylindrical, septate, smooth, thick-walled, brown at the base, paler towards apex. Conidiogenous cells polyblastic, terminal, later becoming intercalary, pale brown, integrated, with several sympodial proliferations, bearing tiny, protuberant, circular scars. Conidia 14–18 μm long, 5–7 μm wide (\( {\bar{x}} \) = 16 × 6 μm, n = 20), solitary, fusiform, obtuse at both ends, mostly 3-septate, sometimes 4-septate, slightly constricted at septa, brown to dark brown in central cells and subhyaline at end cells, smooth. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Jizu Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2016, S.M. Tang, S-889 (MFLU 18–1601, holotype), ex-type living culture MFLUCC 18–1606.

Notes: Aquapteridospora fusiformis resembles A. lignicola in having macronematous, solitary, unbranched, septate conidiophores which are brown at the base and paler towards apex, polyblastic, terminal conidiogenous cells and fusiform, septate conidia. Both of these species also share similar size of conidiophores and conidia (Yang et al. 2015). However, A. fusiformis differs from A. lignicola in having pale brown to brown colonies, smooth conidia without a sheath, while the conidia of A. lignicola has large guttules in the middle cells and a conspicuous sheath. Phylogenetic analysis also shows that A. fusiformis and A. lignicola are distinct from other species, but they cluster together with strong support (Fig. 1, clade 15). To further support A. fusiformis as a new species, we compared nucleotide differences with A. lignicola (MFLU 15–1172) following the guidelines of Jeewon and Hyde (2016). Comparison of the 789 nucleotides across the LSU region reveals 9 bp differences. Based on the differences of morphology and DNA nucleotide, we introduce our isolate as new species in Aquapteridospora.


Magnaporthales Thongk et al.

Ceratosphaeriaceae Z.L. Luo, H.Y. Su & K.D. Hyde, fam. nov.

Index Fungorum: IF 555643; Facesoffungi number: FoF 05415

Asexual morph Harpophora-like. Phialides or short conidiophores arising on aerial hyphae, with conidial heads slimy, inconspicuous, and transparent. Conidiogenous cells phialidic, ampulliform to lageniform, terminal or intercalary, cylindrical. Conidia cylindrical, hyaline, aseptate, smooth. Sexual morph Stromata absent. Ascomata globose to pyriform, deeply immersed to almost superficial, dark brown to black, carbonaceous, with a long cylindrical, black or yellow crystals neck. Periphyses well-developed. Peridium composed of a large number of layers of very thick-walled rather small cells in the neck region. Interascal tissue of paraphyses thin-walled, probably evanescent at maturity. Asci 8-spored, unitunicate, cylindrical, fairly thin-walled, the apex truncate, with a conspicuous J-apical ring. Ascospores arranged biseriately, narrowly cylindric-fusiform, or filiform, the ends acute, thin-walled, hyaline, septate, guttulate, smooth-walled.

Type genus: Ceratosphaeria Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)

Notes: Réblová (2006) accommodated Ceratosphaeria in Magnaporthaceae based on their phylogenetic analyses of combined LSU and SSU sequence data. Phylogenetically, our multi-gene analyses show that Ceratosphaeria species formed a distinct clade in Magnaporthales and shares close relationship with Pseudohalonectriaceae (Fig. 15). Morphologically, Pseudohalonectriaceae is characterized by erumpent to immersed ascomata with a protruding, greenish yellow, bright yellow to brown neck, cylindrical to clavate asci with a nonamyloid, thimble-shaped, refractive, apical apparatus and cylindrical or ellipsoidal, straight to curved, multi-septate ascospores (Hongsanan et al. 2017). Ceratosphaeriaceae is distinct from Pseudohalonectriaceae in having narrowly cylindric-fusiform to filiform, longer ascospores. We therefore introduce a new family Ceratosphaeriaceae to accommodate Ceratosphaeria.

Fig. 15
figure 15

Phylogram generated from maximum likelihood analysis based on LSU and TEF1α sequence data for species of Magnaporthales (with Amplistroma erinaceum and A. caroliniana as outgroup). The best scoring RAxML tree with a final likelihood value of − 10365.588175 is presented. RAxML bootstrap support values equal to or greater than 60% are given before the forward slash (black). Bayesian posterior probability equal to or higher than 0.95 are given after the forward slash (red). Hyphen (‘-’) indicates a value lower than 60% for RAxML and Bayesian posterior probability lower than 0.95. Newly generated sequences are in red. Ex-type or ex-epitype strains are in bold


Ceratosphaeria Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)

Asexual morph Harpophora-like. Phialides or short conidiophores arising on aerial hyphae, conidial heads slimy, inconspicuous, and transparent. Conidiogenous cells phialidic, ampulliform to lageniform, terminal or intercalary, hyaline to subhyaline, cylindrical. Conidia cylindrical with curvature, hyaline, narrowly rounded at both ends, aseptate, smooth. Sexual morph Stromata absent. Ascomata globose to pyriform, often irregular in shape due to compression, deeply immersed to almost superficial, dark brown to black, carbonaceous, with a long cylindrical black or yellow crystals, thick-walled neck which is easily detached, scattered to densely aggregated. Peridium composed of a large number of layers of very thick-walled rather small cells in the neck region. Interascal tissue of paraphyses thin-walled, probably evanescent at maturity, periphyses well-developed. Asci 8-spored, unitunicate, cylindrical, fairly thin-walled, the truncate apex, with a conspicuous, J-, apical ring. Ascospores arranged biseriately, narrowly cylindric-fusiform, filiform, the ends acute, often slightly curved, thin-walled, hyaline, guttulate, smooth-walled.

Type species: Ceratosphaeria lampadophora (Berk. & Broome) Niessl, Verh. nat. Ver. Brünn 14: 203 (1876)

Notes: The genus Ceratosphaeria was introduced by Niessl (1876) with C. lampadophora as the type species and it is morphologically characterized in having globose to pyriform, immersed to almost superficial, dark coloured stromatic ascomata, leathery to fragile perithecial walls, cylindric-clavate, short-stipitate asci, truncate to broadly rounded at the apex, with an apical annulus and allantoid to suballantoid, pale brown, aseptate ascopores (Niessl 1876; Réblová 2006). In this study, we introduce two new species in Ceratosphaeria. Five species in this genus have been recorded from freshwater habitats.

Ceratosphaeria aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555644, Facesoffungi number: FoF 05416, Fig. 11

Etymology: Referring to the aquatic habitat of this fungus.

Holotype: MFLU 18–2323

Saprobic on decaying wood submerged in freshwater habitats. Asexual morph Undetermined. Sexual morphAscomata 269–361 μm high, 196–284 μm diam., deeply immersed with neck erumpent through host surface, globose to subglobose, brown to yellow brown, occurring solitary. Neck long, surface smooth, at times with yellow crystals. Peridium 29–43 μm thick, composed of an inner layer of flattened hyaline cells, a middle layer of small, polygonal to irregular, pale brown cells, an outer layer of irregular, yellow brown, pseudoparenchymatic cells. Paraphyses 4–7 μm wide, longer than asci, long tapering above. Asci 86–124 × 13–21 µm (\( {\bar{x}} \) = 105 × 17 µm, n = 30), 8-spored, unitunicate, cylindrical to broadly fusiform, with a narrow apical ring. Ascospores 89–95 × 4–7 µm (\( {\bar{x}} \) = 92 × 5.5 µm, n = 30), filiform, mostly 3-septate, guttulate, hyaline, smooth-walled.

Material examined: CHINA, Yunnan Province, saprobic on decaying wood submerged in a freshwater river, April 2015, Z.L. Luo, S-639 (MFLU 18–2323, holotype; HKAS 92859, isotype), ex-type living culture MFLUCC 18–1337.

Notes: Ceratosphaeria aquatica resembles C. phialidica in having deeply immersed ascomata, and filiform, hyaline ascospores (Huhndorf et al. 2008). However, Ceratosphaeria aquatica differs from C. phialidica in having broader cylindrical asci (13–21 vs. 5.5–6.5 μm) and guttulate, septate, larger ascospores (89–95 × 4–7 vs. 65–85 × 1.3–1.7 μm). Ceratosphaeria aquatica also shares similar morphological characters with C. lignicola with filiform, guttulate, hyaline ascospores. However, Ceratosphaeria aquatica differs from C. lignicola in having 3-septate, wider ascospores (4–7 vs. 3.5–4.5 μm). Phylogenetic analysis also shows that C. aquatica is distinct from C. lignicola (Fig. 15).

Ceratosphaeria lampadophora (Berk. & Broome) Niessl

Sphaeria lampadophora Berk. & Broome, Ann. Mag. nat. Hist., Ser. 3 3: 372 (1859)

Distribution: Australia, on submerged wood (Hyde et al. 1997)

Asexual morph: Harpophora-like. Detailed descriptions and illustrations refer to Réblová (2006).

Notes: LSU sequence data is available.

Ceratosphaeria phialidica (Shearer) Huhndorf et al.

Pseudohalonectria phialidica Shearer Can. J. Bot. 67(7): 1950 (1989)

Distribution: USA, submerged wood (Shearer 1989).

Asexual morph: Harpophora-like.

Notes: Holotype NY-01050492 and NY-01050493, other specimen collected from freshwater habitats: NY-03380687. Sequence data is not available.

Ceratosphaeria pusilla (Fuckel) Sacc

Ceratostoma pusillum Fuckel Jb. nassau. Ver. Naturk. 23-24: 128 (1870) [1869-70]

Distribution: USA, submerged wood in southern Illinois swamps (Shearer and Crane 1986).

Asexual morph: Undetermined

Notes: Sequence data is not available.

Ceratosphaeria lignicola D.F. Bao, Z.L. Luo, H.Y. Su & K.D. Hyde, sp. nov.

Index Fungorum number: IF 555645, Facesoffungi number: FoF 05417, Fig. 12

Etymology: Referring to this fungus dwelling on wood.

Holotype: MFLU 18–1457

Saprobic on decaying wood, submerged wood in freshwater. Asexual morph Undetermined. Sexual morphAscomata 390–470 μm diam., 500–600 μm high, solitary or aggregated, deeply immersed, subglobose, dark brown to black, with a yellow, cylindrical, periphysate neck. Peridium 13.5–17.5 μm wide, composed of an inner layer of flattened hyaline cells, a middle layer of small, polygonal to irregular, brown cells, an outer layer of irregular, yellow brown, pseudoparenchymatic cells. Paraphyses 4.5–6.5 μm wide, numerous, septate, hyaline, tapering distally, smooth. Asci 106–116 × 11–13 μm (\( {\bar{x}} \) = 111 × 12 μm, n = 20), 8-spored, unitunicate, broadly cylindrical, thin-walled, with a refractive apical apparatus. Ascospores 94–102 × 3.5–4.5 μm (\( {\bar{x}} \) = 98 × 4 μm, n = 30), hyaline, filiform, tapering at both ends, guttulate, smooth-walled.

Material examined: THAILAND, Khwaeng Hua Mak, Khet Bang Kapi Krung Thep Maha Nakhon, saprobic on decaying wood submerged in a freshwater stream, October 2017, Z.L. Luo, B-96, (MFLU 18–1457, holotype), ex-type living culture, MFLUCC 18–0342.

Notes: Ceratosphaeria lignicola resembles C. phialidica in having deeply immersed ascomata in the substrate (Huhndorf et al. 2008). Ceratosphaeria lignicola differs from C. phialidica in having broader cylindrical asci (11–13 vs. 5.5–6.5 μm) and larger, guttulate, ascospores (94–102 × 3.5–4.5 vs. 65–85 × 1.3–1.7 μm). Ceratosphaeria lignicola also shares similar morphological characters with C. lampadophora (Huhndorf et al. 2008). However, Ceratosphaeria lignicola differs from C. lampadophora in having longer, filiform, aseptate ascospores (94–102 vs. 52–72 μm), while C. lampadophora have fusiform, 5–7-septate ascospores. Phylogenetic analysis also shows that Ceratosphaeria lignicola is distinct from C. lampadophora (Fig. 15).


Magnaporthaceae P.F. Cannon

Aquafiliformis Z.L. Luo, K.D. Hyde & H.Y. Su, gen. nov.

Index Fungorum number: IF 555646, Facesoffungi number: FoF 05418

Etymology: “Aqua” referring to the aquatic habitats, “filiformis” referring to the filiform ascospores.

Saprobic on decaying wood. Asexual morph Undetermined. Sexual morphAscomata immersed with neck erumpent through host surface, globose to subglobose, dark brown to black, solitary. Peridium composed of an inner layer of flattened hyaline cells and an outer layer of irregular, heavily pigmented, thick-walled, pale brown to dark brown cells of textura angularis. Paraphyses hyaline, septate, unbranched. Asci 8-spored, unitunicate, cylindrical to clavate, hyaline. Ascospores filiform, aseptate, guttulate, hyaline, smooth-walled.

Type species: Aquafiliformis lignicola Z.L. Luo, K.D. Hyde & H.Y. Su

Notes: Aquafiliformis morphologically resembles Ceratosphaeria in having globose to subglobose, dark brown to black ascomata, cylindrical, 8-spored, unitunicate asci and filiform, hyaline, guttulate ascospores (Huhndorf et al. 2008). However, the phylogenetic analysis of combined LSU, SSU, RPB2 and TEF1α sequence data show that our newly obtained strains (MFLUCC 16–1341, MFLUCC 18–1338) clusters in Magnaporthaceae, while Ceratosphaeria belongs to the newly introduced Ceratosphaeriaceae (Fig. 1, clade 20, 22). Wijayawardene et al. (2018) accepted 23 genera in the family Magnaporthaceae. Twenty genera with available molecular sequence data are included in our phylogenetic analysis and 17 of them clusters together, while the newly obtained strains MFLUCC 16–1341 and MFLUCC 18–1338 form a separate clade in Magnaporthaceae and close to Muraeriata (SMH 2008a, SMH 2008b) (Fig. 15). However, our strain differs from Muraeriata species in having globose to subglobose ascomata, a peridium composed of an inner layer of flattened hyaline cells and an outer layer of irregular, heavily pigmented, thick-walled, pale brown to dark brown cells of textura angularis and filiform, aseptate ascospores, while Muraeriata species have lageniform to globose ascomata, a peridium with a middle layer of large, polygonal to irregular, hyaline cells that collapse or rupture creating large empty pockets, with an external brown crust and narrowly fusiform, septate ascospores (Huhndorf et al. 2008). The genera Clavatisporella and Herbampulla were also placed in Magnaporthaceae, but the sequence data for these two genera are unavailable, and morphology of our fungus are different from those of Clavatisporella and Herbampulla by its filiform ascospores (Scheuer and Nograsek 1993; Hyde 1995a). Therefore, we introduce a new genus Aquafiliformis to accommodate our collections.

Aquafiliformis lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555647, Facesoffungi number: FoF 05419, Fig. 13

Etymology: Referring to this fungus dwelling on wood.

Holotype: MFLU 18–2325

Saprobic on decaying wood submerged in freshwater habitats. Asexual morph Undetermined. Sexual morphAscomata 275–335 μm high, 300–356 μm diam., immersed with erumpent neck through host surface, globose to subglobose, dark brown to black, solitary. Peridium 24–34 μm thick, composed of an inner layer of flattened hyaline cells and an outer layer of irregular, heavily pigmented, thick-walled, pale brown to dark brown cells of textura angularis. Paraphyses 4.5–7.5 μm wide, hyaline, septate, unbranched. Asci 76–86 × 8–12 µm (\( {\bar{x}} \) = 81 × 10 µm, n = 25), 8-spored, unitunicate, cylindrical to clavate, hyaline. Ascospores 57–69 × 2.5–3.5 µm (\( {\bar{x}} \) = 63 × 3 µm, n = 20), filiform, guttulate, aseptate, hyaline, smooth-walled.

Material examined: CHINA, Yunnan Province, saprobic on decaying wood submerged in a freshwater River, April 2015, X.C. Tao, S-478 (MFLU 18–2325, holotype; HKAS 92814, isotype), ex-type living culture MFLUCC 16–1341; Ibid., saprobic on decaying wood submerged in a freshwater stream in Cangshan Mountain, August 2016, H.W. Shen, S-717 (MFLU 18–2324, paratype), living culture MFLUCC 18–1338.

Notes: Aquafiliformis lignicola resembles Neogaeumannomyces bambusicola in having immersed, globose to subglobose, solitary ascomata, unitunicate, cylindrical, hyaline asci and filiform, hyaline, guttulate ascospores (Liu et al. 2015a, b). However, Aquafiliformis lignicola differs from Neogaeumannomyces bambusicola in having different sizes of ascomata, asci and ascospores. Aquafiliformis lignicola has aseptate ascospores, while Neogaeumannomyces bambusicola has 2–3-septate ascospores. Phylogenetic analysis also support that they belong to different genera (Fig. 15).


Ophioceraceae Klaubauf et al.

Ophioceras Sacc., Syll. fung. (Abellini) 2: 358 (1883)

Asexual morph Undetermined. Sexual morphAscomata globose to elongate-globose, superficial to immersed, black, with a long neck. Peridium thick, blackened. Paraphyses hypha-like, hyaline, septate, numerous. Asci 8-spored, cylindrical, with small, refractive, apical rings. Ascospores filiform, narrowly fusoid to cylindrical, septate, hyaline.

Type species: Ophioceras dolichostomum (Berk. & M.A. Curtis) Sacc., Syll. fung. (Abellini) 2: 358 (1883)

Notes: Saccardo (1883) introduced Ophioceras based on O. dolichostomum (Berk. & M.A. Curtis) Sacc. Ophioceras species are commonly encountered on decaying woody substrates in freshwater habitats worldwide (Hyde 1992b; Hyde and Goh 1998a; Shearer et al. 1999; Tsui et al. 2001a,b; Thongkantha et al. 2009).

