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Submitted 6 June 2021, Accepted 30 August 2021, Published 14 September 2021
Corresponding Authors: Kevin D. Hyde – e-mail – kdhyde3@gmail.com,
Ting-Chi Wen – e-mail – tingchiwen@yahoo.com 875
Taxonomic studies of Coronophorales and Niessliaceae
(Hypocreomycetidae)
Huang SK1,2,3,4,5, Hyde KD3,6*, Maharachchikumbura SSN7, McKenzie EHC8,
Wen TC1,2,5*
1State Key Laboratory Breeding Base of Green Pesticide and Agricultural Bioengineering, Key Laboratory of Green
Pesticide and Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang 550025, China
2The Engineering Research Center of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou
University, Guiyang 550025, Guizhou Province, China
3Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
4School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
5The Mushroom Research Centre, Guizhou University, Guiyang 550025, China
6Innovative Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District, Guangzhou
510225, China
7School of Life Science and Technology, Center for Informational Biology, University of Electronic Science and
Technology of China, Chengdu 611731, China
8Manaaki Whenua-Landcare Research, Private Mail Bag 92170, Auckland, New Zealand
Huang SK, Hyde KD, Maharachchikumbura SSN, McKenzie EHC, Wen TC 2021 – Taxonomic
studies of Coronophorales and Niessliaceae (Hypocreomycetidae). Mycosphere 12(1), 875–992,
Doi 10.5943/mycosphere/12/1/9
Abstract
The number of species in Sordariomycetes has increased dramatically. However, some genera
discovered earlier are poorly known, documented and thus need to be redescribed. This paper
continues the work on ‘Taxonomic studies of some often overlooked Diaporthomycetidae and
Sordariomycetidae’. We provide descriptions, notes and figures or drawings for 71 over-looked
genera belonging to Bertiaceae, Ceratostomataceae, Chaetosphaerellaceae, Coronophoraceae,
Niessliaceae, Nitschkiaceae and Scortechiniaceae in Hypocreomycetidae based on observation of
type materials, authentic specimens or studying the literature. The taxonomic placements of the
genera are re-defined based on morphology and/or phylogenetic results. As a result, Gonatobotrys
and Neotrotteria, which are often overlooked and have unstable taxonomic placements in previous
studies, belong to Ceratostomataceae based on morphology and phylogeny. The lichenicolous
genus, Nitschkiopsis, which lacks DNA sequence data, is placed in Niessliaceae based on its setose
ascomata and hyaline ellipsoidal ascospores similar to Niesslia species. The type species of
Cryptosphaerella lacks molecular data and is morphologically similar to Cryptosphaeria or
Coronophora, both of which have ostiolate ascomata. Therefore, it is placed in Sordariomycetes,
not in Scortechiniaceae, whose species are characterized by ascomata that lack ostioles. In addition,
the taxonomy of species with ascomata lacking ostioles in Cryptosphaerella have been revised and
introduced as two new genera; Neocryptosphaerella and Pseudocryptosphaerella based on the
phylogenetic result. Seven new combinations proposed are Fracchiaea myricoides (≡ Coronophora
myricoides), Neocryptosphaerella celata (≡ Cryptosphaerella celata), N. globosa (≡ Cry. globosa),
Pseudocryptosphaerella costaricensis (≡ Cry. costaricensis), P. cylindriformis (≡ Cry.
cylindriformis), P. elliptica (≡ Cry. elliptica) and P. malindensis (≡ Cry. malindensis).
Mycosphere 12(1): 875–992 (2021) www.mycosphere.org ISSN 2077 7019
Article
Doi 10.5943/mycosphere/12/1/9
876
Keywords – 9 new taxa – Bertiaceae – Ceratostomataceae – Chaetosphaerellaceae –
Coronophoraceae – Nitschkiaceae – Scortechiniaceae – Niessliaceae
Introduction
An increase in fungal taxonomic studies over the last decade has led to the discovery of
numerous new taxa and the revision of extant taxa, and subsequently, this has resulted in a better
understanding of fungal classification (Hyde et al. 2020d, Maharachchikumbura et al. 2021). More
than 100 taxa in Sordariomycetes, such as Chaetomium, Humicola, Podospora, Thielavia and
Zopfiella species, have been repositioned based on molecular data (Wang et al. 2016, 2019a, b, Vu
et al. 2019, Marin-Felix et al. 2020, Hyde et al. 2020e, Huang et al. 2021), while, many new species
have been introduced based on morphology and phylogenetic analyses. This data is scattered in
many publications, and information is available on various webpages (Pem et al. 2019a, Calabon et
al. 2020, Bundhun et al. 2020, Hyde et al. 2020a, Wijesinghe et al. 2021).
A major problem in fungal classification, however, is the number of genera that have not
been studied in recent decades (Dayarathne et al. 2016, Huang et al. 2021). Many have very brief
descriptions and no drawing or photos (Saccardo 1873b, 1880, 1882a, 1886, Penzig & Saccardo
1897, Boudier 1904), and therefore it is tough to place these taxa in a classification scheme. As
more studies have taken place in the tropics and researchers rely on poorly documented genera
from the northern hemisphere, it is extremely hard to link new collections to old names. Similarly,
sequence data for many old northern hemisphere genera are lacking (Berkeley 1875, 1876, Ellis &
Everhart 1892, Lundqvist 1972), and new collections from the tropics with sequence data cannot be
linked to the extant genera. The trend to introduce additional new taxa, however, is not hindered
despite progress in the discovery of many new species from China and Thailand (Hyde et al. 2019,
2020b, c, Jayawardena et al. 2018, Wanasinghe et al. 2018, Hapuarachchi et al. 2019, Luo et al.
2019, Pem et al. 2019b, Phookamsak et al. 2019, Yuan et al. 2020).
Several previous studies have contributed DNA sequence data and taxonomic illustrations of
earlier published taxa. This information has helped to clarify the placement of these taxa (Wang et
al. 2016, 2019a, b, 2020, Marin-Felix et al. 2020). For example, the polyphyletic Lasiosphaeriaceae
with cylindrical to pod-like ascospores was divided into nine families, Bombardiaceae,
Diplogelasinosporaceae, Lasiosphaeriaceae, Lasiosphaeridaceae, Naviculisporaceae,
Neoschizotheciaceae, Podosporaceae, Strattoniaceae and Zygospermellaceae, based on
phylogenetic analyses (Wang et al. 2019a, Vu et al. 2019, Marin-Felix et al. 2020, Huang et al.
2021). The difference between Neurospora and Sordaria was clarified with the ascospores of the
former having pitted or ribbed walls and those of the latter being smooth-walled (Cai et al. 2006,
Huang et al. 2021). Several Cercophora species having ellipsoidal to triangular ascospores were
assigned to Cladorrhinum, Mammaria, Podospora and Triangularia and this taxonomic
arrangement was supported by phylogenetic results (Marin-Felix et al. 2020, Huang et al. 2021).
Wang et al. (2020) sequenced and analysed additional taxa from the Cordycipitaceae but the
affinities and taxonomic placement of many taxa still need to be investigated.
An important class, Sordariomycetes was introduced by Eriksson & Winka (1997). Many
poorly known taxa were initially classified to this class based on morphology (Barr 1983, 1990,
Eriksson & Hawksworth 1993, Lumbsch & Huhndorf 2007, Hyde et al. 2020e).