Ophioceras aquaticus Hu et al.

Distribution: China, Yunnan Province, on submerged wood (Hu et al. 2012c).

Asexual morph: Undetermined

Notes: Holotype IFRD 021–055. ITS, LSU and SSU sequence data obtained from ex-type culture are available.

Ophioceras arcuatisporum Shearer et al.

Distribution: Canada, Manitoba, Lake Rosabella, on submerged grasses; USA, Minnesota, Lake Itasca State Park, Elk lake, on Typha sp., Shagawa lake, on submerged herbaceous debri (Shearer et al. 1999).

Asexual morph: Undetermined

Notes: Holotype ILLS, Crane A-167–1. Sequence data is not available.

Ophioceras commune Shearer et al.

Distribution: China, Hong Kong, Lam Tsuen River, on submerged wood (Tsui et al. 2001b); Egypt, River Nile, on submerged wood (Abdel-Aziz 2016); Japan, Koito River, on submerged wood (Tsui et al. 2001a); Panama, Barro Colorado Islanda, on twig submerged in Allee Creek (Shearer et al. 1999); USA, Illinois, Louisiana, Minnesota, on submerged wood (Shearer et al. 1999), Florida, on submerged decaying wood (Raja et al. 2009b).

Asexual morph: Undetermined

Notes: Holotype ILLS, Shearer 408–2, other specimens collected from freshwater habitats: HKAS 92587, HKAS 92640, HKAS 92569. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Ophioceras dolichostomum (Berk. & M.A. Curtis) Sacc

Sphaeria dolichostoma Berk. & M.A. Curtis, Soc., Bot. 10(no. 46): 388 (1868) [1869]

Distribution: Australia, north Queensland, submerged wood (Hyde 1992b); Japan, Koito River, on submerged wood (Tsui et al. 2001a); USA, Florida, on submerged wood (Conway and Barr 1977); Seychelles, Riviere St Marie-Louis, on submerged wood (Hyde and Goh 1998a).

Asexual morph: Undetermined

Notes: Holotype anon. 51 (anon. 580), other specimen collected from freshwater habitats: BRIP 19330. ITS, LSU, SSU, TEF1α and MCM7 sequence data are available.

Ophioceras fusiforme Shearer et al.

Distribution: USA, Indiana, Shades State Park, small stream, on submerged decorticated woody debris (Shearer et al. 1999).

Asexual morph: Undetermined

Notes: Holotype ILLS, Shearer 837–1. SSU sequence data obtained from ex-type culture is available.

Ophioceras guttulatum Tsui et al.

Distribution: China, Hong Kong, Tai Po. Lain Tsuen River, on submerged wood (Tsui et al. 2001c); China, Yunnan Province, on submerged bamboo (Cai et al. 2006a).

Asexual morph: Undetermined

Notes: Holotype IFRD 8819. Sequence data is not available.

Ophioceras hongkongense Tsui et al.

Distribution: China, Hong Kong, Tai Po. Lain Tsuen River, on submerged wood (Tsui et al. 2001c)

Asexual morph: Undetermined

Notes: Holotype IFRD 8820. LSU and SSU sequence data are available.

Ophioceras leptosporum (S.H. Iqbal) J. Walker

Synonym: Gaeumannomyces leptosporus S.H. Iqbal, Trans. Br. mycol. Soc. 58(2): 346 (1972)

Distribution: UK, submerged plant stalks (Iqbal 1972).

Asexual morph: Undetermined

Notes: Holotype K(M) 35072. ITS, LSU, SSU, RPB1, TEF1α and MCM7 sequence data are available.

Ophioceras submersum D.F. Bao, Z.L. Luo, H.Y. Su & K.D. Hyde, sp. nov.

Index Fungorum number: IF 555648, Facesoffungi number: FoF 05420, Fig. 14

Etymology: Referring to the submerged habitats of the fungus

Holotype: MFLU 18–1459

Saprobic on decaying wood, submerged wood in freshwater. Asexual morph Undetermined. Sexual morphAscomata 500–600 μm diam., 300–400 μm high, scattered, solitary, deeply immersed, subglobose or ellipsoidal, coriaceous, black, with a long black neck. Ostiole central, with straight upright neck at one end, black, periphysate. Peridium 25–31 μm, thick-walled, composed two layers, inner layer of hyaline, small pseudoparenchyma cells, outer layer of pseudoparenchyma cells occluded with brown amorphous material, dark brown cells of textura angularis. Paraphyses 7–10 μm wide, hyaline, septate, constricted at septa, broader at base, tapering to the end, longer than asci, smooth. Asci 115–137 × 10–11 μm (\( {\bar{x}} \) = 126 × 10.5 μm, n = 15), 8-spored, unitunicate, cylindrical, subhyaline, apically rounded. Ascospores 87–109 × 3–4 μm (\( {\bar{x}} \) = 98 × 3.5 μm, n = 20), overlapping in ascus, filiform, slightly curved, thin-walled, multi-septate, rounded at both ends, hyaline, smooth-walled.

Material examined: THAILAND, Sai Khu Waterfall, on submerged decaying wood, 1 September 2017, C.G. Lin, B-25 (MFLU 18–1459, holotype), ex-type living culture, MFLUCC 18–0211.

Notes: Ophioceras submersum clusters in Ophioceras based on multi-gene phylogenetic analyses and is related to O. dolichostomum, O. hongkongense and O. venezuelense (Fig. 15). Ophioceras submersum resembles O. hongkongense in having subglobose, black ascomata with a long black neck, hyaline, septate paraphyses, unitunicate, cylindrical, subhyaline asci and filiform, hyaline, septate, smooth ascospores (Tsui et al. 2001c). However, Ophioceras submersum differs from O. hongkongense by its smaller ascomata and longer asci (Tsui et al. 2001c). Phylogenetic analysis also shows that they are distinct species (Fig. 1, clade 19).

Ophioceras tenuisporum Shearer et al.

Distribution: Panama, Barro Colorado Islanda, on twig submerged in Allee Creek (Shearer et al. 1999);

Asexual morph: Undetermined

Notes: Holotype ILLS, Shearer 652–1. LSU, SSU and β-tubulin sequence data are available.

Ophioceras venezuelense Shearer et al.

Distribution: Venezuela, on submerged decorticated wood (Shearer et al. 1999).

Asexual morph: Undetermined

Notes: Holotype ILLS, Crane A-109–1. SSU sequence data is available.


Pseudohalonectriaceae Hongsanan & K.D. Hyde

Pseudohalonectria Minoura & T. Muroi, Trans. Mycol. Soc. Japan 19(2): 132 (1978)

Asexual morph Hyphomycetous, phialidic. Phialides hyaline, micronematous, flask-shaped. Conidia allantoid, hyaline, aseptate. Sexual morphAscomata immersed or partially immersed, with a long neck, globose to subglobose. Peridium membranous. Paraphyses numerous, septate, hyaline. Asci unitunicate, cylindrical, straight or curved, with J-, thimble-shaped apical ring. Ascospores overlapping uniseriate to biseriate, multi-seriate, filiformes, septate.

Type species: Pseudohalonectria lignicola Minoura & T. Muroi, Trans. Mycol. Soc. Japan 19(2): 132 (1978)

Notes: The genus Pseudohalonectria was introduced to accommodate P. lignicola (Minoura and Muroi 1978). Hongsanan et al. (2017) introduced Pseudohalonectriaceae as a new family within Magnaporthales to accommodate Pseudohalonectria. Sixteen species are accepted in this genus, of which six species have been reported from freshwater habitats worldwide (Minoura and Muroi 1978; Shearer 1989a, b; Hyde et al. 1998b; Cai et al. 2002a).

Pseudohalonectria adversaria Shearer

Distribution: South Africa, Durban, Palmiet River, on submerged wood (Hyde et al. 1998b); USA, Illinois, on submerged woody debris from Deer Pond (Shearer 1989a, b).

Asexual morph: Undetermined

Notes: Holotype ILLS, CS-603–1. Sequence data is not available.

Pseudohalonectria falcata Shearer

Distribution: USA, Illinois, on submerged twig collected from Quiver Creek (Shearer 1989).

Asexual morph: Undetermined

Notes: Holotype NY-01050484, paratype NY-01050483, other specimens collected from freshwater habitats NY-03380667, NY-03380668. LSU, SSU and TEF1α sequence data are available.

Pseudohalonectria fuxianii Cai et al.

Distribution: China, Yunnan Province, Lake Fuxian, on submerged wood (Cai et al. 2002a).

Asexual morph: Undetermined

Notes: Holotype IFRD 8838. Sequence data is not available.

Pseudohalonectria lignicola Minoura & T. Muroi

Distribution: China, Yunnan Province, Lake Fuxian, on submerged wood (Cai et al. 2002a); Japan, Koito River, on submerged wood (Tsui et al. 2001a), Lake Biwa, on submerged wood (Minoura and Muroi 1978); USA, Illinois, Indiana, on submerged wood in streams or Rivers (Shearer 1989a, b).

Asexual morph: Undetermined

Notes: Holotype HUT 40005. ITS, LSU, SSU, RPB1, TEF1α, MCM7 and β-tubulin sequence data are available.

Pseudohalonectria longirostrum Shearer

Distribution: Panama, a twig submerged in Shannon Creek (Shearer 1989).

Asexual morph: Undetermined

Notes: Holotype CS-656-1, NY. Sequence data is not available.

Pseudohalonectria lutea Shearer

Distribution: China, Yunnan Province, Lake Fuxian, on submerged wood (Cai et al. 2002a); Chile, submerged wood collected from a small stream (Shearer 1989a).

Asexual morph: Undetermined

Notes: Holotype NY-01050490, NY-01050491. LSU sequence data is available.


Myrmecridiales Crous

Myrmecridiaceae Crous

Myrmecridium Arzanlou et al., Stud. Mycol. 58: 84 (2007)

Asexual morphColonies flat, with immersed mycelium. Conidiophores arising vertically and clearly distinct from creeping hyphae, unbranched, straight or flexuose, septate, thick-walled. Conidiogenous cells polyblastic, integrated, terminally, cylindrical. Conidia solitary, subhyaline, smooth or finely verrucose, obovoidal or fusiform, conidial secession schizolytic. Sexual morphAscomata solitary or aggregated in small groups, immersed, hyaline to pale brown. Papilla or short necks centrally located, opening flush with the wood surface or slightly projecting. Ostiole periphysate. Clypeus positioned slightly beneath the wood surface. Ascomatal wall two layered. Paraphyses hyaline, septate, slightly constricted at the septa, cylindrical. Asci cylindrical, with a slender, long stipe. Ascospores ellipsoidal, delicately verruculose, septate, hyaline.

Type species: Myrmecridium schulzeri (Sacc.) Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 84 (2007)

Notes: The genus Myrmecridium was introduced by Arzanlou et al. (2007) to accommodate Ramichloridium-like taxa with hyaline mycelium, and pale to unpigmented conidiophores with pimple-like conidiogenous denticles at their apices. Twelve species are accepted in this genus (Peintner et al. 2016; Réblová et al. 2016b; Tibpromma et al. 2017) and two of them were collected from freshwater habitats (Réblová et al. 2016b; Tibpromma et al. 2017).

Myrmecridium aquaticum Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555649, Facesoffungi number: FoF 05421, Fig. 16

Fig. 16
figure 16

Myrmecridium aquaticum (MFLU 18–1595, holotype) a, b Colony on natural substrate. c, d Conidiophore with conidia. e, f Conidiogenous cells with conidia. gi Conidia. j Germinating conidium. Culture on PDA from above (k) and reverse (l). Scale bars: c, d 100 μm, e, f 30 μm, gj 10 μm

Etymology: Referring to the aquatic habitat of this fungus

Holotype: MFLU 18–1595

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies on natural substrate superficial, effuse, hairy, brown. Mycelium immersed, composed of septate, branched, smooth, hyaline hyphae. Conidiophores 211–308 µm long, 5–7 μm wide (\( {\bar{x}} \) = 258 × 6 µm, n = 20), macronematous, mononematous, erect, unbranched, multi-septate, straight or flexuous, cylindrical, percurrently proliferating, brown, paler towards the apex, smooth, nodose at the tip. Conidiogenous cells holoblastic, polyblastic, integrated, terminal, later becoming intercalary, subhyaline to pale brown. Conidia 14–16 µm long, 4–6 μm wide (\( {\bar{x}} \) = 15 × 5 µm, n = 20), acropleurogenous, dry, obovoid, rounded at the apex, pointed at the base, 3-septate, subhyaline, smooth-walled. Sexual morph Undetermined

Material examined: CHINA, Yunnan Province, Lancang River, saprobic on submerged decaying wood, April 2015, X.C. Tao, S-448 (MFLU 18–1595, holotype; HKAS 92833, isotype), ex-type living culture MFLUCC 15–0366, KUMCC 15–0340; Jizu Mountain, saprobic on decaying wood submerged in a freshwater stream, April 2017, N Zhao, S-1158, living culture MFLUCC 18–1489; Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, March 2014, X.Y. Liu, S-001.

Notes: Myrmecridium aquaticum resembles M. sorbicola in having solitary, erect, unbranched, multi-septate, brown conidiophores, integrated, terminal and intercalary conidiogenous cells and obovoid, smooth conidia rounded at the apex (Crous et al. 2018). However, Myrmecridium aquaticum differs from M. sorbicola in having larger conidiophores (211–308 vs. 50–200 µm), 3-septate and longer conidia (14–16 vs. 8–10 µm), while M. sorbicola has almost 1-septate conidia with mucoid sheath surrounding conidium in median region. Phylogenetic analysis shows that Myrmecridium aquaticum is distinct from other Myrmecridium species (Fig. 17).

Fig. 17
figure 17

Phylogram generated from maximum likelihood analysis based on combined ITS and LSU sequence data for species of Myrmecridium (with Acrodictys elaeidicola as outgroup). The best scoring RAxML tree with a final likelihood value of − 4631.789675 is presented. RAxML bootstrap support values equal to or greater than 75% are given before the forward slash (black). Maximum parsimony bootstrap support values equal to or greater than 75% are given after the forward slash (red). Hyphen (‘-’) indicates a value lower than 75% for RAxML and Maximum parsimony. Newly generated sequences are in red. Ex-type strains are in bold

Myrmecridium fluviae Hyang B. Lee & T.T.T. Nguyen

Distribution: Korea, Jeonnam Province, Yeongsan River located in Gwangju, from a freshwater sample (Tibpromma et al. 2017).

Sexual morph: Undetermined

Notes: Holotype CNUFC YR61–1; ITS and LSU sequence data are available.

Myrmecridium montsegurinum Réblová & J. Fourn

Distribution: France, Midi-Pyrénées: Ariège, Montségur, Le Lasset stream along D9 road, on submerged wood of Fraxinus excelsior, Fagus sylvatica, Hedera helix and Alnus glutinosa (Réblová et al. 2016b)

Asexual morph: Undetermined

Notes: Holotype PRM 934684, other specimens collected from freshwater habitats: PRM 934685, PRM 934686. ITS, LSU, SSU and RPB2 sequence data are available.


Ophiostomatales Benny & Kimbr.

Ophiostomataceae Nannf.

Subbaromyces Hesselt., Bull. Torrey bot. Club 80: 511 (1953)

Asexual morphConidiophores branched, septate. Conidia hyaline, smooth-walled, asepate, exogenously formed, ellipsoid. Sexual morphAscomata partially submerged, later superficial, membranous, syringe-shaped, beak divided into two portions by a large pronounced collar, with upper portion tapering to a small ostiole, surrounded by a fringe of hyphae. Paraphyses absent. Asci 8-spored, unitunicate, clavate, with autodigestible wall. Ascospores 2-celled, with cells of equal size, hyaline, surrounded by a gelatinous envelope, released as a mucus-like droplet at tip of perithecium.

Type species: Subbaromyces splendens Hesselt., Bull. Torrey bot. Club 80: 511 (1953)

Notes: The genus was established by Hesseltine (1953) for a taxon collected from trickling filter rocks in New York, USA. Two species were accepted within this genus (Hesseltine 1953; Chary and Ramarao 1974). The other accepted species, S. aquaticus, was introduced by Chary and Ramarao (1974) for a species isolated from water samples collected in India. A phylogenetic analysis based on SSU sequence data of S. splendens was provided by Jones et al. (1999), and it showed that S. splendens is closely related to Curvularia brachyspora. In updated classifications, Subbaromyces was placed in the family Ophiostomataceae (Ophiostomatales, Sordariomycetes) (Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2017, 2018).

Subbaromyces aquaticus Manohar. & P. R. Rao

Distribution: India, on submerged wood in freshwater (Chary and Ramarao 1974).

Asexual morph: Undetermined

Notes: Sequence data is not available.


Phomatosporales Senan. et al.

Phomatosporaceae Senan. & K.D. Hyde

Phomatospora Sacc., Nuovo G. bot. ital. 7: 306 (1875)

Asexual morphSporothrix-like, reported from culture (Rappaz 1992). Sexual morphAscomata solitary to rarely gregarious, immersed or becoming erumpent with age, globose or subglobose, light brown, dark brown to black, coriaceous, sometimes developing under a small blackened clypeus, ostiolate, papillate. Papilla short or rarely somewhat long, central or eccentric, cylindrical, sometimes covered with black, amorphous material around the upper region, periphyses hyaline, short, filiform. Peridium comprising small, brown pseudoparenchymatous cells forming a textura angularis to textura prismatica or inner, hyaline, thick-walled cells of textura angularis and outer, brown, cells of textura angularis. Paraphyses comprising hypha-like, filamentous, septate or aseptate, slightly constricted at the septa, distally tapering, hyaline. Asci 8-spored, unitunicate, cylindrical or oblong-fusiform, thin-walled, short stalked or sessile, apex oblong with J-, apical apparatus. Ascospores uniseriate, rarely biseriate, overlapping uniseriate to biseriate, ellipsoidal to fusiform, 0–3-septate, not constricted at the septum, sometimes bi-guttulate, guttules located at the ends of the cell, or longitudinally striate, sometimes with filamentous appendages at both ends, hyaline.