Hypocreomycetidae was listed in Sordariomycetes and previously accommodated Coronophorales,
Hypocreales, Melanosporales and Microascales (Huhndorf et al. 2004b, Lumbsch & Huhndorf
2007). Subsequently, Conioscyphales, Falcocladiales, Glomerellales, Pleurotheciales, Savoryellales
and Torpedosporales were also placed in this subclass based on morphology and phylogenetic
results (Jones et al. 2015, Maharachchikumbura et al. 2015, 2016). Parasympodiellales was initially
treated as a member of Sordariomycetes (Hernández-Restrepo et al. 2017), and Hyde et al. (2020e)
accepted this order in Hypocreomycetidae based on the divergent time estimates. Members of the
Conioscyphales, Fuscosporellales, Pleurotheciales and Savoryellales clustered as a distinct,
independent clade Savoryellomycetidae (Hongsanan et al. 2017). Therefore, there are currently
seven orders, viz Coronophorales, Falcocladiales, Glomerellales, Hypocreales, Microascales,
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Parasympodiellales and Torpedosporales included in Hypocreomycetidae and its divergence time
has been estimated as 256 MYA (Hyde et al. 2020e).
Coronophorales is a poorly known order, which has turbinate or globose ascomata with long
necks or Quellkörper structures and/or Munk pores in the cells of the ascoma wall (Mugambi et al.
2010, Carneiro de Almeida et al. 2016), and is found on woody substrates worldwide (Mugambi &
Huhndorf 2010, Maharachchikumbura et al. 2016, Hyde et al. 2020e). Several members of this
order were initially placed in Sordariales (Nannfeldt 1932, 1975, Müller & von Arx 1973,
Nannfeldt & Santesson 1975, Barr 1990, Subramanian & Sekar 1990). However, Huhndorf et al.
(2004b) clarified the affinities of Coronophorales based on phylogenetic analyses and subsequently,
this order contained six families, Bertiaceae, Ceratostomataceae, Chaetosphaerellaceae,
Coronophoraceae, Nitschkiaceae and Scortechiniaceae (Nannfeldt 1975, Subramanian & Sekar
1990, Mugambi & Huhndorf 2010, Maharachchikumbura et al. 2015, 2016, Hyde et al. 2017,
2020e). The taxa in Ceratostomataceae usually have long-necked ascomata and ellipsoidal to
broadly fusiform ascospores (Marin-Felix et al. 2018, Hyde et al. 2020e). Bertiaceae,
Chaetosphaerellaceae, Coronophoraceae and Nitschkiaceae have black, tuberculate ascomata with
ostioles or lacking ostioles, but have obconical Quellkörper structures (Saccardo 1882a, Patouillard
& Lagerheim 1895, Subramanian & Sekar 1990, Réblová 1999a, b). Scortechiniaceae is mainly
characterized by Quellkörper structures in the turbinate ascomata and allantoid to ellipsoidal
ascospores (Huhndorf et al. 2004b).
Hypocreales, as the largest order in Hypocreomycetidae, is common silvicolous fungi with a
worldwide distribution (Rehner & Samuels 1995, Rossman et al. 1999, Maharachchikumbura et al.
2016). Hyde et al. (2020e) listed 14 families of the order. Some of them with medicinal and edible
values are often mentioned in recent studies, such as Ophiocordycipitaceae and Cordycipitaceae
(Quandt et al. 2014, Kepler et al. 2017). However, the saprobic Niessliaceae, as an earlier member,
has rarely been mentioned over the past decade (Tang et al. 2007, Lombard et al. 2015,
Maharachchikumbura et al. 2016, Hyde et al. 2020e). This family usually has membranaceous
ascomata with setae or hairs and ellipsoidal to filiform ascospores and accommodates 21 genera,
but many members lack sequence data (Rehner & Samuels 1995, Samuels & Barr 1997, Rossman
et al. 1999, Maharachchikumbura et al. 2016, Hyde et al. 2020e).
In this study, we revisit rarely mentioned taxa in Coronophorales and Niessliaceae. Most of
them were found many years ago and lack detailed descriptions, photoplates and molecular data.
These facts make it hard to recognize and demarcate species, and it is not easy to compare them
with new collections in subsequent studies. Therefore, we re-examine the type material and/or
authentic specimens together with the literature of the taxa in Coronophorales and Niessliaceae and
provide illustrations and descriptions. Morphology and phylogenetic analyses are used to determine
their affinities and taxonomic placement.
Materials & Methods
Material examination
Specimens were loaned from BPI, BR, CUP, F, FH, G, HBG, IMI, MAF, NY, PC, PDD,
PRM, S, TRTC, UPS and W herbaria#. Examination of specimens was done following the method
of Senanayake et al. (2020). Samples were observed in pure water and/or 5% KOH reagent for
sectioning; sometimes, they were stained in Congo red or cotton blue to observe the reaction.
Specimens were examined using a Motic SMZ 168 stereomicroscope. Micromorphological
# Institution: BPI: U.S. National Funngus Collections, USDA-ARS, USA; BR: Meise Botanic Garden, Belgium; CUP:
Cornell University, USA; F: Field Museum of Natural History, USA; FH: Harvard University, USA; G: Conservatoire
et Jardin botaniques de la Ville de Genève, Switzerland; HBG: University of Hamburg, Germany; IMI: CABI
Bioscience UK Centre, UK; MAF: Facultad de Farmacia, Universidad Complutense, Madrid, Spain; NY: The New
York Botanical Garden, USA; PC: Muséum National d’Histoire Naturelle, France; PDD: Manaaki Whenua - Landcare
Research, New Zealand; PRM: National Museum, Czech Republic; S: Swedish Museum of Natural History, Sweden;
TRTC: Royal Ontario Museum, Canada; UPS: Museum of Evolution, Sweden; W: Naturhistorisches Museum Wien,
Austria.
878
characters were examined using a Nikon ECLIPSE 80i compound microscope, and images were
captured with a Canon EOS 550D and/or a Canon EOS 600D digital camera. Taro soft ® Image
Framework program version 0.9.0.7 was used for measurements. Some of the type specimens were
in poor condition or could not be loaned. For these taxa, we redrew the morphological features
based on the original publications. The images used for photoplates were processed with Adobe
Photoshop CS6 software (Adobe Systems, USA).
Phylogenetic analyses
Sequences were obtained from GenBank mostly following previous publications and aligned
by using the default settings of MAFFT version 7 (http://mafft.cbrc.jp/alignment/server/index.html)
(Katoh & Standley 2013). These datasets were manually aligned using Bioedit 7.0.9.0 (Hall 1999).