Type species: Phomatospora berkeleyi Sacc., Nuovo G. bot. ital. 7(4): 306 (1875)

Notes: Phomatospora was placed in Ascomycota genera incertae sedis based on phylogenetic analysis (Vijaykrishna et al. 2006). Senanayake et al. (2016) established the family Phomatosporaceae (Phomatosporales) to accommodate the genera Phomatospora, Lanspora and Tenuimurus. Members of the genus Phomatospora are widely distributed in freshwater, marine and terrestrial habitats. Seven species of Phomatospora are known from freshwater habitats (Shearer and Raja http://fungi.life.uiuc.edu/world_records; Nordén et al. 2015).

Phomatospora aquatica Minoura & T. Muroi

Distribution: Australia, Queensland, submerged wood in stream (Vijaykrishna and Hyde 2006); Japan, on basal wood submerged in Pond Nanatsuike (Minoura and Muroi 1978).

Asexual morph: Undetermined

Notes: Holotype HUT 40004. Sequence data is not available.

Phomatospora berkeleyi Sacc

Distribution: Australia, Queensland, submerged wood in stream (Vijaykrishna and Hyde 2006); USA, Illinois, Tinley Park Forest Preserve, Typha pond, on submerged stems of Typha latifolia; Wisconsin, Trout lake, on submerged stems of Carex comosa, Big Muskellunge lake, on submerged stems of Scirpus brevicaudatus, Allequash lake, on submerged stems of Typha latifolia (Fallah and Shearer 1998).

Asexual morph: Undetermined

Notes: Holotype K(M) 49573, other specimens were collected from freshwater: ILLS 53088, ILLS 53089, ILLS 53091. Sequence data is not available. Phomatospora berkeleyi was originally collected from dead stalks of Solanum on terrestrial habitats (Saccardo 1875). Fallah and Shearer (1998) collected this species from freshwater habitats and provided descriptions and illustrations.

Phomatospora helvetica H. Wegelin

Distribution: Norway, Aust-Agder, Arendal, Langevoll, Nedenes, on submerged wood in small stream in temperate deciduous forest (Nordén et al. 2015).

Asexual morph: Undetermined

Notes: Sequence data is not available.

Phomatospora luteotingens J. Fourn. & Lechat

Distribution: France, Ariège: Caussou, Caussou Brook, on submerged branch of Fraxinus excelsior, Hautes Pyrénées: Asque, La Gourgue, Arros stream, on submerged twig of Alnus glutinosa; Spain, Asturias: road to La Endriga, on submerged wood of Alnus glutinosa (Fournier and Lechat 2010)

Asexual morph: Undetermined

Notes: Holotype LIP, Fournier 09145. Sequence data is not available.

Phomatospora muskellungensis Fallah & Shearer

Distribution: USA, Wisconsin, Big Muskellunge lake, submerged wood of Typha latifolia, Allequash lake, on submerged stems of Typha latifolia (Fallah and Shearer 1998).

Asexual morph: Undetermined

Notes: Holotype ILLS 53011. Sequence data is not available.

Phomatospora striatigera Scheuer

Distribution: Australia, on submerged Carex gracili in freshwater (Scheuer 1988)

Asexual morph: Undetermined

Notes: Holotype GZU, Scheuer, 18 Jun. 1981, Epitype CBS 133932. ITS and LSU sequence data are available.

Phomatospora triseptata Raja & Shearer

Distribution: USA, Florida, Big Cypress National Preserve, on submerged soft, decorticated woody debris (Raja and Shearer 2008).

Asexual morph: Undetermined

Notes: Holotype ILL 40114. Sequence data is not available.


Sporidesmiales Crous

Sporidesmiaceae Fr.

Sporidesmium Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 41 (1809)

Asexual morph Descriptions and illustrations refer to Su et al. (2016). Sexual morph Descriptions and illustrations refer to Zhang et al. (2017a, b).

Type species: Sporidesmium atrum Link, Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 41 (1809)

Notes: The genus Sporidesmium was established by Link (1809) with S. atrum Link as the type. Sporidesmium is a large and heterogeneous genus with 483 epithets referred to the genus in Index Fungorum (December 2018). However, many previously described species were revised and transferred to over 30 genera (Iturriaga et al. 2008). Studies based on phylogenetic analyses have been carried out to further re-examine the classification of Sporidesmium-like taxa, given that the generic delimitations based on morphological characters appear to be questionable, and it has been found that Sporidesmium and its morphology similar genera are clearly not monophyletic, and even they are distributed among different families and orders within Dothideomycetes and Sordariomycetes (Shenoy et al. 2006; Su et al. 2016; Yang et al. 2018a). Sporidesmium was only found as an asexual morph genus until Zhang et al. (2017a, b) introduced the first sexual morph Sporidesmium thailandense which was collected from freshwater habitats in Thailand.

Sporidesmium aquaticivaginatum J. Yang & K.D. Hyde

Distribution: Thailand, Prachuap Khiri Khan Province, Hua Hin, stream flowing outside Kaeng Krachan National Park, on submerged wood (Hyde et al. 2016b).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–1159, isotype HKAS 95046. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Sporidesmium cangshanense Z.L. Luo & K.D. Hyde, nom. nov.

Facesoffungi number: FoF 05422

Sporidesmium aquaticum H.Y. Su, Z.L. Luo & K.D. Hyde, Fungal Divers 80:398 (2016)

Distribution: China, Yunnan Province, Dali city, on submerged wood in a freshwater stream in Cangshan Mountain (Su et al. 2016).

Sexual morph: Undetermined

Notes: Holotype HKAS 84029. LSU sequence data is available. Sporidesmium aquaticum was introduced by Arambarri et al. (1989). Su et al. (2016) introduced a new Sporidesmium species also called Sporidesmium aquaticum which is a homonym and thus invalid. We replace the latter invalid name, Sporidesmium aquaticum, as Sporidesmium cangshanense.

Sporidesmium dulongense Luo et al.

Distribution: China, Yunnan Province, saprobic on decaying wood submerged in Dulong River (Hyde et al. 2019).

Sexual morph: Undetermined

Notes: Holotype HKAS 92659. ITS, LSU, RPB2 and TEF1α sequence data are available.

Sporidesmium fluminicola H.Y. Su & K.D. Hyde

Distribution: China, Yunnan Province, Dali, Cangshan Mountain, saprobic on decaying wood submerged in stream (Su et al. 2016).

Sexual morph: Undetermined

Notes: Holotype HKAS 84003. ITS, LSU and SSU sequence data are available.

Sporidesmium gyrinomorphum Yang et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on decaying wood submerged in a freshwater stream (Yang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–0851, isotype GZAAS 17–0004. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Sporidesmium lageniforme Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555650, Facesoffungi number: FoF 05423, Fig. 18

Fig. 18
figure 18

Sporidesmium lageniforme (MFLU 18–1594, holotype) a, b Colonies on substrate. c, d Conidiophore with conidia. e Conidiophores. f Conidiogenous cells gi Conidia j Germinating conidium. Culture on PDA from above (k) and reverse (l). Scale bars: ce 50 μm, fl 20 μm

Etymology: Referring to the lageniform conidia of the fungus

Holotype: MFLU 18–1594

Saprobic on decaying wood submerged in freshwater. Asexual morphColonies on the substratum superficial, effuse, scattered, hairy, black. Mycelium mostly immersed, comprising of branched, septate, smooth-walled, brown hyphae. Conidiophores 105–141 μm long, 4–6 μm wide (\( {\bar{x}} \) = 123 × 5 μm, n = 20), macronematous, mononematous, erect, straight or flexuous, unbranched, 4–6-septate, greyish brown to dark brown, smooth. Conidiogenous cells monoblastic, integrated, terminal, determinate, cylindrical, dark brown. Conidia 38–48 μm long, 13–17 μm wide (\( {\bar{x}} \) = 43 × 15 μm, n = 20), acrogenous, solitary, lageniform, truncate at base, tapering at apex, 5–7-septate, subhyaline to greyish at immature, dark olivaceous-brown at maturity, hyaline towards the apex, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Nujiang River, saprobic on decaying submerged wood, October 2016, Z.L. Luo, S-880 (MFLU 18–1594, holotype), ex-type living culture DLUCC 0880.

Notes: Sporidesmium lageniforme resembles S. dulongense in having unbranched, dark brown, septate conidiophores, monoblastic, terminal, determinate conidiogenous cells and acrogenous, solitary, septate conidia truncate at base, tapering at apex. Sporidesmium lageniforme differs from S. dulongense in having shorter conidia (38–48 vs. 50–58 μm) and conidia of S. dulongense have sphaerical guttules in each cell while conidia of S. lageniforme mostly have no guttules. Phylogenetic analysis also shows that Sporidesmium lageniforme and S. dulongense are distinct species (Fig. 1, clade 7).


Sporidesmium lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555651, Facesoffungi number: FoF 05424, Figs. 19, 20

Fig. 19
figure 19

Sexual morph ofSporidesmium lignicola (MFLU 18–2326, holotype) a, b Appearance of necks on substrate. c Section through ascoma. d, e Structure of peridium. f Paraphyses. gi Asci. jo Ascospores. Scale bars: c 100 μm, d 50 μm, fi 30 μm, jo 5 μm

Fig. 20
figure 20

Asexual morph ofSporidesmium lignicola (DLU 1376) a Colonies on natural substrate. b Conidiophore with conidia. cm Conidia. Scale bars: bm 20 μm

Etymology: Referring to the fungus dwelling on wood.

Holotype: MFLU 18–2326

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse on natural substrate, scattered, pale brown to brown. Mycelium immersed, composed of septate, branched, brown, smooth hyphae. Conidiophores 50–70 μm long, 3–4 μm wide (\( {\bar{x}} \) = 60 × 3.5 μm, n = 20), macronematous, mononematous, unbranched, erect, straight or flexuous, brown to dark brown, septate, smooth. Conidiogenous cells holoblastic, monoblastic, terminal, determinate, dark brown, cylindrical. Conidia 21–27 μm long, 4.5–6.5 μm wide (\( {\bar{x}} \) = 24 × 5.5 μm, n = 20), acrogenous, solitary, dry, clavate or cylindrical, straight or slight curved, truncate at base, brown, mostly 3-septate, formed in chains, smooth. Sexual morphAscomata 190–330 μm high, 160–340 μm diam., immersed with neck erumpent through host surface, subglobose to ellipsoid, scattered, 2–4 locules, pale brown to brown. Ostiole 185–365 μm long, 78–122 μm wide, cylindrical, central or lateral, brown at the base, becoming hyaline towards the apex. Peridium 30–44 μm thick, two-layered, outer layer comprising pale brown to brown, oblong and rounded cells, inner layer comprising several layers of hyaline, large cells of textura angularis or irregular cells. Paraphyses 2.5–4.5 μm wide, hyaline, unbranched, septate, slightly constricted at the septum. Asci 87–115 × 8–10 µm (\( {\bar{x}} \) = 101 × 9 µm, n = 35), 8-spored, unitunicate, cylindrical, with a tapering pedicel, apically rounded, with a distinct, relatively small, refractive, wedge-shaped apical ring. Ascospores 13–15 × 6–8 µm (\( {\bar{x}} \) = 14 × 7 µm, n = 35), uni-seriate, fusiform, aseptate or uniseptate, guttulate, hyaline, smooth-walled.

Material examined: CHINA, Yunnan Province, saprobic on decaying wood submerged in a freshwater river, April 2015, X.C. Tao, S-429 (MFLU 18–2326, holotype), ex-type living culture KUMCC 15–0266; Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2017, H.W. Shen, S-1376 (DLU 1376, paratype), living culture DLUCC 1376.

Notes: The sexual morph of Sporidesmium lignicola resembles Sporidesmium thailandense in having subglobose to ellipsoid, scattered, immersed ascomata with necks erumpent through the host surface, hyaline, unbranched, septate paraphyses, cylindrical asci apically rounded with an apical ring and fusiform, hyaline ascospores (Zhang et al. 2017). However, Sporidesmium lignicola differs from S. thailandense in having ascomata with 2–4 locules, a two-layered peridium, smaller asci (87–115 × 8–10 vs. 160–220 × 11–14 µm), and aseptate or uniseptate, smaller ascospores (13–15 × 6–8 vs. 23–28 × 8–10 µm) with small guttules, while S. thailandense has 3-septate ascospores with 2–5 prominent guttules. The asexual morph of Sporidesmium lignicola can be easily distinguished from other Sporidesmium asexual morph species in having clavate or cylindrical conidia formed in chains. Phylogenetically, our two newly obtained strains cluster together with high support (100% ML) and is distinct from other Sporidesmium species (Fig. 1, clade 7). We therefore introduce a new species Sporidesmium lignicola with both asexual and sexual morphs.

Sporidesmium olivaceoconidium J. Yang & K.D. Hyde

Distribution: Thailand, Chiang Rai Province, stream flowing in Tham Luang Nang Non Cave, on submerged wood (Hyde et al. 2016b).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–1175. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Sporidesmium pyriformatum J. Yang & K.D. Hyde

Distribution: Thailand, Khiri Khan Province, Hua Hin, stream flowing outside Kaeng Krachan National Park, on submerged wood (Hyde et al. 2016b).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–1155. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Sporidesmium submersum H.Y. Su & K.D. Hyde

Distribution: China, Yunnan Province, Dali, Cangshan Mountain, saprobic on decaying wood submerged in stream (Su et al. 2016).

Sexual morph: Undetermined

Notes: Holotype HKAS 84004. ITS, LSU and SSU sequence data are available.

Sporidesmium thailandense Dong et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged wood in a small River (Zhang et al. 2017a, b; Yang et al. 2018a).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2709, other specimen collected from freshwater habitats MFLU 15–1152. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available.

Sporidesmium tropicale M.B. Ellis

Distribution: Thailand, Prachuap Khiri Khan Province, on decaying wood submerged in a freshwater stream (Yang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype IMI 37498, other specimen collected from freshwater MFLU 17–0850. ITS, LSU, SSU, RPB2 and TEF1α sequence data are available. Sporidesmium tropicale was found on dead branches of woody plants and is widely distributed in tropical areas (Ellis 1958; Wu and Zhuang 2005). Yang et al. (2018a, b) collected this species from freshwater habitats in Thailand and provided the illustrations and descriptions.


Tirisporellales Suetrong et al.

Tirisporellaceae Suetrong et al.

Thailandiomyces Pinruan, Sakayaroj, Hyde & Jones, Fungal Diversity 29: 91 (2008)

Asexual morphColonies on natural substrate effuse, black. Mycelium superficial. Conidiophores macronematous, mononematous, erect, brown, paler towards the apex, straight or flexuous, branched or unbranched. Conidiogenous cells enteroblastic, monophialidic, integrated, terminal, with a large and distinct collarette; collarette funnel-shaped. Conidia cylindrical, ellipsoid or obovoid, thick-walled, brown, aseptate. Sexual morphAscomata scattered to gregarious, semi-immersed to superficial, globose, black, coriaceous, ostiolate, with long cylindrical necks, periphysate with short hyaline cells. Peridium composed of compressed cells oftextura angularis, black outside, becoming brown inwardly. Paraphyses present but deliquescent, irregular in width, rarely septate, tapering towards the apices, embedded in a mucilaginous matrix. Asci 8-spored, unitunicate, cylindrical to clavate, apedicellate, free-floating, apically truncate, with a J- subapical ring. Ascospores uniseptate, obliquely overlapping 2-seriate, fusoid, straight or broadly curved, with bipolar appendages, hyaline, smooth-walled.

Type species: Thailandiomyces bisetulosus Pinruan et al., Fungal Divers 29: 91 (2008)

Notes: The genus Thailandiomyces was introduced by Pinruan et al. (2008) to accommodate one ascomycete species collected from submerged trunk of Licuala longicalycata in Thailand. This species was apparently linked with a Craspedodidymum asexual morph in culture. This genus remains monotypic. Based on phylogenetic analyses, Suetrong et al. (2015) introduced a new family Tirisporellaceae, typified by a new genus Tirisporella E.B.G. Jones, K.D. Hyde & Alias. The genus Thailandiomyces phylogenetically resides in this family.

Thailandiomyces bisetulosus Pinruan et al.

Distribution: Thailand, on submerged trunk of Licuala longicalycata (Pinruan et al. 2008).

Asexual morph: Craspedodidymum licualae Pinruan

Notes: Holotype BBH, Pinruan, Wah 110. LSU and SSU sequence data are available.


Togniniales Senan. et al.

Togniniaceae Réblová et al.