The phylogenetic analyses of the combined the nuclear ribosomal large subunit (LSU), nuclear
ribosomal small subunit (SSU), internal transcribed spacer (ITS), translation elongation factor
(TEF), RNA polymerase II (RPB2) and beta-tubulin (TUB) sequence data were performed using
maximum-likelihood (ML) and Bayesian inference (BI). The evolutionary model was obtained
using jModeltest 2.1.1 under the Akaike information criterion (AIC) (Darriba et al. 2012). The
selected models which are not implemented in MrBayes were substituted by the closest over-
parameterized model (Huelsenbeck & Rannala 2004). The TIM2, TIM3, TrN and TrNef models
were replaced by the GTR model (Lecocq et al. 2013). The proportion of invariable sites (I) and
gamma distributed rates (G) assured in jModeltest were conserved in all models. The ML analyses
were run with IQ-TREE (Nguyen et al. 2015, Chernomor et al. 2016), using models for each
partitioned gene, with 1000 rapid bootstrap replicates. BI was implemented by MrBayes v.3.0b4
(Ronquist & Huelsenbeck 2003) with the best-fit model of sequence evolution. Phylogenetic tree
was viewed with FigTree v1.4.0 (http://tree.bio.ed.ac.uk/software/figtree/) and processed by Adobe
Illustrator CS5. The final alignment and trees were deposited in TreeBASE.
Table of contained information of genera in Coronophorales and Niessliaceae in
Hypocreomycetidae. Those not in agreement with Wijayawardene et al. 2020 are marked with an
asterisk (*).
Phylum Ascomycota Caval.-Sm.
Class Sordariomycetes O.E. Erikss. & Winka
Subclass Hypocreomycetidae O.E. Erikss. & Winka
Coronophorales Nannf.
Bertiaceae Smyk
Bertia De Not.
Gaillardiella Pat.
Ceratostomataceae G. Winter
Arxiomyces P.F. Cannon & D. Hawksw.
Dactylidispora Y. Marín et al.
Echinusitheca Y. Marín et al.
Erythrocarpon Zukal
Gonatobotrys Corda 1839*
Harzia Costantin
Melanospora Corda
Microthecium Corda
Neotrotteria Sacc.*
Pseudomicrothecium Y. Marín et al.
Pustulipora P.F. Cannon
Rhytidospora Jeng & Cain
Scopinella Lév.
Setiferotheca Matsush.
879
Syspastospora P.F. Cannon & D. Hawksw.
Vittatispora P. Chaudhary et al.
Chaetosphaerellaceae Huhndorf et al.
Chaetosphaerella E. Müll. & C. Booth
Crassochaeta Réblová
Neochaetosphaerella Lar.N. Vassiljeva et al.
Spinulosphaeria Sivan.
Coronophoraceae Höhn.
Coronophora Fuckel
Nitschkiaceae (Fitzp.) Nannf.
Acanthonitschkea Speg.
Biciliosporina Subram. & Sekar
Botryola Bat. & J.L. Bezerra
Fracchiaea Sacc.
Groenhiella Jørg. Koch et al.
Janannfeldtia Subram. & Sekar
Lasiosphaeriopsis D. Hawksw. & Sivan.
Loranitschkia Lar.N. Vassiljeva
Nitschkia G.H. Otth ex P. Karst.
Rhagadostoma Körb.
Rhagadostomella Etayo
Tortulomyces Lar.N. Vassiljeva et al.
Scortechiniaceae Huhndorf et al.
Biciliospora Petr.
Coronophorella Höhn.
Euacanthe Theiss.
Neocryptosphaerella S.K. Huang & K.D. Hyde *
Neofracchiaea Teng
Pseudocatenomycopsis Crous & L.A. Shuttlew.
Pseudocryptosphaerella S.K. Huang & K.D. Hyde *
Scortechinia Sacc.
Scortechiniella Arx & E. Müll.
Scortechiniellopsis Sivan.
Tympanopsis Starbäck
Hypocreales Lindau
Niessliaceae Kirschst.
Atronectria Etayo
Circinoniesslia Samuels & M.E. Barr
Cryptoniesslia Scheuer
Eucasphaeria Crous
Malmeomyces Starb.
Melchioria Penz. & Sacc.
Miyakeomyces Hara
Myrmaeciella Lindau
Myrtacremonium Crous
Neoeucasphaeria Crous
Niesslia Auersw.
= Hyaloseta A.W. Ramaley*
Nitschkiopsis Nannf. & R. Sant.*
Paraniesslia K.M. Tsui et al.
Pseudohyaloseta Tibpromma & K.D. Hyde
880
Pseudonectriella Petr.
Pseudorhynchia Höhn.
Rosasphaeria Jaklitsch & Voglmayr
Taiwanascus Sivan & H.S. Chang
Trichosphaerella E. Bommer et al.
Valetoniella Höhn.
Valetoniellopsis Samuels & M.E. Barr
Other genera in Hypocreomycetidae
Glomerellales genera incertae sedis
Ascocodinaea Samuels, Cand. & Magni
Torpedosporales E.B.G. Jones, Abdel-Wahab & K.L. Pang
Etheirophoraceae Rungjind., Somrith. & Suetrong
Etheirophora Kohlm. & Volkm.-Kohlm.
Swampomyces Kohlm. & Volkm.-Kohlm.
Juncigenaceae E.B.G. Jones, Abdel-Wahab & K.L. Pang
Juncigena Kohlm., Volkm.-Kohlm. & O.E. Erikss.
Sordariomycetes genera incertae sedis
Cryptosphaerella Sacc.*
Hypocreomycetidae O.E. Erikss. & Winka, Myconet 1(1): 6 (1997)
Eriksson & Winka (1997) introduced the subclass Hypocreomycetidae closely related to
Savoryellomycetidae based on phylogenetic results (Hongsanan et al. 2017). There are six orders in
this subclass viz. Coronophorales, Falcocladiales, Glomerellales, Hypocreales, Microascales,
Torpedosporales (Wijayawardene et al. 2020) and most contain species as saprobes in aquatic
and/or terrestrial habitats (Maharachchikumbura et al. 2016, Hyde et al. 2020e).
Coronophorales Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 54 (1932)
The genera in Coronophorales have a worldwide distribution as saprobes on wood and are
characterized by superficial ascomata, sometimes with a subiculum, Quellkörper (a mucilaginous
mass of thick-walled cells as conical structure within the cupulate or collapsed ascoma, believed to
induce rupture of the ascoma and release ascospores) and Munk pores (a small pore with annular
thickened rim, between cells of the ascoma wall) (Mugambi & Huhndorf 2010). Asci are clavate to
subglobose with subcylindrical to ellipsoidal ascospores and asexual morphs are usually
hyphomycetous and phialidic (Mugambi & Huhndorf 2010, Maharachchikumbura et al. 2016,
Hyde et al. 2020e). This order was considered as a synonym of Nitschkiaceae and Sordariales (Barr
1990, Müller & von Arx 1973, Nannfeldt 1932, Nannfeldt & Santesson 1975, Subramanian &
Sekar 1990). However, multi-gene analyses were used to show that Coronophorales which
accommodates Bertiaceae, Ceratostomataceae, Chaetosphaerellaceae, Coronophoraceae,
Nitschkiaceae and Scortechiniaceae is distinct, and is sister to Falcocladiales in Sordariomycetidae
(Hongsanan et al. 2017, Huang et al. 2021, this study, Fig. 1).
Bertiaceae Smyk, Ukr. bot. Zh. 38(6): 47 (1981)
Saprobic on wood. Sexual morph: Subiculum inconspicuous or composed of brown to dark
brown, branched, septate hyphae. Ascomata solitary or gregarious, superficial to erumpent, usually
sitting on a subiculum, cupuliform, turbinate, globose to subglobose, carbonaceous to
membranaceous, dark brown to black, tuberculate or smooth, collapsing, collapsing laterally or not
collapsing, with ostioles or a papillate ostiole-like structure, without an external opening,
periphysate, Quellkörper absent. Peridium composed of carbonaceous to membranaceous, dark
881
brown, brown to hyaline cells of textura angularis to textura prismatica, mostly Munk pores
present. Paraphyses few, inflated, branched. Asci 8-spored, unitunicate, cylindrical to clavate, long
pedicellate, apical ring indistinct or absent. Ascospores 2–3-seriate or irregularly arranged, hyaline
to brown, ellipsoidal to fusiform, 0 to multi-septate, smooth-walled, mostly with guttules. Asexual
morph: Undetermined (adapted from Patouillard & Lagerheim 1895, Mugambi & Huhndorf 2010,
Maharachchikumbura et al. 2015, Hyde et al. 2020e).
Figure 1 – Phylogram generated from maximum likelihood analysis based on combined LSU,
SSU, TEF, RPB2, TUB and ITS sequence data with the confidence values of bootstrap (BS)
proportions from the Maximum Likelihood (ML) analysis (ML-BS > 50%, before the backslash)
and the posterior probabilities (PP) from the Bayesian (BY) analysis (BY-PP > 0.90, after the
backslash) above corresponding nodes. The ‘--’ indicates a lack of statistical support (< 50% for
ML-BS and < 0.90 for BY-PP). One hundred and ninety-eight strains are included in the combined
882
analyses which comprised 4581 characters (842 characters for LSU, 1025 characters for SSU, 950
characters for TEF, 876 characters for RPB2, 391 characters for TUB, 497 characters for ITS) after
alignment. Strains of Xylariomycetidae are used as outgroup taxa. The best score in IQ-TREE
explores with a final likelihood value of -90984.2293 is presented. The model of each partitioned
gene is LSU: GTR+I+G; SSU: GTR+G; TEF: GTR+I+G; RPB2: GTR+I+G; TUB: GTR+I+G;
ITS: GTR+G. Sequences generated indicated in bold. Strain numbers are noted after the species
names and ex-type strains marked with ‘T’ after the culture number. Alignment is available at
TreeBASE (URL: http://purl.org/phylo/treebase/phylows/study/TB2:S28271).
Figure 1 – Continued.
883
Type genus – Bertia De Not.
Notes – Smyk (1981) introduced the family Bertiaceae and initially included Bertia which
has tuberculate ascomata and fusiform ascospores. Subsequently, Mugambi & Huhndorf (2010) re-
evaluated the Coronophorales and transferred the genus Gaillardiella to Bertiaceae based on multi-
gene analysis. Therefore, Bertia and Gaillardiella were confirmed as members of Bertiaceae, and
they have tuberculate ascomata, cylindrical to clavate asci and ellipsoidal to fusiform ascospores
(Huhndorf et al. 2004b, Mugambi & Huhndorf 2010). In this study, strains of these two genera
clustered with 100%ML/1.00BY support and are closely related to Nitschkiaceae (Fig. 1).
Bertia De Not., G. bot. ital. 1(1): 334 (1844)
Saprobic on wood. Sexual morph: Subiculum inconspicuous or composed of brown to dark
brown, branched, septate hyphae. Ascomata solitary or gregarious, superficial to erumpent, usually
sitting on a subiculum, cupuliform, turbinate, globose to subglobose, carbonaceous to
membranaceous, dark brown to black, tuberculate or smooth, collapsing, collapsing laterally or not
collapsing, with ostioles or a papillate ostiole-like structure, without an external opening,
periphysate. Peridium carbonaceous or membranaceous, outer layer composed of dark brown to
brown cells of textura angularis, Munk pores present; inner layer composed of hyaline cells of
textura prismatica. Paraphyses few, inflated, branched. Asci 8-spored, unitunicate, cylindrical to
clavate, long pedicellate, apical ring indistinct or absent. Ascospores 2–3-seriate or irregularly
arranged, hyaline to brown, ellipsoidal to fusiform, 0 to multi-septate, smooth-walled, mostly with
guttules. Asexual morph: Undetermined (adapted from Mugambi & Huhndorf 2010,
Maharachchikumbura et al. 2015, Hyde et al. 2020e).
Notes – Bertia is typified by B. moriformis and its species mostly have tuberculate ascomata
and hyaline ascospores (De Notaris 1844, Mugambi & Huhndorf 2010). It is closely related to
members of Nitschkiaceae based on multi-gene analyses (Huhndorf et al. 2004b, Mugambi &
Huhndorf 2010, Maharachchikumbura et al. 2015, Hyde et al. 2020e, Fig. 1).
Bertia moriformis (Tode) De Not., G. bot. ital. 1(1): 335 (1844) Fig. 2
Basionym – Sphaeria moriformis Tode, Fung. mecklenb. sel. (Lüneburg) 2: 22 (1791)
Facesoffungi number: FoF 01113
Saprobic on decorticated branches. Sexual morph: Ascomata 0.7–1 × 0.4–0.8 mm (x
̄ = 0.8 ×
0.6 mm, n = 10), scattered to gregarious, superficial or erumpent through the surface of the host,
globose to subglobose, dark brown to black, carbonaceous, tuberculate, collapsing laterally or not
collapsing, with a short papillate ostiole-like structure, without an external opening, periphysate.
Peridium 90–250 µm, comprising three layers, outer layer composed of dark tissues, carbonaceous;
middle layer composed of membranaceous, dark brown to brown cells of textura angularis, Munk
pores present; inner layer composed of membranaceous, hyaline cells of textura prismatica.
Paraphyses 9–13 µm wide, few, inflated, branched. Asci (120–)140–155(–180) × (13–)15–20(–23)
µm (x
̄ = 148 × 18 µm, n = 30), 8-spored, unitunicate, clavate, with long tapering pedicel, apical ring
absent. Ascospores (22–)25–35(–40) × (3–)5–7(–9) µm (x
̄ = 30 × 6.5 µm, n = 50), 2–3-seriate or
irregularly arranged, hyaline to pale brown, cylindrical to fusiform, 0–1-septate, slightly curved,
smooth-walled, with guttules. Asexual morph: Undetermined.
Material examined – Italy, Riva Valdobbia, on decorticated branch of Rhododendron sp.
(Ericaceae), 30 April 1859, Carestia (S-F298).
Known hosts and distribution – on decayed wood in Germany (type locality), Italy and USA
(Tode 1791, Huhndorf et al. 2004b).
Notes – Sphaeria moriformis was collected in Germany in March or April 1790 (Tode 1791).
De Notaris (1844) established Bertia based on S. moriformis. We were unable to obtain the type
material but re-examined a specimen collected in Italy in 1859. This sample is similar to B.
moriformis described by Tode (1791), which has gregarious, oval, tuberculate ascomata and
fusiform ascospores. Bertia species aggregate into a cluster based on phylogenetic result with
100%ML/1.00BY support (Fig. 1).
884
Figure 2 – Bertia moriformis: (S-F298). a Material. b–d Ascomata on the host. e Ascoma in cross
section. f–g Peridium. h Wall cells with Munk pores (arrows). i–k Asci with inflated paraphyses.
l–o Ascospores. Notes: k, m–o stained in Melzer’s reagent. Scale bars: b = 1 mm, c–e = 500 µm,
f = 200 µm, g = 100 µm, i–k = 50 µm, h, l–m = 20 µm, n–o = 10 µm.