Phaeoacremonium Gams et al., Mycologia 88: 789 (1996)

Asexual morphMycelium consisting of branched, septate hyphae, single or bundled. Conidiophores branched in the basal region or unbranched, arising from aerial or submerged hyphae, erect, nearly cylindrical when unbranched, slightly tapering, straight or flexuous, variable in length, up to 7-septate, mostly pale brown, paler towards the tip. Conidiogenous cells mostly monophialidic, discrete or integrated, terminal or intercalary, sometimes polyphialidic, sparsely warted, pale brown to hyaline, verruculose or smooth. Conidia aggregated into round, slimy heads at the apices of phialides, aseptate, hyaline, smooth-walled; oblong-ellipsoidal to obovate, cylindrical, allantoid or reniform, uncommonly fusiform-ellipsoidal or globose, becoming guttulate with age. Sexual morphAscomata aggregated or solitary, superficial to immersed, nonstromatic, globose to subglobose, dark, opaque, long-necked. Paraphyses abundant, broadly cellular, slightly constricted at the septa, branching, hyaline, slightly tapering apically or thread-like towards the apex. Asci 8-spored, unitunicate, arising in acropetal succession, appearing spicate when mature, ascal apex thickened without a discharge mechanism, basally bluntly obtuse, sessile. Ascospores mostly biseriate or in a single row, allantoid, reniform, cylindrical or oblong-ellipsoidal, aseptate, hyaline.

Type species: Phaeoacremonium parasiticum (Ajello, Georg & C.J.K. Wang) Gams et al., Mycologia 88(5): 794 (1996)

Notes: Phaeoacremonium has recently been monographed, and comprises 46 species (Gramaje et al. 2015; Maharachchikumbura et al. 2016). Phaeoacremonium species are saprobic on plants, or pathogenic on human and animals or soil fungi (Mostert et al. 2006; Maharachchikumbura et al. 2016). Hu et al. (2012c) introduced a new species, Togninia aquatica, collected from freshwater habitats. Gramaje et al. (2015) combined this species as Phaeoacremonium aquaticum.

Phaeoacremonium aquaticum (Hu et al.) Gramaje et al.

Togninia aquatica Hu et al. Mycologia 104(6): 1482 (2012)

Distribution: China, Yunnan Province, Jinghong City, Mengla County, on submerged wood in a small stream (Hu et al. 2012c).

Asexual morph: Undetermined

Notes: Holotype IFRD 023–047. ITS sequence data is available.

Phaeoacremonium ovale Huang et al.

Distribution: China, Yunnan Province, Baoshan City, on submerged decayed wood in a stream along the roadside (Huang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype HKAS 99550, isotype MFLU 18–1076. ITS, actin and β-tubulin sequence data are available.


Trichosphaeriales M.E. Barr

Trichosphaeriaceae G. Winter

Brachysporium Sacc., Syll. fung. (Abellini) 4: 423 (1886)

Asexual morphColonies effuse, brown, velvety. Mycelium mostly immersed, composed of branched, septate, subhyaline to brown hyphae. Conidiophores mononematous, macronematous, erect, straight or slightly flexuous, smooth, thick-walled, septate, unbranched, cylindrical, brown in the bottom, paler and tapering toward the apex. Conidiogenous cells holoblastic, terminal, integrated, hyaline, denticulate, proliferating sympodially. Conidia acropleurogenous, septate, smooth, thick-walled, fusoid to limoniform, polar cells subhyaline, narrowing at the apex, median cells brown. Sexual morphAscomata partly immersed to superficial, solitary, globose to subglobose, conical around the ostiole, papillate, dark brown to black, glabrous. Peridium leathery to fragile, consisting of two regions; outer region of carbonaceous, dark brown, angular to rectangular cells; inner region of hyaline, thin-walled, elongated, compressed cells. Ostiolar canal periphysate. Paraphyses persistent, branched, hyaline, septate, irregular in width. Asci 8-spored, unitunicate, cylindrical-clavate, with long, slender stipe, broadly rounded to truncate at the apex, with distinct refractive apical annulus. Ascospores biseriate, ellipsoidal to fusiform to oblong-lemon-shaped, at first aseptate, followed by formation of a median septum, hyaline, smooth-walled.

Type species: Brachysporium obovatum (Berk.) Sacc., Syll. fung. (Abellini) 4: 427 (1886)

Notes: The asexual morph genus Brachysporium was established by Saccardo in 1880. Réblová and Seifert (2004a) introduced a new genus Cryptadelphia, with type species C. groenendalensis (Sacc., E. Bomm. & M. Rouss.) Réblová & Seifert to accommodate six sexual morphs of Brachysporium. Many Brachysporium species were reported saprobic on decaying wood of different substrates (Réblová and Seifert 2004a; Markovskaja and Treigien 2007). Some species were also described from marine habitats, e.g. B. belgolandicum Schaumann and B. helgolandicum Schaumann. Among the accepted Brachysporium species, only two are known from freshwater habitats (Lamore and Goos 1978; Raja et al. 2009b).

Brachysporium obovatum (Berk.) Sacc.

Helminthosporium obovatum Berk., Annals and Magazine of Natural History 6: 434 (1841)

Distribution: USA, Florida, on submerged substrate in lentic habitat in Ocala National Forest (Raja et al. 2009b).

Sexual morph: see Réblová and Seifert (2004a)

Notes: Sequence data is not available.

Brachysporium nigrum (Link) S. Hughes

Distribution: USA, Rhode Island, on wood in the Saugatucket River (Lamore and Goos 1978).

Sexual morph: see Réblová and Seifert (2004a).

Notes: LSU, SSU and RPB2 sequence data are available.


Unisetosphaeria Pinnoi et al., Mycoscience 44: 377 (2003)

Asexual morph Undetermined. Sexual morphAscomata immersed to superficial, scattered, pyriform, hyaline to light brown, dark brown near the apex, coriaceous, ostiolate, papillate. Papilla periphysate, surrounded by short dark hairs. Seta single, composed of several rows of brown cells, arising from the ostiolar region. Peridium composed of angular brown-walled cells. Paraphyses sparse, obscure, comprising short rows of ovoid to oblong cells. Asci 8-spored, unitunicate, clavate, short pedicellate, apically truncate, with a refractive, J-, apical ring. Ascospores 2-seriate, septate, hyaline.

Type species: Unisetosphaeria penguinoides Pinnoi et al., Mycoscience 44(5): 378 (2003)

Notes: This monotypic genus was introduced by Pinnoi et al. (2003). Its taxonomic placement was between Chaetosphaeriaceae and Trichosphaeriaceae. However, Unisetosphaeri penguinoides has several incompatible characters of ascomata, paraphyses and asci which are against the Chaetosphaeriaceae. The characters of ascomata, peraphyses, asci and ascospores are more congruent for Trichosphaeriaceae (Pinnoi et al. 2003). Thus, it was suggested to assign this genus in Trichosphaeriaceae based on morphology. This suggestion was followed by Maharachchikumbura et al. (2015, 2016).

Unisetosphaeria penguinoides Pinnoi et al.

Distribution: Thailand, Narathiwat Province, on submerged petiole of Eleiodoxa conferta (Pinnoi et al. 2003).

Asexual morph: Undetermined

Notes: Holotype BBH Aom 103. Sequence data is not available.


Xenospadicoidales Hern.-Restr. et al.

Xenospadicoidaceae Hern.-Restr. et al.

Neospadicoides Z.L. Luo, K.D. Hyde & H.Y. Su, gen. nov.

Index Fungorum number: IF 555652, Facesoffungi number: FoF 05425

Etymology: Referring to the genus similar to Spadicoides.

Saprobic on decaying wood. Asexual morphColonies effuse, hairy, brown to dark brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores macronematous, mononematous, solitary or in groups, erect, unbranched, septate, straight or flexuous, cylindrical, brown, paler towards the apex, smooth. Conidiogenous cells holoblastic, enteroblastic, percurrent, polytretic, integrated, terminal. Conidia acrogenous or acropleurogenous, fusiform, obovoid, septate, smooth-walled. Sexual morph Undetermined.

Type species: Neospadicoides lignicola Z.L. Luo, K.D. Hyde & H.Y. Su

Notes: Neospadicoides is morphologically similar to Spadicoides in having effuse, hairy colonies on natural substrate, mycelium composed of septate, branched, smooth hyphae, conidiophores macronematous, mononematous, erect, straight or flexuous, cylindrical, smooth, conidiogenous cells integrated, terminal and smooth conidia (Hughes 1958; Seifert et al. 2011). However, Neospadicoides differs from Spadicoides in having unbranched conidiophores and acrogenous or acropleurogenous, septate conidia while Spadicoides have branched or unbranched conidiophores, tretic conidiogenous cells and acropleurogenous, aseptate or septate conidia (Hughes 1958; Goh and Hyde 1996a; Seifert et al. 2011). The phylogenetic analysis show that our Neospadicoides species cluster together with good support value and form a distinct subclade from Spadicoides within Xenospadicoidaceae (Xenospadicoidales) (Fig. 1, clade 6).

Neospadicoides aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555654, Facesoffungi number: FoF 05426, Fig. 21

Fig. 21
figure 21

Neospadicoides aquatica (MFLU 18–1605, holotype) ac Conidiophores with conidia. d, e Conidiogenous cells with percurrent proliferations. fk Conidium. Scale bars: ac 40 μm, d, e 25 μm, fk 10 μm

Etymology: Referring to the aquatic habitat of this fungus.

Holotype: MFLU 18–1605

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, brown to dark brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores (80–)126–194(–218) µm long, 5–7 μm wide (\( {\bar{x}} \) = 160 × 6 µm, n = 20), macronematous, mononematous, solitary or in groups, erect, unbranched, septate, straight or flexuous, cylindrical, brown, paler towards the apex, smooth, pecurrently proliferating. Conidiogenous cells holoblastic, integrated, terminal, subhyaline, percurrently proliferating. Conidia 18–22 µm long, 7–9 μm wide (\( {\bar{x}} \) = 20 × 8 µm, n = 20), acrogenous or acropleurogenous, subhyaline to pale brown when young, brown to dark brown when matured, fusiform to cylindrical, rounded at the apex, truncate at the base, 2-septate, with thick and dark band at septa, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2015, X.J. Su, S-701 (MFLU 18–1605, holotype), ex-type living culture MFLUCC 17–2217.

Notes: Neospadicoides aquatica resembles Spadicoides americana in having macronematous, mononematous, erect, unbranched, septate, solitary or in groups conidiophores paler towards the apex, integrated, terminal conidiogenous cells and 2-septate, smooth conidia (Wongsawas et al. 2008). However, Neospadicoides aquatica differs from Spadicoides americana in having larger conidia (18–22 × 7–9 vs. 10–14.5 × 4–6.5 μm) which are rounded at the apex, truncate at the base.

Neospadicoides lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555653, Facesoffungi number: FoF 05427, Fig. 22

Fig. 22
figure 22

Neospadicoides lignicola (MFLU 18–1606, holotype) a, b Conidiophores with conidia. c Conidophore apex with discrete conidiogenous cells. df Conidia. g Germinating conidium. Culture on PDA from above (h) and reverse (i). Scale bars: a, b 50 μm, c 30 μm, dg 20 μm

Etymology: Referring to this fungus dwelling on wood.

Holotype: MFLU 18–1606

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, brown to dark brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores (70–)134–276(–303) µm long, 5–7 μm wide (\( {\bar{x}} \) = 205 × 6 µm, n = 20), macronematous, mononematous, solitary or in groups, erect, unbranched, septate, straight or flexuous, cylindrical, brown, paler towards the apex, smooth. Conidiogenous cells holoblastic, discrete, terminal and intercalary, subhyaline to pale brown. Conidia 20–24 µm long, 9–13 μm wide (\( {\bar{x}} \) = 22 × 11 µm, n = 20), acrogenous or acropleurogenous, pale brown to brown, obovoid, rounded at the apex, truncate at the base, almost 2–3-septate, with dark band at septa, guttulate, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on decaying wood submerged in a freshwater stream, July 2015, X.J. Su, S-735 (MFLU 18–1606, holotype), ex-type living culture MFLUCC 17–2444.

Notes: Neospadicoides lignicola resembles N. yunnanensis in having macronematous, mononematous, solitary or in groups, erect, unbranched, septate, straight or flexuous, cylindrical, brown conidiophores paler towards the apex, integrated, terminal conidiogenous cells and obovoid, septate, guttulate conidia (this study). However, Neospadicoides lignicola differs from N. yunnanensis in having larger conidia (20–24 × 9–13 vs. 7.5–10.5 × 4–6 μm), and conidiogenous cells of N. yunnanensis are with pale-colored pores at the conidiogenous loci. Phylogenetic analysis also shows that N. lignicola and N. yunnanensis are different species (Fig. 1, clade 6). To further support Neospadicoides lignicola as a new species, we compared nucleotide differences with N. yunnanensis following the guidelines of Jeewon and Hyde (2016). Comparison of the 521 nucleotides across the ITS region reveals 40 bp differences (7.67%) including 3 gaps.

Neospadicoides yunnanensis Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555655, Facesoffungi number: FoF 05428, Fig. 23

Fig. 23
figure 23

Neospadicoides yunnanensis (MFLU 18–2329, holotype) a Colonies on substrate. b Conidiophore with conidiogenous cells. c, d Conidiophore with conidia. eh Conidia. Scale bars: c, d 50 μm, b, e 20 μm, fh 5 μm

Etymology: Referring to the location where this fungus was collected, Yunnan Province, China.

Holotype: MFLU 18–2329

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies on decaying wood effuse, pale brown, hairy. Mycelium partly immersed in the substrata, composed of pale brown, septate, smooth, branched hyphae. Conidiophores 113–153 µm long, 4–6 μm wide (\( {\bar{x}} \) = 133 × 5 µm, n = 20), macronematous, mononematous, erect, unbranched, dark brown, paler towards the apex, straight or slightly flexuous, cylindrical, septate, smooth, thick-walled, occasionally swollen at the apex. Conidiogenous cells polytretic, integrated, terminal and intercalary, with pale colored pores remaining at the conidiogenous loci. Conidia 7.5–10.5 µm long, 4–6 μm wide (\( {\bar{x}} \) = 9 × 5 µm, n = 20), acropleurogenous, solitary, pale brown, obovoid, guttulate, mostly 2-septate, comprising a proximal euseptum and a distal distoseptum, with dark band at basal septa, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Lancang River, saprobic on decaying wood submerged, December 2017, X.J. Su, S-1499 (MFLU 18–2329, holotype), ex-type living culture DLUCC 1499.

Notes: Neospadicoides yunnanensis resembles Spadicoides hodgkissa in having macronematous, mononematous, erect, solitary, unbranched, dark brown, paler towards the apex, straight or slightly flexuous, cylindrical, septate, smooth conidiophores occasionally swollen at the apex, polytretic, integrated, terminal and intercalary conidiogenous cells with pale colored pores remaining at the conidiogenous loci and solitary, obovoid, guttulate conidia comprising a proximal euseptum and a distal distoseptum (Ho et al. 2002b). However, Neospadicoides yunnanensis differs from Spadicoides hodgkissa in having pale brown conidia with dark band at basal euseptum while Spadicoides hodgkissa has versicolored conidia comprising a proximal euseptum and a distal distoseptum and the distal distoseptum possesses a conspicuous central pore which is surrounded by a pigmented ring, appearing as dolipores in side view.


Spadicoides S. Hughes, Can. J. Bot. 36: 805 (1958)

Asexual morph Descriptions and illustrations refer to Réblová et al. (2018). Sexual morph Undetermined.

Type species: Spadicoides bina (Corda) S. Hughes, Can. J. Bot. 36: 806 (1958)

Notes: The genus Spadicoides was introduced by Hughes (1958) with S. bina as the type species. Goh and Hyde (1996a) briefly discussed the generic concept of Spadicoides. Thirty-one Spadicoides species were revised, of which 21 species were accepted. Presently, 55 epithets are listed in Index Fungorum (December 2018), and most of the species are reported from terrestrial habitats (Subramanian and Vittal 1974; Kuthubutheen and Nawawi 1991a; Wong et al. 2002; Li 2010; Xia et al. 2014; Ma et al. 2016). Seven species are known from freshwater habitats (Goh and Hyde 1996a; Ho et al. 2002b; Cai et al. 2004a, 2006a, b, c; Zhuang 2001).

Spadicoides americana C.J.K. Wang

Distribution: China, Zhejiang Province, Hangzhou city, on submerged wood in a stream at Nine Creeks (Wongsawas et al. 2008).

Sexual morph: Undetermined

Notes: Specimen collected from freshwater habitats HMZFW 00002. Sequence data is not available.

Spadicoides atra (Corda) S. Hughes

Chloridium atrum Corda, Icones fungorum hucusque cognitorum 1: 17, t. 4:235 (1837)

Psilonia atra (Corda) Corda, Icones fungorum hucusque cognitorum 4: 27, t. 6:84 (1840)

Catenularia atra (Corda) Sacc., Sylloge Fungorum 4: 304 (1886)

Distribution: China, Hong Kong, on submerged wood (Zhuang 2001).

Sexual morph: Undetermined

Notes: LSU, SSU and RPB2 sequence data are available.

Spadicoides bambusicola Zhou et al.

Distribution: China, Yunnan Province, on submerged wood (Cai et al. 2006a).

Sexual morph: Undetermined

Notes: Holotype HKU (M) 8333 (now in IFRD). Sequence data is unavailable.

Spadicoides cordanoides Goh & K.D. Hyde

Distribution: Australia, Queensland, on submerged wood (Goh and Hyde 1996a).

Sexual morph: Undetermined

Notes: Holotype BRIP 23201. Sequence data is not available. This species is known only from Australia on submerged wood in freshwater habitats (Goh and Hyde 1996a).

Spadicoides hodgkissa Ho et al.

Distribution: China, Hong Kong, Tai Po Kau Forest Stream (Ho et al. 2002b)

Sexual morph: Undetermined

Notes: Holotype HKU(M) 6155 (now in IFRD). Sequence data is unavailable.

Spadicoides minuta Cai et al.

Distribution: China, Yunnan Province, on Phyllostachys bambusoides submerged in a small stream (Cai et al. 2004a).

Sexual morph: Undetermined

Notes: Holotype IFRD 8850. Sequence data is unavailable.