Gaillardiella Pat., in Patouillard & Lagerheim, Bull. Soc. mycol. Fr. 11(4): 226 (1895)
Saprobic on wood. Sexual morph: Ascomata gregarious, superficial to erumpent, globose to
subglobose, coriaceous, dark brown to black, tuberculate, glabrous, collapsing or not, with ostioles
885
or a short papillate ostiole-like structure, without an external opening. Peridium composed of
coriaceous to membranaceous, brown to hyaline cells of textura angularis to textura prismatica,
sometimes with Munk pores. Paraphyses filiform. Asci 8-spored, unitunicate, cylindrical to clavate,
long pedicellate, apical ring indistinct or absent. Ascospores 2–3-seriate or irregularly arranged,
hyaline to brown, ellipsoidal to clavate, 0–1-septate, smooth-walled, mostly with guttules. Asexual
morph: Undetermined (adapted from Patouillard & Lagerheim 1895, Mugambi & Huhndorf 2010).
Notes – Gaillardiella was placed in Bertiaceae because it has tuberculate ascomata and
ellipsoidal to fusiform ascospores similar to Bertia, and its generic type, G. pezizoides, nests with
Bertia based on phylogenetic results (Patouillard & Lagerheim 1895, Huhndorf et al. 2004b,
Mugambi & Huhndorf 2010, Fig. 1).
Gaillardiella pezizoides Pat., in Patouillard & Lagerheim, Bull. Soc. mycol. Fr. 11(4): 226 (1895)
Fig. 3
Facesoffungi number: FoF 10053
Saprobic on decorticated branches. Sexual morph: Subiculum composed of brown to dark
brown, branched, septate hyphae. Ascomata 300–550 µm diam., superficial, sitting on thin
subiculum, scattered to gregarious, coriaceous, black, cupuliform, tuberculate, glabrous, collapsing
laterally, with a short, papillate ostiole-like structure, without an external opening. Peridium 150–
200 µm thick, comprising two layers, outer layer composed of coriaceous, dark brown to black
cells of textura angularis, Munk pores present; inner layer composed of membranaceous, hyaline
cells of textura prismatica. Asci (75–)80–95(–125) × (7–)8–10(–13) µm (x
̄ = 90 × 9 µm, n = 20), 8-
spored, unitunicate, subclavate to cylindrical, with a long pedicel, evanescent. Ascospores (8–)10–
12(–14) × 3–4.5(–6) µm (x
̄ = 11 × 4 µm, n = 50), hyaline to brown, oval to ellipsoidal, 1-septate,
smooth-walled, with an apical germ pore, guttulate. Asexual morph: Undetermined (adapted from
Patouillard & Lagerheim 1895).
Material examined – Puerto Rico, El Yunque National Forest, El Verde Research Area, 16-ha
Grid, Luquillo Mts, 350 to 425 m, on decorticated wood, 26 January 1996, S.M. Huhndorf (F-
SMH2070).
Known hosts and distribution – on decayed wood in Ecuador (type locality), Puerto Rico and
Kenya (Patouillard & Lagerheim 1895, Mugambi & Huhndorf 2010).
Notes – The strains of Gaillardiella pezizoides were sequenced by Mugambi & Huhndorf
(2010). In this study, these strains are sister to Bertia sinensis (57%ML/1.00BY, Fig. 1). We were
unable to obtain the type material of G. pezizoides and therefore, re-examined and illustrated an
authentic specimen SMH2070 collected by Huhndorf.
Ceratostomataceae G. Winter [as ‘Ceratostomeae’], Rabenh. Krypt. -Fl., Edn 2 (Leipzig) 1.2: 247
(1885)
Saprobic on wood or vegetation or dung or parasitic on other fungi or closely associated with
other fungi or isolated from soil. Sexual morph: Ascomata perithecial or without an external
opening, solitary or gregarious, superficial to semi-immersed, carbonaceous, coriaceous or
membranaceous, opaque or translucent, yellow to brown, usually glabrous, globose to subglobose,
with or lacking ostioles, usually with long necks. Necks cylindrical to conical or absent, comprising
fasciculate, pale yellow to brown, cylindrical, septate hyphae, straight or slightly curved, with a
tapering or jagged apex, sometimes swelling near the apex, ascospores usually accumulating in
necks. Peridium membranaceous, composed of yellow to brown cells of textura angularis or
textura prismatica. Asci 4–8-spored, or multi-spored, unitunicate, cylindrical to clavate, pedicellate
or sessile, apical ring absent, evanescent. Ascospores uni-, bi-seriate or irregularly arranged, hyaline
to dark brown, ellipsoidal to broadly fusiform, aseptate, with a germ pore at one or each end,
smooth or ornamented, mostly with guttules, without sheath. Asexual morph: Hyphomycetous.
Conidiophores semi-macronematous, mononematous, hyaline to brown, branched or unbranched,
phialidic or bulbils, sometimes swollen. Conidia aggregated, hyaline to brown, usually aseptate,
globose to citriform or ellipsoidal, with guttules (adapted from Zukal 1885, Patouillard &
886
Lagerheim 1895, Whaley & Barnett 1963, Jeng & Cain 1977, Hoch 1977, Walker & Minter 1981,
Cannon 1982, Cannon & Hawksworth 1982, Horie et al. 1986, Vakili 1989, Matsushima 1995,
García et al. 2004, Mugambi & Huhndorf 2010, Fan et al. 2012, Maharachchikumbura et al. 2015,
Chaudhary et al. 2016, Marin-Felix et al. 2018, Crous et al. 2019b, Hyde et al. 2020e).
Figure 3 – Gaillardiella pezizoides: (F-SMH2070). a Material. b Ascoma on wood. c Ascoma
cross section. d–e Peridium (arrow points to Munk pores). f–g Asci. h Ascospores (arrow points to
germ pore). Scale bars: b–c = 500 µm, d = 200 µm, e–g = 50 µm, h = 20 µm.
Melanospora Corda
Notes – The family Ceratostomataceae was introduced by Winter (1885) and is characterized
by opaque to semi-translucent, yellow to brown ascomata with cylindrical to clavate asci and
887
brown to dark brown, aseptate ascospores without a sheath (Hyde et al. 2020e). Ceratostomataceae
was considered as a synonym of Melanosporaceae based on the similar morphology of the
hyphomycetous asexual morph (Cannon & Hawksworth 1983, Horie et al. 1986, Barr 1990,
Hawksworth et al. 1995, Gams 2000). Subsequently, this family was accommodated in
Coronophorales (= Melanosporales) based on phylogenetic study (Zhang & Blackwell 2002, Zhang
et al. 2006, Hibbett et al. 2007, Maharachchikumbura et al. 2015, 2016, Hongsanan et al. 2017,
Hyde et al. 2017, 2020e, Wijayawardene et al. 2020). Wijayawardene et al. (2020) listed 14 genera
in this family, viz Arxiomyces, Dactylidispora, Echinusitheca, Erythrocarpon, Harzia,
Melanospora, Microthecium, Pseudomicrothecium, Pustulipora, Rhytidospora, Scopinella,
Setiferotheca, Syspastospora and Vittatispora. In this study, phylogenetically, Gonatobotrys is
sister to Vittatispora and Neotrotteria is basal to other Ceratostomataceae; and these two genera are
accepted as additional members of Ceratostomataceae (Fig. 1). Therefore, 16 genera are accepted in
Ceratostomataceae based on morphology and/or phylogenetic results.