Spadicoides obovata (Cooke & Ellis) S. Hughes

Acrothecium obovatum Cooke & Ellis, Grevillea 5 (34): 50 (1876)

Spondylocladium obovatum (Cooke & Ellis) S. Hughes, Can. J. Bot. 31 (5): 634 (1953)

Distribution: China, Hong Kong, on submerged wood (Zhuang 2001).

Sexual morph: Undetermined

Notes: Sequence data is unavailable.


Torrentispora Hyde et al., Mycol. Res. 104(11): 1399 (2000)

Asexual morph Undetermined. Sexual morph Updated description refer to Réblová et al. (2018).

Type species: Torrentispora fibrosa Hyde et al., Mycol. Res. 104(11): 1399 (2000)

Notes: Hyde et al. (2000) introduced the genus Torrentispora with single species, T. fibrosa. Réblová et al. (2018) revisited Torrentispora with nine species in this genus and assigned it to the new family Xenospadicoidaceae within the new order Xenospadicoidales based on morphology and DNA sequence data. Torrentispora comprises seven species from freshwater habitats.

Torrentispora aquatica (Vijaykr. et al.) Réblová & A.N. Mill

Fusoidispora aquatica Vijaykr. et al., Sydowia 57:272 (2005)

Distribution: China, Hong Kong, New Territories, Tai Po Kau Country Park, Tai Po Kau Forest Stream, on submerged wood (Vijaykrishna et al. 2005).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 17484 (now in IFRD), isotype PDD 78746. LSU sequence data is available.

Torrentispora biatriispora (K.D. Hyde) Réblová & A.N. Mill.

Annulatascus biatriisporus K.D. Hyde, Nova Hedwigia 61: 120 (1995)

Pseudoannulatascus biatriisporus (K.D. Hyde) Z.L. Luo et al., Phytotaxa 239:179 (2015)

Distribution: Australia, on submerged wood (Hyde 1995b); China, Hong Kong, Tsuen Wan, Shing Mun Reservoir, on submerged wood (Tsui et al. 2002); Costa Rica, on submerged wood (Barbosa et al. 2013); Seychelles, on submerged wood (Hyde and Goh 1998a).

Asexual morph: Undetermined

Notes: Holotype BRIP 21481, other specimen collected from freshwater habitats: ILL 40816. ITS, LSU, SSU and RPB2 sequence data are available.

Torrentispora calembola Réblová & A.N. Mill.

Distribution: France, on submerged wood of Fraxinus excelsior (Réblová et al. 2018).

Asexual morph: Undetermined

Notes: Holotype PRA-12744 (M.R. 3726), isotype PRA-12745. ITS, LSU and SSU sequence data are available.

Torrentispora crassiparietis Fryar & K.D. Hyde

Distribution: Brazil, on submerged bark (Barbosa et al. 2013); Brunei, on submerged wood (Fryar and Hyde 2004; Fryar et al. 2004); Costa Rica, on submerged wood (Barbosa et al. 2013).

Asexual morph: Undetermined

Notes: Holotype IFRD 8863, other specimens collected from freshwater habitats: HUEFS 158094, ILL 40817, ILL 40818, ILL 40819, ILL 40820. Sequence data is unavailable.

Torrentispora fibrosa Hyde et al.

Distribution: China, Hong Kong, New Territories, Tai Po, Lam Tsuen River and Tai Po Kau Forest Reserve, Tai Po Kau Forest Stream, on submerged decaying wood (Hyde et al. 2000); New Zealand, Lake Christabel track, on decaying wood of Nothofagus sp. (Réblová et al. 2018).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 4519 (now in IFRD), other specimen collected from freshwater habitats PDD 110879. LSU, SSU and RPB2 sequence data are available.

Torrentispora fusiformis Fryar & K.D. Hyde

Distribution: Brunei, on submerged wood (Fryar and Hyde 2004).

Asexual morph: Undetermined

Notes: Holotype IFRD 8864. Sequence data is not available. This species is known only from Brunei on submerged wood in brackish and freshwater habitats (Fryar and Hyde 2004).

Torrentispora pilosa Shearer & F.R. Barbosa

Distribution: Costa Rica, on submerged wood (Barbosa et al. 2013).

Asexual morph: Undetermined

Notes: Holotype ILL 40814. Sequence data is not available. This species is known only from Costa Rica, on submerged wood (Barbosa et al. 2013).


Diaporthomycetidae family incertae sedis

Acrodictyaceae J.W. Xia & X.G. Zhang

Acrodictys M.B. Ellis, Mycol. Pap. 79: 6 (1961)

Asexual morph Descriptions and illustrations refer to Xia et al. (2017). Sexual morph Undetermined.

Type species. Acrodictys bambusicola M.B. Ellis, Mycol. Pap. 79: 6 (1961)

Notes: Acrodictys was introduced by Ellis (1961), with A. bambusicola M.B. Ellis as the type species. There are 45 epithets listed in Index Fungorum (December 2018). Until 2016, identification of Acrodictys-like species was based on morphology. Xia et al. (2017) provided ITS, LSU, SSU and β-tubulin sequence data, detailed descriptions and illustrations for eight Acrodictys species, including a new species. In this study, we introduce a new species and a new combination for this genus. Three Acrodictys species have been reported from freshwater habitats.

Acrodictys aquatica (N.G. Liu & K.D. Hyde) Z.L. Luo & K.D. Hyde, comb. nov.

Barbatosphaeria aquatica N.G. Liu & K.D. Hyde, in Hyde et al., Mycosphere 9(2): 384 (2018)

Index Fungorum: IF 655656; Facesoffungi number: FoF 05438

Descriptions and illustrations refer to Hyde et al. (2018).

Distribution: Thailand, Chiang Rai Province, Muang, Ban Nang Lae Nai, on decaying wood submerged in a freshwater stream (Hyde et al. 2018).

Sexual morph: Undetermined

Notes: Holotype MFLU 18–0040. ITS and LSU sequence data are available. Hyde et al. (2018) introduced the species Barbatosphaeria aquatica based on a collection obtained from freshwater habitat in Thailand. Our phylogenetic analysis based on multi-genes shows that the strain from ex-type culture of Barbatosphaeria aquatica clusters in the genus Acrodictys (Fig. 1, clade 9). Morphology of Barbatosphaeria aquatica also fits well with Acrodictys species in having mononematous, erect, unbranched conidiophores; monoblastic, terminal, integrated conidiogenous cells; solitary, dry, clavate, green to brown conidia, with transverse septa, deeply constricted at the septa, distinctly verruculose (Hyde et al. 2018). Based on morphology and phylogenetic analyses, we synonymize Barbatosphaeria aquatica under Acrodictys aquatica.

Acrodictys fluminicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555657, Facesoffungi number: FoF 05429, Fig. 24

Fig. 24
figure 24

Acrodictys fluminicola (DLU 0274, holotype) a, b Colonies on natural substrate. c, d Conidiophores with conidia. e, f Conidiophores with conidiogenous cells. go Conidia. p Germinating conidium. Culture on MEA, q from above, r from reverse. Scale bars: cf 50 μm, gp 15 μm

Etymology: Referring to this fungus dwelling in a stream.

Holotype: DLU 0274

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, dark brown to black. Conidiophores 98–142 µm long, 4–6 μm wide (\( {\bar{x}} \) = 120 × 5 µm, n = 20), macronematous, mononematous, erect, unbranched, straight or flexuous, septate, dark brown at the base, narrower and paler toward the apex, smooth, thick-walled. Conidiogenous cells monoblastic, integrated, terminal, cylindrical, lageniform to doliiform, subhyaline to pale brown, smooth. Conidia 24–30 µm long, 13–17 μm wide (\( {\bar{x}} \) = 27 × 15 µm, n = 20), acrogenous, solitary, muriform, broadly clavate, obovoid to pyriform, usually with 2–3 transverse septa and a few longitudinal septa, with conspicuous pores in each cells, slightly constricted at the septa, pale brown at the basal cells and brown at the other part, basal cell obconical, truncate at base. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, October 2014, L.W. Wang, S-274 (DLU 0274, holotype), ex-type living culture KUMCC 15–0240.

Notes: Acrodictys fluminicola resembles the generic type A. bambusicola in having macronematous, unbranched and septate conidiophores dark brown at the base, narrower and paler toward the apex, solitary, muriform, obovoid to pyriform conidia slightly constricted at the septa, with obconical basal cell and the size of conidiophores and conidia of these two species are almost similar (Xia et al. 2017). However, A. fluminicola differs from A. bambusicola in has 2–3 transverse septa conidia with conspicuous guttulae in each cell, while A. bambusicola have 3–4 transverse septa. Phylogenetic analysis also shows that Acrodictys fluminicola and A. bambusicola are distinct species (Fig. 1, clade 9). Phylogenetically, A. aquatica clusters with A. hainnensis (Fig. 1, clade 9), however, Acrodictys fluminicola morphologically differs from A. hainnensis in having longer conidiophores (98–142 vs. up to 35 µm) and larger conidia (24–30 × 13–17 vs. 15–22 × 7–13 µm).

Acrodictys liputii Cai et al.

Distribution: Philippines, Negros Occidental, Bario Alegria, Liput River, on submerged bamboo culm (Cai et al. 2002b).

Sexual morph: Undetermined

Notes: Holotype PDD 74789, isotype IFRD 8640. ITS, LSU and SSU sequence data are available.


Barbatosphaeriaceae Zhang et al.

Barbatosphaeria Réblová, Mycologia 99(5): 727 (2008) [2007]

Asexual morph Undetermined. Sexual morph Description and illustrations refer to Réblová et al. (2015a).

Type species: Barbatosphaeria barbirostris (Dufour) Réblová, Mycologia 99(5): 727 (2008)

Notes: Barbatosphaeria is a non-stromatic perithecial ascomycete introduced for a single species Barbatosphaeria barbirostris and placed in the Sordariomycetes genera incertae sedis based on nuc28S rDNA sequences (Réblová 2007). Réblová et al. (2015a) revisited Barbatosphaeria and introduced seven species for this genus. In this study, we introduce one new species and this is the only known Barbatosphaeria species collected from freshwater habitats.

Barbatosphaeria lignicola Z.L. Luo, H.Y. Su & K.D. Hyde, sp. nov.

Index Fungorum number: IF 555658, Facesoffungi number: FoF 05430, Fig. 25

Fig. 25
figure 25

Barbatosphaeria lignicola (HKAS 84005, holotype) a Specimen. b, c Appearance of necks on substrate. d Horizontal section through ascomata. e, f Section through ascomata. g, h Structure of peridium. i Paraphyses. jn Ascus. op Ascospore. Scale bars: e, f 150 μm, gi 30 μm, jp 5 μm

Etymology: Referring to this fungus dwelling on wood

Holotype: HKAS 84005

Saprobic on decaying wood submerged in freshwater habitats. Asexual morph Undetermined. Sexual morphAscomata immersed, globose to subglobose, dark brown to black, 380–400 μm high, 270–320 μm diam., aggregated in circular groups, occasionally solitary. Neck black, cylindrical, straight to slightly flexuous, converging radially. Peridium 25–40 μm thick, leathery to fragile, 2-layered. Outer layer consisting of thick-walled, brown, polyhedral to elongate cells of textura prismatica to textura angularis, towards the interior grading into several layers of thin-walled pale brown to subhyaline flattened cells. Paraphyses septate, slightly constricted at the septa, wider near the base, tapering toward the tip, longer than asci. Asci 19–31 × 4–6 μm (\( {\bar{x}} \) = 25 × 5 µm, n = 30), 8-spored, unitunicate, clavate in sporiferous part, tapering toward the stipe, ascal apex broadly rounded to obtuse. Ascospores 5–7 × 1–2 μm (\( {\bar{x}} \) = 6 × 1.5 μm, n = 20), arranged obliquely 1–2-seriately in the sporiferous part of the ascus, allantoid, U- to horseshoe shaped, narrow to pinted at both ends, aseptate, non-constricted, hyaline.

Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in Heilong stream, 15 March 2014, Z.L. Luo, S-105, (HKAS 84005, holotype).

Notes: Barbatosphaeria lignicola mostly resembles B. hippocrepida in having globose to subglobose, dark brown to black ascomata, black, cylindrical neck, septate paraphyses wider near the base, tapering toward the tip, longer than asci and slightly constricted at the septa, unitunicate, 8-spored asci and allantoid, U- to horseshoe shaped, hyaline, aseptate, non-constricted, similar size of ascospores (Réblová and Štěpánek 2009). However, B. lignicola differs from B. hippocrepida in having smaller ascomata (380–400 × 270–320 vs. 350–500 × 400–650 μm), ascospores tending to group in the apex of the sporiferous part while ascospores of B. hippocrepida tend to group in the middle of the sporiferous part, leaving the apex empty. Phylogenetic analyses show that Barbatosphaeria lignicola is distinct from other Barbatosphaeria species (Fig. 26).

Fig. 26
figure 26

Phylogram generated from maximum likelihood analysis based on combined ITS, LSU and RPB2 sequence data for species of Barbatosphaeria (with Natantiella ligneola as outgroup). The best scoring RAxML tree with a final likelihood value of − 9327.825160 is presented. RAxML bootstrap support values equal to or greater than 75% are given before the forward slash (black). Maximum parsimony bootstrap support values equal to or greater than 75% are given after the forward slash (red). Hyphen (‘-’) indicates a value lower than 75% for RAxML and maximum parsimony. Newly generated sequences are in red. Ex-type strains are in bold


Conlariaceae Zhang et al.

Conlarium F. Liu & L. Cai, Mycologia 104(5): 1180 (2012)

Asexual morph Hyphomycetous, for description and illustration refer to Zhang et al. (2017a, b). Sexual morph Description and illustration refer to Liu et al. (2012).

Type species: Conlarium dupliciascospora F. Liu & L. Cai, Mycologia 104(5): 1180 (2012)

Notes: Currently, two species are accepted in the genus Conlarium, and both C. aquaticum and C. dupliciascospora are reported from freshwater habitats in China and Thailand respectively (Liu et al. 2012; Zhang et al. 2017a, b).

Conlarium aquaticum Dong et al.

Distribution: Thailand, Prachuap Khiri Khan, on submerged wood in a small River (Zhang et al. 2017).

Sexual morph: Undetermined

Notes: Holotype MFLU 15–2703. ITS, LSU, SSU and TEF1α sequence data obtained from ex-type culture are available.

Conlarium dupliciascosporum F. Liu & L. Cai

Distribution: China, Guangdong Province, Zhaoqing, Dinghu Mountain, on submerged wood in a stream (Liu et al. 2012).

Asexual morph: See Liu et al. (2012)

Notes: Holotype HMAS 243129. ITS, LSU and SSU sequence data are available, ex-type strain CGMCC3.14938.


Riomyces Ferrer et al., Mycologia 104(4): 876 (2012)

Asexual morph Undetermined. Sexual morph Descriptions and illustration refer to Ferrer et al. (2012).

Type species: Riomyces rotundus A. Ferrer, A.N. Mill., Sarmiento & Shearer, Mycologia 104(4): 876 (2012)

Notes: There is only one species was accepted in the genus Riomyces and was only known from the type locality.

Riomyces rotundus Ferrer et al.

Distribution: Costa Rica, Alajuela and Heredia, on submerged wood (Ferrer et al. 2012).

Asexual morph: Undetermined

Notes: Holotype ILL AF303–1, other specimen collected from freshwater habitats: ILL AF303–3. LSU and SSU sequence data are available.


Junewangiaceae J.W. Xia & X.G. Zhang

Dictyosporella Abdel-Aziz, Fungal Divers 75: 144 (2015)

Asexual morph Description and illustrations see Ariyawansa et al. (2015). Sexual morph Description and illustrations see Zhang et al. (2017a, b).

Type species: Dictyosporella aquatica Abdel-Aziz, Fungal Divers 75: 145 (2015)

Notes: Ariyawansa et al. (2015) introduced the genus Dictyosporella to accommodate the asexual morph species D. aquatica Abdel-Aziz, collected on submerged decaying stems of Phragmites australis in Nile River, Egypt, and accommodated it in family Annulatascaceae. Zhang et al. (2017a, b) introduced a sexual morph species Dictyosporella thailandensis W. Dong, H. Zhang & K.D. Hyde for this genus which was collected from freshwater, and they moved this genus from Annulatascaceae to Diaporthomycetidae genera incertae sedis. Song et al. (2018a) introduced the third species Dictyosporella hydei H.Y. Song & D.M. Hu based on morphology and phylogenetic analyses, but the placement of this genus was not mentioned in their study. Our phylogenetic analysis based on multi-gene shows that the genus Dictyosporella clusters in Junewangiaceae with strong support (Fig. 1, clade 1). According to our phylogenetic result, Dictyosporella is transferred from Diaporthomycetidae genera incertae sedis to Junewangiaceae. Three species were accepted in this genus and all were collected from freshwater habitats.

Dictyosporella aquatica Abdel-Aziz

Distribution: Egypt, Sohag, River Nile, on submerged decayed stem of Phragmites australis (Ariyawansa et al. 2015).

Sexual morph: Undetermined

Notes: Holotype CBS H-22127, isotype MFLU 15–1510. LSU and SSU sequence data are available.

Dictyosporella hydei H.Y. Song & D.M. Hu

Distribution: China, Yunnan Province, Mengla, on submerged wood in a small stream (Song et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype IFRD 021–044. LSU sequence data is available. This species is only known from the type locality.

Dictyosporella thailandensis Dong et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged bamboo in a small River (Zhang et al. 2017a, b).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2706. ITS, LSU, SSU and RPB2 sequence data are available.