Arxiomyces P.F. Cannon & D. Hawksw., Trans. Br. mycol. Soc. 81(3): 644 (1983)
Saprobic on wood and/or isolated from soil. Sexual morph: Ascomata perithecial, gregarious,
semi-immersed or erumpent through attached bark, globose to subglobose, dark brown to black,
sometimes semi-translucent, glabrous or slightly hairy, ostiolate, with long necks. Asci 4- or 8-
spored, unitunicate, obovoid to clavate, with long tapering pedicel, apical ring indistinct or absent,
evanescent. Ascospores irregular or bi-seriate, hyaline becoming brown or dark brown, ovoid to
ellipsoidal, aseptate, smooth-walled, mostly with guttules, with a rounded apex and truncated base,
usually with a collapsed germ pore. Asexual morph: Undetermined (adapted from Cannon &
Hawksworth 1983, Horie et al. 1986).
Notes – Arxiomyces accommodates three species, A. campanulatus, A. vitis and A.
zuberiensis, and is characterized by ellipsoidal ascospores with a rounded apex and truncated base,
but no molecular data is available for this genus (Cannon & Hawksworth 1982, 1983, Horie et al.
1986). These characteristics are similar to Melanospora; thus, Arxiomyces was placed in
Ceratostomataceae (Cannon & Hawksworth 1982, 1983, Horie et al. 1986, Maharachchikumbura et
al. 2016, Hyde et al. 2020e).
Arxiomyces vitis (Fuckel) P.F. Cannon & D. Hawksw., Trans. Br. mycol. Soc. 81(3): 644 (1983)
Fig. 4
Basionym – Ceratostoma vitis Fuckel, Jb. nassau. Ver. Naturk. 23-24: 129 (1870)
Facesoffungi number: FoF 10054
Saprobic on wood. Sexual morph: Ascomata 150–200 µm (x
̄ = 170 µm, n = 10) diam. at
venter, perithecial, gregarious, semi-immersed or erumpent through attached bark, globose to
subglobose, dark brown to black, glabrous or slightly hairy 3.5–5.5 µm wide, ostiolate, with long,
cylindrical necks 500–1500 × 40–70 µm, often curved. Peridium 25–50 µm, membranaceous,
comprising two layers, outer layer composed of dark brown to brown cells of textura angularis;
inner layer composed of hyaline cells of textura angularis. Asci 14–17(–19) × 11–15 µm (x
̄ = 16 ×
13 µm, n = 30) ascospore-bearing part, 4- or 8-spored, unitunicate, obovoid to clavate, with long
tapering pedicel, apical ring indistinct or absent, quickly evanescent. Ascospores 6–9 × 4.5–7 µm (x
̄
= 7.5 × 5.5 µm, n = 50), irregular or bi-seriate, aseptate, hyaline, ovoid to ellipsoidal, becoming
brown or dark brown, with rounded apex and truncated base with an obvious collapsed germ pore,
collapsing as a small, central, conical protuberance, smooth-walled, mostly with guttules. Asexual
morph: Undetermined.
Material examined – Germany, Hessen, Oestrich-Winkel (9° 0’ 19” E / 49° 46’ 28” N), on
decaying radices of Vitis vinifera, in winter, K.W.G. Fuckel (G-G00266143, holotype); Germany,
Saxony, at the churchyard of the Königstein fortress, on rotting roots of Humulus lupulus, 2 April
1886, K.W. Krieger (NY-03380967).
Known hosts and distribution – on decayed wood in Germany (type locality) (Fuckel 1870b).
Notes – We re-examined the type specimen, G00266143, and provided an illustration.
888
Figure 4 – Arxiomyces vitis: a, c, g–j, l–o, q (G-G00266143, holotype), k, p (Drawn following
observation by microscope); b, d–f (NY-03380967). a Material. b Gregarious ascomata on the host.
c Semi-immersed ascomata. d Ascoma with long neck. e Squashed neck. f Venter of ascoma cross
section. g Hairs at base. h Neck with fasciculate cells. i Irregular cells at venter of ascoma.
j–k Asci. l–q Ascospores (q stained in Melzer’s reagent). Scale bars: c–d = 500 µm, e–f = 200 µm,
h = 100 µm, g = 50 µm, i–l = 20 µm, m–q = 5 µm.
889
Dactylidispora Y. Marín, Stchigel, Guarro & Cano, in Marin-Felix et al., MycoKeys 44: 89 (2018)
Isolated from soil. Sexual morph: Ascomata perithecial or without an external opening,
superficial, globose to pyriform, yellow to brown, membranaceous, glabrous or setose, with or
lacking ostioles. Necks conical, with crown of setae surrounding the ostioles. Peridium composed
of yellow to pale brown cells of textura angularis, membranaceous. Asci 8-spored, unitunicate,
broadly clavate, evanescent. Ascospores irregular or bi-seriate, aseptate, hyaline becoming brown
to dark brown, broadly fusiform to citriform, umbonate and truncate at both ends, smooth-walled,
germ pore with a raised rim at each end, mostly with guttules. Asexual morph: Conidia hyaline,
subglobose to ovoid, smooth-walled (adapted from Marin-Felix et al. 2018).
Notes – Dactylidispora was introduced based on Sphaerodes ellipsospora and S.
singaporensis and, those two are sister to Melanospora kurssanoviana based on phylogenetic
results (Marin-Felix et al. 2018). This genus is characterized by citriform ascospores with a germ
pore surrounded by a raised rim at each end (Marin-Felix et al. 2018). These features are similar to
Pustulipora and Vittatispora; although Pustulipora has ascomata without neck and 4-spored asci
(Cannon 1982), and Vittatispora has a hyaline ridge between the germ pores on ascospores
(Chaudhary et al. 2006). In this study, Dactylidispora species clustered in Ceratostomataceae
(80%ML/1.00BY, Fig. 1).
Dactylidispora ellipsospora (Takada) Y. Marín, Stchigel, Guarro & Cano, in Marin-Felix et al.,
MycoKeys 44: 90 (2018)
Basionym – Microthecium ellipsosporum Takada, in Kobayasi et al., Bull. natn. Sci. Mus.,
Tokyo 16(3): 527 (1973)
Facesoffungi number: FoF 10145
Isolated from soil. Sexual morph: Ascomata superficial, globose, without an external opening,
yellow to brown, membranaceous. Peridium composed of yellow to pale brown cells of textura
angularis. Asci 8-spored, unitunicate, broadly clavate, evanescent. Ascospores irregular or bi-
seriate, aseptate, hyaline becoming brown to dark brown, broadly fusiform to citriform, umbonate
and truncate at both ends, smooth-walled, germ pore with a raised rim at each end, mostly with
guttules. Asexual morph: Undetermined (adapted from García et al. 2004, Marin-Felix et al. 2018).
Known hosts and distribution – on forest soil in Papua New Guinea (type locality) (Takada
1973).
Notes – Dactylidispora ellipsospora was initially named Microthecium ellipsosporum
(Takada 1973). Marin-Felix et al. (2018) sequenced its ex-type strain, NBRC 31376, and
established this species as the generic type of Dactylidispora. In this study, D. ellipsospora is sister
to D. singaporennsis (95%ML/1.00BY) in Ceratostomataceae (Fig. 1).
Echinusitheca Y. Marín, Stchigel, Dania García, Guarro, A.N. Mill. & Cano, in Marin-Felix et al.,
MycoKeys 44: 91 (2018)
Isolated from soil. Sexual morph: Ascomata superficial or immersed, globose, without an
external opening, pale brown to brown, semi-translucent, membranaceous, surrounded by setae.
Setae straight, pale brown to brown, 0–1-septate, verrucose to tuberculate, tapering, sometimes
branched. Peridium composed of brown to hyaline cells of textura angularis to textura globulosa.
Asci 8-spored, unitunicate, globose to subglobose, sessile. Ascospores irregular, aseptate, hyaline
becoming brown to dark brown, ellipsoidal, smooth-walled, with a germ pore at each end, mostly
with guttules. Asexual morph: Undetermined (adapted from Marin-Felix et al. 2018).
Notes – Marin-Felix et al. (2018) established the monotypic Echinusitheca typified by E.
citrispora. It is similar to Melanospora in having translucent ascomata and ellipsoidal ascospores;
however, the former has globose ascomata and globose to subglobose asci, whereas the latter has
ascomata with long necks and clavate asci (Zhang & Blackwell 2002, Marin-Felix et al. 2018).
It nests in Ceratostomataceae based on multi-gene phylogeny (Marin-Felix et al. 2018, this study,
Fig. 1).
890
Echinusitheca citrispora Y. Marín, Stchigel, Dania García, Guarro, A.N. Mill. & Cano, in Marin-
Felix et al., MycoKeys 44: 92 (2018)
Facesoffungi number: FoF 10146
Isolated from soil. Sexual morph: Ascomata superficial or immersed, globose, without an
external opening, pale brown to brown, semi-translucent, membranaceous, surrounded by setae.
Setae straight, pale brown to brown, 0–1-septate, verrucose to tuberculate, tapering, sometimes
branched. Peridium composed of brown to hyaline cells of textura angularis to textura globulosa.
Asci 8-spored, unitunicate, globose to subglobose, sessile. Ascospores irregular, aseptate, hyaline
becoming brown to dark brown, ellipsoidal, smooth-walled, with a germ pore at each end, mostly
with guttules. Asexual morph: Undetermined (adapted from Marin-Felix et al. 2018).
Known hosts and distribution – on forest soil in the USA (type locality) (Marin-Felix et al.
2018).
Notes – Marin-Felix et al. (2018) proposed that Echinusitheca citrispora is similar to
Arxiomyces and Scopinella species in having dark semi-translucent ascomata. However, E.
citrispora has smooth-walled, ellipsoidal ascospores, while Arxiomyces has ascospores with a
rounded apex and truncated base, and Scopinella has ascospores with two prominent longitudinal
germ slits (Marin-Felix et al. 2018). The type strain Echinusitheca citrispora (CBS 137837) nests
in Ceratostomataceae in the phylogenetic results (Marin-Felix et al. 2018, this study, Fig. 1).
Erythrocarpon Zukal, Verh. zool.-bot. Ges. Wien 35: 337 (1886)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial, globose, reddish
brown, membranaceous, surrounded by septate, filiform hyphae, ostiolate. Peridium composed of
brown to hyaline cells of textura angularis. Asci 8-spored, unitunicate, cylindrical. Ascospores uni-
seriate, hyaline to reddish brown, broadly fusiform to ellipsoidal, with germ pore at each end,
aseptate, smooth-walled, with guttules. Asexual morph (associated): Hyphomycetous.
Conidiophores mononematous, erect, hyaline to brown, septate, branched. Conidia hyaline to
brown, fusiform and 0–4-septate, or globose to subglobose and aseptate, or stellated, verrucose
(adapted from Zukal 1885).
Notes – The monotypic Erythrocarpon was introduced by Zukal (1885) and is characterized
by reddish brown, hairy and globose ascomata with broadly fusiform, reddish brown ascospores
with germ pore at each end (Zukal 1885). It has two types of accompanied asexual morph on
natural substrate, one has fusiform, septate conidia, and the other has globose or stellate, verrucose
conidia (Zukal 1885).
Type species – Erythrocarpon microstomum Zukal, Verh. zool.-bot. Ges. Wien 35: 337 (1886)
Fig. 8g–k
Facesoffungi number: FoF 10147
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary, superficial, globose, reddish
brown, membranaceous, surrounded by septate, filiform hyphae, ostiolate. Peridium composed of
hyaline to brown cells of textura angularis. Asci 8-spored, unitunicate, cylindrical. Ascospores uni-
seriate, hyaline to reddish brown, broadly fusiform to ellipsoidal, with germ pore at each end,
aseptate, smooth-walled, with guttules. Asexual morph: Hyphomycetous. Conidiophores
mononematous, erect, hyaline to brown, septate, branched. Conidia hyaline to brown, fusiform and
0–4-septate or globose to star-shaped and aseptate, verrucose (adapted from Zukal 1885).
Known hosts and distribution – on decayed wood in Austria (type locality) (Zukal 1885).
Notes – Erythrocarpon microstomum is similar to Melanospora zamiae in having ellipsoidal
to broadly fusiform ascospores, but the latter has translucent ascomata with a long neck; therefore,
it was considered as a member of Ceratostomataceae (Zukal 1885, Maharachchikumbura et al.
2016, Hyde et al. 2020e). There is no molecular data available for this species.
Gonatobotrys Corda, Pracht-Fl. Eur. Schimmelbild.: 9 (1839)
Saprobic on wood or vegetation. Sexual morph: Undetermined. Asexual morph:
891
Hyphomycetous. Mycelium composed of hyaline to light brown, superficial, effused, smooth-
walled, branched, septate hyphae. Conidiophores mononematous, macronematous, hyaline, erect,
septate, branched or unbranched, smooth-walled. Conidiogenous cells terminal to intercalary,
polyblastic, globose to subglobose, hyaline to pale brown, swollen. Conidia solitary, obovoid to
ellipsoidal, with a raised rim at base, 0–multi-septate, holoblastic, hyaline to pale brown, smooth to
verrucose, conidial secession with conspicuous denticles on the conidia and conidiogenous cells
(adapted from Hoch 1977, Walker & Minter 1981, Whaley & Barnett 1963).
Notes – Gonatobotrys was introduced by Corda (1839), and is well known as fungicolous
hyphomycetes of Alternaria and Fusarium species (Hoch 1977, Walker & Minter 1981, Whaley &
Barnett 1963). The type species Gonatobotrys simplex was considered as the asexual morph of
Melanospora damnosa, and was synonymised as M. simplex (Vakili 1989, Réblová et al. 2016).
However, Crous et al. (2020b) proposed that characters of hyphomycetous Gonatobotrys were
never observed in Melanospora cultures. Melanospora damnosa is distinct from G. simplex based
on phylogenetic results (Vu et al. 2019, Crous et al. 2020b). In this study, Gonatobotrys is sister to
Vittatispora (56%ML/0.97BY) in Ceratostomataceae, but distinct from Melanospora clade based
on multi-gene phylogeny (Fig. 1). Therefore, we accept Gonatobotrys as a genus in
Ceratostomataceae.