Junewangia W.A. Baker & Morgan-Jones, Mycotaxon 81: 307 (2002)

Asexual morph Description and illustrations see Xia et al. (2018). Sexual morph Undetermined.

Type species: Junewangia sphaerospora W.A. Baker & Morgan-Jones, Mycotaxon 81: 312 (2002)

Notes: Junewangia was established by Baker et al. (2002) to accommodate four species of Acrodictys M.B. Ellis. The species of Junewangia are characterized by slightly flared conidiophores with annellidic, percurrent proliferation, cylindrical or narrowly cuneate conidiogenous cells, subglobose to globose conidia with a prominent, protruding basal cell, uniformly pigmented, and a series of largely angular or oblique septa, with schizolytic secession. Seven species are accepted in the genus Junewangia and only one of them was collected from freshwater habitats in China (Song et al. 2018b).

Junewangia aquatica H.Y. Song & D.M. Hu

Distribution: China, Yunnan Province, Mengla, stream in rubber trees field, on submerged wood (Song et al. 2018b).

Sexual morph: Undetermined

Notes: Holotype HFJAU 0700; ITS, LSU, SSU sequence data are available. This species is only known from the type locality.


Sporidesmiella P.M. Kirk

Asexual morph Descriptions and illustrations see Wu and Zhuang (2005); Sexual morph Undetermined.

Type species: Sporidesmiella claviformis P.M. Kirk, Trans. Br. Mycol. Soc. 79: 479 (1982)

Notes: Kirk (1982) introduced the genus Sporidesmiella P.M. Kirk to accommodate the species previously described in the genera Sporidesmium and Endophragmia. Sporidesmiella resembles Repetophragma but differs in its distoseptate conidia (Kirk 1982; Subramanian 1992). In our phylogenetic analysis, Sporidesmiella clusters in Junewangiaceae while Repetophragma belongs in Pseudosporidesmiaceae (Fig. 1, clade 1, 37). In this study, we report three Sporidesmiella species which were collected from freshwater habitats with one new taxon.

Sporidesmiella aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555659, Facesoffungi number: FoF 05431, Fig. 27

Fig. 27
figure 27

Sporidesmiella aquatica (MFLU 18–1602, holotype) a Colony on natural substrate. b, c Conidiophore with conidia. d, e Conidiogenous cells with conidia. fh Conidia. i Germinating conidium. Culture on PDA from above (j) and reverse (k). Scale bars: b, c 100 μm, di 50 μm

Etymology: Referring to the aquatic habitat of this fungus

Holotype: MFLU 18–1602

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, hairy, yellow brown to brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores 178–228 µm long, 8–10 μm wide (\( {\bar{x}} \) = 203 × 9 µm, n = 20), macronematous, mononematous, erect, unbranched, septate, straight or flexuous, cylindrical, yellow–brown, paler towards the apex, smooth. Conidiogenous cells holoblastic, polyblastic, sympodial, integrated, terminal, subterminal, subhyaline to pale brown, with percurrent or sympodial proliferations. Conidia 51–59 µm long, 18–22 μm wide (\( {\bar{x}} \) = 55 × 20 µm, n = 20), acropleurogenous, dry, clavate or obovoid, rounded at the apex, truncate at the base, 3–4-distoseptate, pale brown, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, October 2016, Z.L. Luo, S-777 (MFLU 18–1602, holotype), ex-type living culture DLUCC 0777; Laojunshan Mountain, saprobic on decaying wood submerged in a freshwater stream, August 2017, H.W. Shen, S-1339 (MFLU 18–2331, paratype), living culture DLUCC 1339.

Notes: Sporidesmiella aquatica resembles S. hyalosperma in having macronematous, mononematous, erect, unbranched, septate, cylindrical conidiophores, integrated, terminal conidiogenous cells with percurrent or sympodial proliferations and acrogenous, dry, clavate or obovoid conidia (Kirk 1982; Wu and Zhuang 2005). However, S. aquatica differs from S. hyalosperma in having polyblastic conidiogenous cells and larger conidia (51–59 × 18–22 vs. 22–25 × 9–12 μm).

Sporidesmiella novae-zelandiae (S. Hughes) Madrid et al.

Sporidesmium hyalospermum var. novae-zelandiae S. Hughes, N.Z. Jl Bot. 16(3): 349 (1978)

Sporidesmiella hyalosperma var. novae-zelandiae (S. Hughes) P.M. Kirk, Trans. Brit. Mycol. Soc. 79: 479 (1982)

Exserohilum novae-zelandiae (S. Hughes) H.P. Upadhyay & Mankau, Mycologia 83: 373 (1991)

Facesoffungi number: FoF 05432, Fig. 28

Fig. 28
figure 28

Sporidesmiella novae-zelandiae (MFLU 18–1604) a Colony on natural substrate. b, c Conidiophore with conidia. d, e Conidiogenous cells with conidia. fi Conidia. j Germinating conidium. Culture on PDA from surface (k) and reverse (l). Scale bars: b, c 30 μm, di 15 μm

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, hairy, yellow brown to brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores 132–186 µm long, 5–7 μm wide (\( {\bar{x}} \) = 159 × 6 µm, n = 20), macronematous, mononematous, erect, unbranched, septate, straight or flexuous, cylindrical, yellow–brown, paler towards the apex, smooth. Conidiogenous cells holoblastic, polyblastic, sympodial, integrated, terminal, subhyaline to pale brown. Conidia 25–29 µm long, 12–14 μm wide (\( {\bar{x}} \) = 27 × 13 µm, n = 20), acrogenous, dry, pale brown, clavate or obovoid, rounded at the apex, truncate at the base, 2–4-distoseptate, basal cell cut off by a dark brown septum, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Dulong River, saprobic on decaying wood submerged, October 2016, Z.L. Luo, S-951 (MFLU 18–1604), living culture DLUCC 0951; Laojunshan Mountain, saprobic on decaying wood submerged in a freshwater stream, August 2017, H.W. Shen, S-1256 (MFLU 18–2332), living culture DLUCC 1256.

Notes: Holotype PDD 30420, isotype DAOM 159962, other specimen collected from freshwater habitats: MFLU 18–1604. We provide the ITS, LSU, SSU, RPB2 and TEF1α sequence data for this species. The species Sporidesmiella novae-zelandiae was originally described as Sporidesmium hyalospermum var. novae-zelandiae by Hughes (1978). This species and the type variety were reallocated to Sporidesmiella by Kirk (1982) and Sporidesmium hyalospermum var. novae-zelandiae was later transferred to Exserohilum as E. novae-zelandiae by Upadhyay and Mankau (1991). Hernández-Restrepo et al. (2018) considered that Exserohilum novaezelandiae should be retained in Sporidesmiella and combined it as S. novae-zelandiae. During an investigation on lignicolous freshwater fungi in China, a Sporidesmiella-like fungus was collected from Yunnan Province and the morphological characters of our new collection fits well with S. novae-zelandiae (Hughes 1978; Kirk 1982; Wu and Zhuang 2005; Hernández-Restrepo et al. 2018). Phylogenetic analysis shows that our isolates cluster with other Sporidesmiella species with good bootstrap support value and form in a distinct clade (Fig. 1, clade 1). Based on the morphology and phylogeny, we identify our isolates as Sporidesmiella novae-zelandiae.

Sporidesmiella hyalosperma (Corda) P.M. Kirk

Helminthosporium hyalospermum Corda, Icon. fung. (Prague) 1: 13 (1837)

Facesoffungi number: FoF 05433, Fig. 29

Fig. 29
figure 29

Sporidesmiella hyalosperma (MFLU 18–1603) a Colony on natural substrate. b Conidiophores with conidia. c Conidiogenous cells. d, e Conidia. f, g Germinating conidia. Culture on PDA from above (h) and reverse (i). Scale bars: b 50 μm, cg 20 μm

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies effuse, hairy, yellow brown to brown. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth, pale brown hyphae. Conidiophores 116–144 µm long, 5–7 μm wide (\( {\bar{x}} \) = 130 × 6 µm, n = 20), macronematous, mononematous, erect, unbranched, septate, straight or flexuous, cylindrical, brown, paler towards the apex, smooth. Conidiogenous cells holoblastic, polyblastic, sympodial, integrated, terminal, later becoming subterminal, subhyaline to pale brown, with percurrent or sympodial proliferations. Conidia 17–21 µm long, 8–10 μm wide (\( {\bar{x}} \) = 19 × 9 µm, n = 20), acropleurogenous, dry, clavate or obovoid, rounded at the apex, truncate at the base, 3–4-distoseptate, pale brown, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Jinsha River, saprobic on decaying wood submerged, April 2015, Z.L. Luo, S-563 (MFLU 18–1603, HKAS 92987), living culture KUMCC 15–0431; Cangshan Mountain, saprobic on decaying wood submerged in a freshwater stream, August 2017, H.W. Shen, S-1400, living culture MFLUCC 18–1312; Ibid., saprobic on decaying wood submerged in a freshwater stream, August 2017, H.W. Shen, S-1518 (MFLU 18–2330), living culture DLUCC 1518.

Notes: Holotype PDD 30420, specimen collected from freshwater habitats: MFLU 18–1603. We provide the ITS, LSU, SSU, RPB2 and TEF1α sequence data for this species in this study. Sporidesmiella hyalosperma is the most common species in the genus Sporidesmiella. Morphologically, our isolates fit well with the Sporidesmiella hyalosperma in having macronematous, mononematous, solitary, erect, unbranched, septate, straight or flexuous, cylindrical, brown, smooth conidiophores paler towards the apex, holoblastic, integrated, terminal, subhyaline to pale brown conidiogenous cells with percurrent or sympodial proliferations and acrogenous, dry, clavate or obovoid, distoseptate conidia rounded at the apex, truncate at the base and even the size of conidiophores and conidia are also similar (Kirk 1982; Wu and Zhuang 2005). Therefore, we identify our collections as Sporidesmiella hyalosperma, and it is the first time to report this species from freshwater.


Papulosaceae Winka & O.E. Erikss.

Brunneosporella Ranghoo & K.D. Hyde, Mycol. Res. 105(5): 625 (2001)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Ranghoo et al. (2001).

Type species: Brunneosporella aquatica Ranghoo & K.D. Hyde, Mycol. Res. 105(5): 625 (2001)

Notes: Ranghoo et al. (2001) introduced the genus Brunneosporella with single species B. aquatica which was collected from freshwater habitats. This species is only known from the type locality.

Brunneosporella aquatica Ranghoo & K.D. Hyde

Distribution: China, Hong Kong, New Territories, Plover Cove Reservoir, on submerged wood (Ranghoo et al. 2001).

Asexual morph: Undetermined

Notes: Holotype IFRD 8717. ITS and LSU sequence data are available.


Fluminicola Wong et al., Fungal Diversity Res. Ser. 2: 190 (1999)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations refer to Wong et al. (1999b) and Zhang et al. (2017a, b).

Type species: Fluminicola bipolaris Wong et al., Fungal Divers 2: 190 (1999)

Notes: Wong et al. (1999b) established the genus Fluminicola with F. bipolaris as type species. Zhang et al. (2017a, b) introduced three species which were collected from a freshwater River in southern Thailand.

Fluminicola aquatica Dong et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged wood in a small River (Zhang et al. 2017a, b).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2710. ITS, LSU and SSU sequence data are available.

Fluminicola bipolaris Wong et al.

Distribution: Philippines, Mindanao, Bukidnon, Impalutao, Natigbasan Creek, on submerged wood (Wong et al. 1999b).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 3127 (now in IFRD). Sequence data is not available.

Fluminicola saprophytica Dong et al.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged bamboo in a small River (Zhang et al. 2017a, b).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2694. ITS, LSU, SSU and RPB2 sequence data are available.

Fluminicola thailandensis Dong et al.

Facesoffungi number: FoF 03346, Fig. 30

Fig. 30
figure 30

Fluminicola thailandensis (MFLU 15–0085) a Specimen. b, c Appearance of necks on substrate. d Vertical section of ascomata. e Structure of peridium. f Paraphyses. gi Ascus. jm Ascospores. n Germinating ascospore. o Colony on PDA. Scale bars: d, gi 40 μm, e, f, n 20 μm, jm 10 μm

Saprobic on decaying wood submerged in freshwater habitats. Asexual morph Undetermined. Sexual morphAscomata 138–156 μm high, 100–120 μm diam., immersed with neck erumpent through host surface, subglobose to ellipsoid, solitary, dark brown to black. Ostiole central, with straight upright, black neck. Peridium 12–18 μm thick, consisting two-layers, outer layer comprising 3–4 layers of dark brown, thick-walled cells of textura angularis, inner layer comprising 2–3 layers of pale brown to hyaline, thin-walled cells of textura angularis. Paraphyses 4–6 μm wide, hyaline, unbranched, septate, slightly constricted at the septum. Asci 135–181 × 11–15 µm (\( {\bar{x}} \) = 158 × 13 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, tapering to a point, apically rounded or slightly obtuse, with an indistinct, refractive, small, discoid apical ring. Ascospores 19–21 × 7–9 µm (\( {\bar{x}} \) = 20 × 8 µm, n = 20), uniseriate, fusiform, 3-septate, slightly constricted at the septa, with 2 prominent guttules, with a thin mucilaginous sheath, hyaline, smooth-walled.

Material examined: THAILAND, Chiang Rai Province, saprobic on decaying wood submerged in a freshwater stream, November 2013, Z.L. Luo, ZL-10 (MFLU 15–0085), living culture MFLUCC 14–0037.

Distribution: Thailand, Prachuap Khiri Khan Province, on submerged wood in a small River (Zhang et al. 2017a, b), Chiang Rai Province, saprobic on decaying wood submerged in a freshwater stream (This study).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2704, other specimen collected from freshwater MFLU 15–0085. ITS, LSU, SSU and TEF1α sequence data are available. Fluminicola thailandensis was introduced by Zhang et al. (2017a, b) based on an isolate collected from southern Thailand. During our investigation of lignicolous freshwater fungi in northern Thailand, an Annulatascaceae-like fungus was collected from Chiang Rai Province. Morphologically, the new isolate resembles Fluminicola species in having immersed ascomata with black neck, cylindrical asci with apical ring and fusiform, septate, guttulate ascospores with a thin mucilaginous sheath. Phylogenetic analysis based on LSU, SSU, RPB2 and TEF1α sequence data shows that the new strain (MFLUCC 14–0037) clusters with Fluminicola species with good support, and close to F. thailandensis and F. saprotrophitica. We compared the ITS sequences and noted that there are 3 nucleotide differences (including 2 gaps) between the ex-type strain of Fluminicola thailandensis (MFLUCC 15–0984) and our strain (MFLUCC 14–0037). Based on morphology and phylogeny, we identify our species as Fluminicola thailandensis.


Wongia Khemmuk et al., IMA Fungus 7(2): 249 (2016)

Asexual morphColonies effuse, scattered, hairy, dark brown. Mycelium partly immersed, composed of branched, septate, brown, smooth hyphae. Conidiophores macronematous, mononematous, solitary, erect, straight or flexuous, unbranched, septate, dark brown, smooth. Conidiogenous cells polyblastic, denticulate, integrated, terminal, subhyaline. Conidia acrogenous, clavate to fusiform, 3-septate, guttulate, dark brown at central two cells, paler at end cells, rounded and narrow at apex, truncate at base. Sexual morph Descriptions refer to Khemmuk et al. (2016).

Type species: Wongia garrettii (P. Wong & M.L. Dickinson) Khemmuk et al., IMA Fungus 7(2): 250 (2016)

Notes: Khemmuk et al. (2016) re-examined Magnaporthe garrettii and M. griffinii that are pathogenic on roots of couch and hybrid couch (Wong et al. 2012) and established the genus Wongia. Wongia is the fourth genus to be placed in Papulosaceae, along with Brunneosporella (Ranghoo et al. 2001), Fluminicola (Wong et al. 1999a, b) and Papulosa (Kohlmeyer and Volkmann-Kohlmeyer 1993). In this study, we introduce an asexual Wongia species which was collected from freshwater habitats in northwestern Yunnan Province, China.

Wongia aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555660, Facesoffungi number: FoF 05434, Fig. 31

Fig. 31
figure 31

Wongia aquatica (MFLU 18–1596, holotype) ac Conidiophore with conidia. d Conidiogenous cells. ei Conidia. j Germinating conidium. Scale bars: ac 45 μm, dj 20 μm

Etymology: Referring to the aquatic habitat of this fungus

Holotype: MFLU 18–1596

Saprobic on submerged decaying wood. Asexual morphColonies effuse, scattered, hairy, dark brown. Mycelium partly immersed, composed of branched, septate, brown, smooth hyphae. Conidiophores 54–90 µm long, 3–6 μm wide (\( {\bar{x}} \) = 72 × 4 µm, n = 20), macronematous, mononematous, solitary, erect, straight or flexuous, unbranched, septate, dark brown, smooth. Conidiogenous cells polyblastic, sympodial, denticulate, integrated, terminal, subhyaline, with pale brown scar. Conidia 17–21 µm long, 5–7 μm wide (\( {\bar{x}} \) = 19 × 6 µm, n = 20), acropleurogenous, fusiform, guttulate, 3-septate, dark brown at central two cells, paler at end cells, rounded and narrow at apex, truncate at base, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Dulong River, saprobic on decaying wood submerged, October 2016, Z.L. Luo, S-912 (MFLU 18–1596, holotype), ex-type living culture MFLUCC 18–1607.

Notes: There are no asexual morphs reported for Wongia garrettii and W. griffinii. Wongia aquatica is the first asexual morph species in the genus Wongia. Phylogenetically, W. aquatica shares a sister relationship to W. griffinii with high bootstrap support (96% ML) (Fig. 1, clade 11).


Pseudoproboscisporaceae Zhang et al.

Diluviicola Hyde et al., Fungal Diversity Res. Ser. 1: 141 (1998)

Asexual morph Undetermined. Sexual morph Description and illustrations see Hyde et al. (1998b).

Type species: Diluvicola capensis K.D. Hyde, S.W. Wong & E.B.G. Jones, Fungal Divers 1: 134 (1998)

Notes: Hyde et al. (1998b) introduced the genus Diluviicola based on a collection from Brunei. Zhang et al. (2017a, b) introduced the second species in this genus, Diluviicola aquatica W. Dong, H. Zhang & K.D. Hyde, based on a collection made from submerged bamboo in a river in Thailand.

Diluviicola aquatica Dong et al.

Distribution: Thailand, Prachuap Khiri Khan, on submerged bamboo in a small River (Zhang et al. 2017a, b).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2701. ITS, LSU, SSU and RPB2 sequence data obtained from ex-type culture are available.

Diluviicola capensis Hyde et al.

Distribution: Brunei, Temburong, Kuala Belalong Field Studies Centre, Sungai Esu, on submerged wood (Hyde et al. 1998c).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 3125 (now in IFRD). Sequence data is unavailable.


Pseudoproboscispora Punith., Kew Bull. 54(1): 234 (1999)

Asexual morph Undetermined. Sexual morph Description and illustrations see Wong and Hyde (1999a, b).

Type species: Pseudoproboscispora aquatica (S.W. Wong & K.D. Hyde) Punith., Kew Bull. 54(1): 234 (1999)

Notes: Index Fungorum (2018) lists three species in the genus Pseudoproboscispora. Previous studies showed that the species of Pseudoproboscispora are distributed worldwide and all of them were reported from freshwater habitats (Wong and Hyde 1999a, b; Campbell et al. 2003a; Zhang et al. 2017a, b).

Pseudoproboscispora aquatica (S.W. Wong & K.D. Hyde) Punith.

Proboscispora aquatica S.W. Wong & K.D. Hyde, Mycol. Res. 103: 82 (1999)

Distribution: Australia, North Queensland, near Cairns, Crystal Cascades, on submerged wood; Philippines, Negros Occidental, Bacolod, Kaliban River, on submerged wood (Wong and Hyde 1999a, b). Japan, Koito River, on submerged wood (Tsui et al. 2001a)

Asexual morph: Undetermined

Notes: Holotype HKU (M) 2792 (now in IFRD). Sequence data is not available.

Pseudoproboscispora caudae-suis (Ingold) J. Campb. et al.

Ceriospora caudae-suis Ingold, Trans. Br. Mycol. Soc. 34: 210 (1951)

Distribution: UK, Westmorland, Lake Windermere, on submerged, decorticated twigs of Fraxinus; USA, Arkansas, Lee County, Florida, Hamilton County, Illinois, Johnson County, Lousiana, Caldwell Parish, Mississippi, Franklin County, New York, Adirondack Park, North Carolina, Cheoah River, Oregon, Florence County, Pennsylvania, Columbia County, Tennessee, Great Smoky Mountains National Park, West Virginia, Pocahontas County, Wisconsin, Adams County, on submerged wood in lentic or lotic habitats (Campbell et al. 2003a).

Asexual morph: Undetermined

Notes: Holotype K(M) 109308; Sequence data is not available.

Pseudoproboscispora thailandensis Dong et al.

Distribution: Thailand, Prachuap Khiri Khan, on submerged bamboo in a small River (Zhang et al. 2017a, b).

Asexual morph: Undetermined

Notes: Holotype MFLU 15–2705. ITS, LSU and SSU sequence data are available.


Rhamphoriaceae Réblová

Rhamphoria Niessl, Verh. nat. Ver. Brünn 14: 204 (1876)

Asexual morph Hyphomycetous (Phaeoisaria-like). Sexual morphAscomata partly immersed in the wood, black, single or sometimes in pairs, flask-shaped with an ostiolar neck or depressed cup shape with the neck arising sideways. Peridium composed of thick-walled, dark brown polygonal cells which become thin-walled and hyaline towards interior. Paraphyses filiform, hyaline and numerous. Asci 8-spored, unitunicate, cylindrical, long-stalked. Ascospores monostichous in the ascus, ovoid to oblong, hyaline to subhyaline, smooth-walled.

Type species: Rhamphoria delicatula Niessl, Verh. nat. Ver. Brünn 14: 206 (1876)

Notes: 15 Rhamphoria epithes are listed in Index Fungorum (December 2018) and only one species was reported from freshwater habitats (Ho et al. 2001).

Rhamphoria pyriformis (Pers.) Höhn

Sphaeria pyriformis Pers., Syn. meth. fung. (Göttingen) 1: 64 (1801)

Distribution: China, Hong Kong, Tai Po Kau Forest, on submerged wood in a stream (Ho et al. 2001)

Asexual morph: Undetermined

Notes: Herbarium records CBS H-115, CBS H-17785. ITS, LSU and RPB2 sequence data are available. Ho et al. (2001) reported that Rhamphoria pyriformis was found from freshwater habitats in China, but they did not provide illustrations and descriptions for this species.


Rhodoveronaea Arzanlou et al., Stud. Mycol. 58: 89 (2007)

Asexual morphColonies velvety, floccose; surface olivaceous-grey to dark olivaceous-green, reverse olivaceousblack. Hyphae pale olivaceous, smooth, thin-walled. Conidiophores arising vertically from creeping hyphae, straight or flexuose, simple, thick-walled, red-brown, with inflated basal cell. Conidiogenous cells polyblastic, terminally integrated, sympodial, smooth, thick-walled, pale brown, rachis straight, occasionally geniculate, with crowded, slightly prominent conidium-bearing denticles, denticles flat-tipped, slightly pigmented. Conidia solitary, ellipsoidal to obovoidal, aseptate to multiseptate, with a protruding base and a marginal basal frill, pale brown, thinor slightly thick-walled, smooth. Conidial secession schizolytic (Arzanlou et al. 2007). Sexual morphAscomata nonstromatic, gregarious or solitary, dark brown to black, venter subglobose to conical, immersed; neck conical, emerging above the substratum, straight or slightly curved, ostiolum periphysate. Peridium leathery, two-layered. Paraphyses septate, hyaline, tapering towards the tip, longer than the asci. Asci 8-spored, unitunicate, cylindrical, broadly rounded at the apex, long-stipitate. Ascospores fusiform, aseptate to multiseptate, hyaline (Réblová 2009).

Type species: Rhodoveronaea varioseptata Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 91 (2007)

Notes: Arzanlou et al. (2007) introduced the genus Rhodoveronaea with single asexual species R. varioseptata. Réblová (2009) provided the sexual morph for this species. In this study, we introduce the second species R. aquatica, which was collected from freshwater habitats in northwestern Yunnan, China.

Rhodoveronaea aquatica Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555661, Facesoffungi number: FoF 05435, Fig. 32

Fig. 32
figure 32

Rhodoveronaea aquatica (MFLU 18–1593, holotype). a, b Conidiophores with conidia. c, d Conidiogenous cells with conidia. eo Conidia. Scale bars: ac 50 μm, do 10 μm

Etymology: Referring to the aquatic habitat of this fungus

Holotype: MFLU 18–1593

Saprobic on decaying, submerged wood in freshwater habitats. Asexual morph Hyphomycetous. Colonies effuse, inconspicuous, brown, hairy. Conidiophores 182–310 μm long, 9–13 μm wide (\( {\bar{x}} \) = 246 × 11 μm, n = 20), macronematous, mononematous, cylindrical, arising vertically from creeping hyphae, straight or flexuose, simple, thick-walled, septate, red-brown, paler at apex. Conidiogenous cells polyblastic, terminally integrated, sympodial, smooth, thick-walled, pale brown at the base, paler towards the apex. Conidia 23–27 μm long, 9–11 μm wide (\( {\bar{x}} \) = 25 × 10 μm, n = 20), acropleurogenous, ellipsoid to obovoid, apically rounded, with a flat basal scar, 1–3-septate, pale brown, smooth-walled. Conidial secession schizolytic. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Gaoligong Mountain, saprobic on submerged decaying wood in a freshwater stream, 10 July 2015, X.J. Su, S-718 (MFLU 18–1593, holotype), ex-type living culture MFLUCC 18–1339.

Notes: Rhodoveronaea aquatica resembles R. varioseptata in having macronematous, mononematous, cylindrical, straight or flexuose, septate, red-brown conidiophores, terminally integrated, polyblastic, sympodial, smooth conidiogenous cells, ellipsoid to obovoid, pale brown, septate conidia (Arzanlou et al. 2007; Réblová 2009). However, R. aquatica differs from R. varioseptata in having longer conidiophores (182–310 vs. 80–125 μm) and larger conidia (23–27 × 9–11 μm vs. 8–15 × 3–4 μm). Phylogenetic analysis also shows that Rhodoveronaea aquatica and R. varioseptata are distinct species (Fig. 1, clade 8).


Woswasiaceae Zhang et al.

Cyanoannulus Raja et al., Mycotaxon 88: 8 (2003)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Raja et al. (2003).

Type species: Cyanoannulus petersenii Raja, J. Campb. & Shearer

Notes: The genus Cyanoannulus was introduced by Raja et al. (2003) with single species Cyanoannulus petersenii Raja, J. Campb. & Shearer which was collected from freshwater habitats. And this species is known only from the type locality.

Cyanoannulus petersenii Raja et al.

Distribution: USA, North Carolina and Tennessee, Great smoky Moubtains National Park, on submerged wood (Raja et al. 2003).

Asexual morph: Undetermined

Notes: Holotype ILL, Raja & Hamburger R044-1. ITS and LSU sequence data are available.


Diaporthomycetidae genera incertae sedis

Aquaticola Ho et al., Fungal Diversity Res. Ser. 3: 88 (1999)

Asexual morph Undetermined. Sexual morph Descriptions and illustration see Ho et al. (1999c).

Type species: Aquaticola hyalomura W.H. Ho, C.K.M. Tsui, Hodgkiss & K.D. Hyde, Fungal Divers 3: 88 (1999)

Notes: The genus Aquaticola was introduced for tropical freshwater fungi occurring on submerged wood in Asia with two species and was placed in the family Annulatascaceae (Ho et al. 1999c). Subsequently, Tsui et al. (2003) described three additional Aquaticola species from tropical freshwater environments in Asia and Australia. Réblová et al. (2016a) transferred Aquaticola ellipsoidea to the genus Atractospora as At. ellipsoidea. Four freshwater species are accepted in Aquaticola.

Aquaticola hyalomura Ho et al.

Distribution: China, Hong Kong, Tai Po Kau Forest Stream, on submerged decaying wood (Ho et al. 1999c).

Asexual morph: Undetermined

Notes: Holotype HKU (M) 2969 (now in IFRD). LSU sequence data is available.

Aquaticola longicolla Tsui et al.

Distribution: Australia, north Queensland, near Ravenshoe, submerged in creek (Tsui et al. 2003); Brunei, Tutong River, on submerged wood (Fryar et al. 2004)

Asexual morph: Undetermined

Notes: Holotype IFRD 8682. Sequence data is not available.

Aquaticola minutiguttulata Tsui et al.

Distribution: China, Hong Kong, Sai Kung, Hang Cho Shui, on submerged wood (Tsui et al. 2003).

Asexual morph: Undetermined

Notes: Holotype IFRD 8683. Sequence data is not available.

Aquaticola triseptata Tsui et al.

Distribution: China, Hong Kong, Tai Po, Lam Tsuen River, on submerged wood (Tsui et al. 2003).

Asexual morph: Undetermined

Notes: Holotype IFRD 8684. Sequence data is not available.


Bullimyces Ferrer et al., Mycologia 104(4): 868 (2012)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Ferrer et al. (2012).

Type species: Bullimyces communis Ferrer et al., Mycologia 104(4): 868 (2012)

Notes: Ferrer et al. (2012) introduced the genus Bullimyces with three freshwater species. All these species are known only from their respective type localities.

Bullimyces aurisporus Ferrer et al.

Distribution: Costa Rica, Alajuela, on submerged wood (Ferrer et al. 2012).

Asexual morph: Undetermined

Notes: Holotype ILL AF316–1. LSU and SSU sequence data are available.

Bullimyces communis Ferrer et al.

Distribution: Costa Rica, Alajuela and Heredia, on submerged wood (Ferrer et al. 2012).

Asexual morph: Undetermined

Notes: Holotype ILL AF281–1, other specimens collected from freshwater habitats: ILL AF281–4, ILL AF281–7, ILL AF281–3. LSU, SSU sequence data are available.

Bullimyces costaricensis Ferrer et al.

Distribution: Costa Rica, Limón, on submerged wood (Ferrer et al. 2012).

Asexual morph: Undetermined

Notes: Holotype ILL AF317–1. LSU and SSU sequence data are available.


Cancellidium Tubaki, Trans. Mycol. Soc. Japan 16(4): 357 (1975)

Asexual morphColonies on natural substrate effuse, black, shiny. Mycelium immersed or superficial, composed of septate, subhyaline to hyaline, smooth-walled hyphae. Conidiophores micronematous, short. Conidiogenous cells integrated, terminal, determinate, cylindrical, subhyaline. Conidia acrogenous, solitary, muriform, dictyosporous, strongly flattened, fan-shaped, obovate to obcordate, brown to almost black, shiny. Sexual morph Undetermined.

Type species: Cancellidium applanatum Tubaki, Trans. Mycol. Soc. Japan 16(4): 358 (1975)

Notes: The genus Cancellidium is typified by C. applanatum which was collected from wood blocks of Ochroma pyramidale (Cav. ex Lam.) Urb. in Kobe, Japan. The genus includes two species (Tubaki 1975; Yeung et al. 2006).

Cancellidium applanatum Tubaki

Distribution: Perú, Cusco, on submerged, decomposing woody and herbaceous debris in freshwater habitats (Zelski et al. 2014).

Sexual morph: Undetermined

Notes: Specimens collected from freshwater: ILL 41205, ILL 41206. LSU sequence data is available.


Ceratostomella Sacc., Michelia 1(no. 4): 370 (1878)

Asexual morph Undetermined. Sexual morph Descriptions and illustrations see Inderbitzin (2000) and Réblová (2006).

Type species: Ceratostomella vestita Sacc., Michelia 1(no. 4): 370 (1878)

Notes: Index fungorum (2018) lists 112 epithets for the genus Ceratostomella and only one species was collected from freshwater habitats (Inderbitzin 2000).

Ceratostomella hyalocoronata Inderb

Distribution: China, Guangdong Province, on submerged wood in a stream (Inderbitzin 2000).

Asexual morph: Undetermined

Notes: Holotype UBC F13874. Sequence data is not available.


Clohiesia K.D. Hyde, Nova Hedwigia 61(1-2): 125 (1995)

Asexual morph: Undetermined. Sexual morph: Descriptions and illustrations see Hyde (1995), Cai & Hyde (2007).

Type species: Clohiesia corticola K.D. Hyde, Nova Hedwigia 61(1–2): 126 (1995)

Notes: Clohiesia K.D. Hyde was established by Hyde (1995) to accommodate the freshwater taxon C. corticola from tropical Australia. Subsequently, another two species Clohiesia curvispora and C. lignicola were introduced by Tsui et al. (1998) and Cai and Hyde (2007) respectively. All these species were collected from freshwater habitats. Presently, Clohiesia is placed in the family Annulatascaceae (Maharachchikumbura et al. 2015, 2016; Wijayawardene et al. 2017). However, Zhang et al. (2016) showed that Clohiesia corticola (HKUCC 3712) is close to Chaetosphaeriales based on phylogenetic analyses. In this study, Clohiesia corticola (HKUCC 3712) is basal to Phyllachoraceae based on phylogenetic analysis, therefore we assign this genus to Diaporthomycetidae genera incertae sedis.

Clohiesia corticola K.D. Hyde

Distribution: Australia, Queensland, on submerged wood (Hyde 1995b); Japan, Koito River, on submerged wood (Tsui et al. 2001a).

Asexual morph: Undetermined

Notes: Holotype BRIP 21485. LSU sequence data is available.

Clohiesia curvispora L. Cai & K.D. Hyde

Distribution: China, Yunnan Province, Jinghong city, on submerged wood in a small forest stream (Cai and Hyde 2007).

Asexual morph: Undetermined

Notes: Holotype HKU(M) 10854 (now in IFRD). Sequence data is not available.

Clohiesia lignicola Tsui et al.

Distribution: China, Hong Kong, on submerged wood (Tsui et al. 1998a)

Asexual morph: Undetermined

Notes: Holotype HKU(M) 5533 (now in IFRD). Sequence data is not available.


Hyalorostratum Raja & Shearer, Mycosphere 1(4): 281 (2010)

Asexual morph Undetermined. Sexual morph Description and illustrations refer to Raja et al. (2010).

Type species: Hyalorostratum brunneisporum Raja & Shearer, Mycosphere 1(4): 281 (2010)

Notes: Hyalorostratum was introduced by Raja et al. (2010) with Hyalorostratum brunneisporum Raja & Shearer as type species which was collected from freshwater habitat in north America. Raja et al. (2010) suggested that Hyalorostratum belongs to the order Diaporthales based on results of their phylogenetic analyses, but they did not assign it to any of families in Diaporthales and referred it as Diaporthales genera incertae sedis. In our phylogenetic analysis, two strains of Hyalorostratum brunneisporum formed a distinct subclade and basal to Jobellisiales, we also assign this genus to Diaporthomycetidae genera incertae sedis. Presently, a single species is accepted in this genus.

Hyalorostratum brunneisporum Raja & Shearer

Distribution: USA, Alaska, Headquarters, Lake Kenai Wildlife Refuge; New Hampshire, Hubbard Brook Forest stream, on submerged woody debris, on submerged woody debris (Raja et al. 2010).

Asexual morph: Undetermined

Notes: Holotype ILL 40792. LSU and SSU sequence data are available.


Pseudostanjehughesia J. Yang & K.D. Hyde, Mycol Prog 17: 609 (2018)

Asexual morph Description and illustrations see Yang et al. (2018). Sexual morph Undetermined.

Type species: Pseudostanjehughesia aquitropica J. Yang & K.D. Hyde, Mycol. Progr. 17(5): 610 (2017)

Notes: The genus Pseudostanjehughesia was introduced by Yang et al. (a, b) with single species which was collected from freshwater habitats in Thailand. Pseudostanjehughesia resembles Stanjehughesia and Linkosia due to the absence or reduced conidiophores, and brown and obclavate conidia. Linkosia is easily distinguished from Pseudostanjehughesia by the lageniform or ampulliform conidiogenous cells and distoseptate conidia. It is difficult to separate Pseudostanjehughesia from Stanjehughesia as both genera share similar morphological characters of conidiophores, conidiogenous cells and conidia. However, they are phylogenetically distinct.

Pseudostanjehughesia aquitropica J. Yang & K.D. Hyde

Distribution: Thailand, Prachuap Khiri Khan Province, on decaying wood submerged in a freshwater stream (Yang et al. 2018a).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–0857, isotype GZAAS 17–0010. ITS, LSU, SSU and TEF1α sequence data are available.

Pseudostanjehughesia lignicola Z.L. Luo, K.D. Hyde & H.Y. Su, sp. nov.

Index Fungorum number: IF 555662, Facesoffungi number: FoF 05436, Fig. 33

Fig. 33
figure 33

Pseudostanjehughesia lignicola (DLU 078, holotype) a, cf Conidia. b, g Germinating conidium. h, i Culture on PDA after 20 days (i from below). Scale bars: ag 20 μm

Etymology: Referring to this fungus dwelling on wood

Holotype: DLU 078

Saprobic on decaying, submerged wood in freshwater habitats. Asexual morph Hyphomycetous. Colonies effuse, dark brown, scattered, glistening. Mycelium partly superficial, composed of brown, septate, branched hyphae. Conidiophores indistinct. Conidiogenous cells holoblastic, monoblastic, integrated, terminal, cylindrical. Conidia 51–62 μm long, 16–22 μm wide (\( {\bar{x}} \) = 57 × 19 μm, n = 20), acrogenous, solitary or group in 2–3 conidia, ovoid, slightly curved, multi-euseptate, verrucose, dark brown at base, tapering and becoming pale brown towards the apex, truncate at the base. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, saprobic on decaying wood submerged in Heilong stream in Cangshan Mountain. 15 March 2014, Z.L. Luo, HLXM 7-4 (DLU 078, holotype) ex-type living culture MFLUCC 15–0352.

Notes: Pseudostanjehughesia lignicola morphologically resembles P. aquitropica in having terminal, monoblastic conidiogenous cells, acrogenous, multi-euseptate, verrucose conidia dark brown at base, tapering and becoming pale brown towards the apex (Yang et al. 2018a). However, P. lignicola differs from P. aquitropica in having ovoid, shorter conidia (51–62 μm vs. 55–120 μm), while P. aquitropica have rostrate conidia which are oval or ellipsoidal when young, fusiform or obclavate when mature. Phylogenetic analysis shows that Pseudostanjehughesia lignicola clusters with P. aquitropica with strong support (100% ML) (Fig. 1, clade 10). Following Jeewon and Hyde’s (2016) recommendations on species delimitation for new species, we delved into pairwise dissimilarities of DNA sequences and noted that there are indeed differences in the ribosomal ITS sequences, 68 noticeable nucleotide differences including 11 gaps among the 485 nucleotides analysed between Pseudostanjehughesia lignicola and P. aquitropica. Pseudostanjehughesia lignicola is the second species for Pseudostanjehughesia and both species are collected from freshwater habitats.


Subclass Hypocreomycetidae O.E. Erikss. & Winka

Coronophorales Nannf. (=Melanosporales N. Zhang & M. Blackw.)

Bertiaceae Smyk

Bertia De Not., G. bot. ital. 1(1): 334 (1844)

Asexual morph Undetermined. Sexual morph Updated descriptions and illustrations see Maharachchikumbura et al. (2016).

Type species: Bertia moriformis (Tode) De Not., G. bot. ital. 1(1): 335 (1844)

Notes: The genus Bertia was introduced by De Notaris (1844) and 48 epithets are listed in Index Fungorum (December 2018). Among these species, only Bertia convolutispora K.D. Hyde was collected from freshwater habitats (Hyde 1995c).

Bertia convolutispora K.D. Hyde

Distribution: Australia, north Queensland, on wood submerged in stream (Hyde 1995c).

Asexual morph: Undetermined

Notes: Holotype BRIP 22512. Sequence data is not available.


Glomerellales Chadef. ex Réblová et al.

Reticulascaceae Réblová & W. Gams

Cylindrotrichum Bonord., Handbuch der allgemeinen Mykologie: 88 (1851)

Asexual morph Hyphomycetous. Colonies in vivo brown to black, hairy, effuse. Setae absent. Conidiophores macronematous, mononematous, cylindrical, straight. Conidiogenous cells usually monophialidic, rarely polyphialidic with up to two lateral enteroblastic openings, collarette hyaline to subhyaline. Conidia cylindrical, slightly tapering, rounded at apex, obtuse at base, 1-septate, not constricted at septum, hyaline, guttulate, smooth. Sexual morphStroma absent. Ascomata superficial, solitary, or gregarious, brown, venter subglobose to conical. Ostiolum periphysate. Peridium fragile, 2-layered. Paraphyses septate, hyaline, filiform, forming a branching and anastomosing “network”. Asci 8-spored, unitunicate, cylindrical-clavate, short-stipitate. Ascospores ellipsoidal to fusiform, septate, hyaline.

Type species: Cylindrotrichum oligospermum (Corda) Bonord., Handb. Allgem. mykol. (Stuttgart): 88 (1851)

Notes: Réblová et al. (2011) introduced a new genus Reticulascus including two species. Reticulascus clavatus with its asexual morph Cylindrotrichum clavatum is a common dweller of submerged wood in lotic sites in France. Cylindrotrichum includes 23 names (Rambelli and Onofri 1987) while Reticulascus includes only two names (Réblová et al. 2011). Given its widespread use, priority, and greater number of names, Réblová et al. (2016b) recommend the use of Cylindrotrichum over Reticulascus.

Cylindrotrichum aquaticum (Luo et al.) Z.L. Luo & K.D. Hyde comb. nov.

Blastophorum aquaticum Luo et al., Fungal Divers 80:177 (2016)

Index Fungorum: IF 555663; Facesoffungi number: FoF 05437

Description: For a complete description of this taxon see Hyde et al. (2016) as Blastophorum aquaticum.

Notes: Holotype DLU 084. ITS and LSU sequence data are available. This fungus was introduced as Blastophorum aquaticum by Hyde et al. (2016). It resembles Cylindrotrichum species in morphology of conidiophores, conidiogenous cells and conidia. In this study, the phylogenetic analysis shows that the isolate from type clusters within the genus Cylindrotrichum with strong support (Fig. 1, clade 42). Based on the morphology and phylogeny, we synonymized Blastophorum aquaticum under Cylindrotrichum aquaticum.

Cylindrotrichum clavatum W. Gams & Hol.-Jech., Studies in Mycology 43: 54 (1976).

Distribution: China, Yunnan Province, on submerged wood (Maharachchikumbura et al. 2018); France, on submerged wood of Alnus glutinosa, Fraxinus sp., Platanus sp. (Réblová et al. 2011).

Asexual morph: see Réblová et al. (2011).

Notes: Holotype CBS 128.76, other specimens collected from freshwater habitats: PRM 915717, PRM 915719, DLU 0575, DLU 0572. ITS, LSU, SSU and RPB2 sequence data are available. Cylindrotrichum clavatum was found on submerged wood in freshwater habitats in France and China respectively (Réblová et al. 2011; Maharachchikumbura et al. 2018).

Cylindrotrichum gorii Lunghini

Distribution: China, Yunnan Province, Dali, on submerged in a stream in Jizu Mountain (Maharachchikumbura et al. 2018).

Sexual morph: Undetermined

Notes: Specimen collected from freshwater habitats: MFLU 17–1965. ITS, LSU and RPB2 sequence data are available.

Cylindrotrichum submersum Z.L. Luo, H.Y. Su & K.D. Hyde sp. nov.

Index Fungorum number: IF 555664, Facesoffungi number: FoF 05439, Fig. 34

Fig. 34
figure 34

a Cylindrotrichum submersum (MFLU 18–2320, holotype) a Colonies on wood. b Conidiophore. c, d Conidiogenous cells. ej Conidium. Colonies on PDA from above (l) and reverse (k). Scale bars: b 40 μm, c, d 20 μm, ej 10 μm

Etymology: Referring to the submerged habitats of this fungus

Holotype: MFLU 18–2320

Saprobic on decaying wood submerged in freshwater habitats. Asexual morphColonies on natural substrate, effuse, superficial, brown to black, hairy, in groups. Conidiophores 77–135(–148) µm long, 4–6 μm wide (\( {\bar{x}} \) = 106 × 5 µm, n = 20), macronematous, mononematous, solitary, cylindrical, erect, straight or slightly flexuous, unbranched, septate, dark brown, becoming pale brown to subhyaline towards the apex, smooth. Conidiogenous cells monophialidic, integrated, terminal, without a conspicuous collarette. Conidia 15–17 µm long, 4–6 μm wide (\( {\bar{x}} \) = 16 × 5 µm, n = 20), acrogenous, cylindrical or clavate, rounded at apex, slightly tapering, guttulate, aseptate, hyaline, smooth-walled. Sexual morph Undetermined.

Material examined: CHINA, Yunnan Province, Jizu Mountain, saprobic on decaying wood submerged in a freshwater stream, October 2016, Z.L. Luo, S-988 (MFLU 18–2320, holotype), ex-type living culture DLUCC 0988.

Notes: Cylindrotrichum submersum resembles C. clavatum in having macronematous, solitary, cylindrical, erect, straight or slightly flexuous, unbranched, septate conidiophores of similar size and cylindrical or clavate, hyaline, smooth conidia. However, Cylindrotrichum submersum differs from C. clavatum in having monophialidic conidiogenous cells, aseptate, guttulate, longer (15–17 vs. 8.5–13 µm) conidia, while the later one has monophialidic or polyphialidic conidiogenous cells, 1–2-septate conidia (Gams and Holubova-Jechova 1976; Maharachchikumbura et al. 2018). Furthermore, they are phylogenetically distinct species (Fig. 1, clade 42). To further support Cylindrotrichum submersum as a new species, we compared nucleotide differences with C. aquaticum (MFLUCC 15–0264) following the guidelines of Jeewon and Hyde (2016). Comparison of the 543 nucleotides across the ITS region reveals 36 bp differences including 6 gaps. Based on the differences of morphology and DNA nucleotide, we introduce our isolate as new species in Cylindrotrichum.


Kylindria DiCosmo et al., Mycologia 75 (6): 970 (1983)

Asexual morph Hyphomycetous. Colonies effuse, hairy, dark. Conidiophores macronematous, mononematous, setiform, solitary, erect, sometimes curved, unbranched, often in fascicles, multiseptate, thick-walled, dark brown below, paler above, smooth-walled. Conidia blastic-phialidic, 1-several-septate, oblong-ellipsoid or cylindrical, apex rounded, base tapered and truncate, hyaline, smooth-walled. Sexual morph Undetermined.

Type species: Kylindria triseptata (Matsush.) DiCosmo et al., Mycologia 75(6): 971 (1983)

Notes: DiCosmo et al. (1983) introduced the genus Kylindria based on Cylindrotrichum triseptatum Matsush. (Matsushima 1975). Presently, there are 21 species included in this genus (Mycobank 2019). Three species of Kylindria have been reported from freshwater habitats in China and UK (Hyde and Goh 1999; Maharachchikumbura et al. 2018).

Kylindria aquatica Luo et al.

Distribution: China, Yunnan Province, on submerged wood (Maharachchikumbura et al. 2018).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–1967, paratype MFLU 17–1966. ITS, LSU and RPB2 sequence data are available.

Kylindria chinensis Maharachch. et al.

Distribution: China, Yunnan Province, on submerged wood (Maharachchikumbura et al. 2018).

Sexual morph: Undetermined

Notes: Holotype MFLU 17–1964. ITS, LSU and RPB2 sequence data are available.


Hypocreales Lindau

Bionectriaceae Samuels & Rossman

Didymostilbe Henn., Hedwigia 41: 148 (1902)

Asexual morphConidiomata synnemayous, solitary, errect, straight, cylindrical to clavate, or subulte-capitate, Conidiophores macronematous, unbranched. Conidiogenous cells enteroblastic phialidic, integrated or discrete, terminal or lateral, hyaline, smooth-walled, cylindrical. Conidia acrogenous, hyaline, 0–1-septate, sometimes slightly constricted at septum, cylindrical to slight clavate, broadly rounded at the apex, subtruncate to obconically truncate at the base, thick-walled, smooth. Sexual morph Undetermined.

Type species: Didymostilbe coffeae Henn., Hedwigia 41: 148 (1902)

Notes: The genus Didymostilbe was established by Hennings (1902) with D. coffeae as the type species. Index Fungorum (2019) list 17 epithets for Didymostilbe. Only Didymostilbe australiensis Goh & K.D. Hyde is collected from freshwater habitats from Queensland, Australia (Hyde and Goh 1997).

Didymostilbe australiensis Goh & K.D. Hyde

Distribution: Australia, Queensland, Cape Tribulation, Mountain Lewis, on decaying wood submerged in a stream (Hyde and Goh 1997).

Sexual morph: Undetermined

Notes: Holotype ML 28 (BRIP). Sequence data is not available. Didymostilbe australiensis was introduced by Hyde and Goh (1997) based on their collection from freshwater habitats in Australia, and it is known only from the type locality.

Hypocreaceae De Not.

Trichoderma Pers., Neues Mag. Bot. 1: 92 (1794)

Asexual morph Hyphomycetous. Colonies usually growing rapidly, mycelium initially submerged, eventually with variably matted, floccose, woolly or arachnoid hyaline aerial mycelium. Conidiophores in most species with a broad main axis branched at regular intervals, usually with successive branches apically and distally progressively shorter and narrower, branches more or less divergent, solitary, paired or in verticils. Conidiogenous cells phialidic, typically disposed in divergent verticels terminally on branches of the conidiophore, or in whorls directly beneath septa along the conidiophore and branches, otherwise paired or solitary and irregularly disposed; cylindrical, subulate, lageniform, ampulliform or subglobose; usually attenuated to a narrow, short-cylindrical, conidium bearing neck. Conidia one-celled, typically green, or otherwise colourless, greyish, or brownish; smooth-walled to distinctly roughened, or with sinuate, bullate or winglike projections from the outer wall, subglobose, obovoid, ellipsoid, oblong or short-cylindrical; accumulating in gloeoid heads, sometimes enclosed in a sac-like sheath visible at high magnifications. Sexual morphStromata solitary or aggregated, rounded or elongate, yellowish-brown to brown with white-yellow margin, becoming pulvinate, more rarely turbinate or discoid, circular to irregular in outline, surface smooth to slightly uneven to granular, broadly attached, margin often becoming free and concolorous with stroma surface. Ostiolar openings visible as slightly raised, purple-brown spots. Ascomata immersed in stroma, densely disposed, globose to subglobose. Peridium colourless, consisting of laterally strongly compressed thin hyphae, basally and apically pseudoparenchymatous, indistinct, scarcely differentiated from and merging with the surrounding tissue, apical part flanking the ostioles conspicuously thickened. Asci cylindrical, with short stipe, sometimes with a knob-like base, with apical minute pore. Ascospores 1-septate, dimorphic, distal part subglobose to oval, sometimes slightly tapered towards the upper end, proximal part oblong to wedge-shaped, with lower end broadly rounded, sometimes verrucose, hyaline.

Type species: Trichoderma viride Pers., Neues Mag. Bot. 1: 92 (1794)

Notes: The genus Trichoderma is cosmopolitan in soils and on decaying wood and vegetable matter. Species of Trichoderma are frequently from dominant components of the soil microflora in widely varying habitats (Kubicek and Harman 1998). Au et al. (1992) reported T. glaucum Abboott from freshwater habitats in Hong Kong, China.

Trichoderma glaucum Abboott

Distribution: China, Hong Kong, on submerged leaves (Au et al. 1992).

Sexual morph: Undetermined.

Notes: ITS sequence data is available in GenBank.


Nectriaceae Tul. & C. Tul.

Aquanectria L. Lombard & Crous, Stud. Mycol. 80: 207 (2015)

Holomorph Descriptions and illustrations refer to Lombard et al. (2015).

Type species: Aquanectria penicillioides (Ingold) L. Lombard & Crous, Stud. Mycol. 80: 207 (2015)

Notes: The aquatic genus Aquanectria was established by Lombard et al. (2015) to accommodate two fungal species previously treated as members of the genera Flagellospora and Heliscus (Ingold 1942; Ranzoni 1956a, b; Hudson 1961). Three species are included in this genus and all were collected from freshwater habitats. In this study, description and illustrations for Aquanectria penicillioides are provided and it is new record for China.

Aquanectria jacinthicolor Huang et al.

Distribution: China, Yunnan Province, Baoshan Cit