Gonatobotrys simplex Corda, Prachtflora: 9 (1839) Fig. 5
Facesoffungi number: FoF 10055
Saprobic on rotting vegetation. Sexual morph: Undetermined. Asexual morph:
Hyphomycetous. Mycelium composed of hyaline to light brown, superficial, effused, smooth-
walled, branched, septate hyphae 2–4 µm (x
̄ = 3, n = 10) wide. Conidiophores 5–8 µm (x
̄ = 6.5, n =
30) wide, mononematous, macronematous, hyaline, erect, septate, unbranched, smooth-walled.
Conidiogenous cells 9–14 µm (x
̄ = 10.5 µm, n = 20) diam., terminal to intercalary, polyblastic,
globose to subglobose, hyaline to pale brown, swollen. Conidia (12–)14–18(–22) × 8–12 µm (x
̄ =
16 × 10 µm, n = 50), solitary, obovoid to ellipsoidal, with a raised rim at base, aseptate, holoblastic,
hyaline to pale brown, smooth to verrucose, conidial secession with conspicuous denticles on the
conidia and conidiogenous cells (adapted from Vakili 1989).
Material examined – New Zealand, Auckland, on dry rootstock of Camelia sp., 27 April
1977, L.T.W. Mattson (PDD 47724).
Known hosts and distribution – on branches of Crataegus oxyacantha (Rosaceae) in
Germany (type locality) (Corda 1839); on dead wood or vegetation in New Zealand, South Africa
and USA (Vakili 1989, Crous et al. 2020b).
Notes – Gonatobotrys simplex was first collected on wood in Germany (Corda 1839) and
Melanospora damnosa was found on culm of wheat in Italy (Saccardo & Berlese 1896). Vakili
(1989) linked these two species based on the fact that both fungi were found on their corn-meal
agar. Thus, Réblová et al. (2016) reduced Gonatobotrys to a synonym of Melanospora.
Subsequently, these species were sequenced by Vu et al. (2019) and Crous et al. (2020b)
respectively. In this study, G. simplex and Vittatispora coorgii formed a clade (56%ML/0.97BY)
far from the Melanospora clade (Crous et al. 2020b, this study, Fig. 1). We could not obtain the
type specimen; therefore, we re-examined the material that has similar characteristics with the
description of G. simplex in Crous et al. (2020b).
Harzia Costantin, Mucéd. Simpl. (Paris): 42 (1888)
Saprobic on wood or leaves or isolated from soil. Sexual morph: Undetermined. Asexual
morph: Hyphomycetous. Mycelium composed of hyaline, smooth-walled, branched, septate hyphae.
Conidiophores mononematous, macronematous, hyaline, subcylindrical to cylindrical, erect, 0–
multi-septate, tapering, smooth-walled. Conidia terminal to intercalary, solitary, obovoid to
ellipsoidal, sometimes with a raised rim at base, aseptate, holoblastic, hyaline to brown, smooth to
verrucose (adapted from Saccardo 1886, Costantin 1888, Poursafar et al. 2017, Crous et al. 2019b).
Notes – The hyphomycetous Harzia was considered to be related to members of Hypocreales
892
(Maharachchikumbura et al. 2016, Wijayawardene et al. 2018), until Schultes et al. (2017) clarified
the species in this genus based on molecular information and accepted Harzia as a member in
Ceratostomataceae. In this study, Harzia species are sister to Melanospora (100%ML/1.00BY) in
Ceratostomataceae, except for Harzia velata which nests in Nectriaceae (Hypocreales)
(100%ML/1.00BY, Fig. 1).
Figure 5 – Gonatobotrys simplex: (PDD-47724). a Material label. b Dry culture. c Mycelium.
d Branched hyphae. e–g Conidiophore with conidia (g is strained in Congo red reagent). h Conidia.
Scale bars: d = 200 µm, e–g = 20 µm, h = 10 µm.
Harzia acremonioides (Harz) Costantin, Mucéd. Simpl. (Paris): 42 (1888)
Basionym – Monosporium acremonioides Harz, Linig. Hyph.: 17 (1872)
893
Facesoffungi number: FoF 10148
Saprobic on wood or leaves. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium composed of hyaline, smooth-walled, branched, septate hyphae. Conidiophores
mononematous, macronematous, hyaline, subcylindrical to cylindrical, straight or slightly curved,
erect, 1–2-septate, tapering, smooth-walled. Conidia terminal, solitary, obovoid to ellipsoidal,
aseptate, holoblastic, hyaline to brown, smooth to verrucose (adapted from Saccardo 1886,
Poursafar et al. 2017).
Known hosts and distribution – on diseased branches and leaves in Austria and Germany
(type locality) (Saccardo 1886); on dune sand in France (Schultes et al. 2017); on diseased wheat
and barley in Iran (Poursafar et al. 2017).
Notes – The generic type, Harzia acremonioides, was initially collected on diseased branches
and leaves in Austria and Germany (Saccardo 1886). Schultes et al. (2017) sequenced the epitype
strain of Monosporium acremonioides (≡ H. acremonioides) and defined this species in
Ceratostomataceae. In our study, H. acremonioides is sister to H. verrucosa (84%ML/0.97BY,
Fig. 1).
Melanospora Corda, Icon. fung. (Prague) 1: 24 (1837)
Saprobic on vegetation or dung or wood or parasitic on other fungi or closely associated with
other fungi. Sexual morph: Ascomata perithecial, solitary, superficial to immersed, globose venter,
usually with long necks, yellow to reddish brown, membranaceous, translucent or semi-translucent,
glabrous, tomentose or hairy, ostiolate. Necks cylindrical to conical or absent, composed of
fasciculate, cylindrical, pale yellow, septate hyphae, straight or slightly curved, with a jagged apex,
ascospores usually accumulating in translucent necks. Peridium membranaceous, composed of
yellow to pale brown cells of textura angularis. Asci 4- or 8-spored, unitunicate, obovoid to broadly
clavate, evanescent. Ascospores irregular or bi-seriate, hyaline becoming brown or reddish brown,
broadly fusiform, ellipsoidal to citriform, umbonate and truncate at both ends, aseptate, smooth-
walled, verrucose or with reticulate ribs on surface, usually laterally collapsing, with a germ pore at
each end, mostly with guttules. Asexual morph (associated): Hyphomycetous. Mycelium consisting
of hyaline to pale brown, branched, septate, smooth-walled hyphae. Conidiophores hyaline,
cylindrical, straight, curved or flexuous, septate, branched or unbranched. Conidia hyaline to
brown, globose to ovoid, 0–1-septate (adapted from Rehner & Samuels 1995, Seifert et al. 2011,
Schultes et al. 2017, Marin-Felix et al. 2018).
Notes – Melanospora was introduced by Corda (1837) and its asexual morph was considered
as Gonatobotrys and Harzia (Vakili 1989, Seifert et al. 2011, Schultes et al. 2017). However, Crous
et al. (2020b) suggested that these asexual morphs have never been found on the culture of
Melanospora. In this study, Melanospora is sister to Harzia (100%ML/1.00BY), and distinct from
Gonatobotrys based on phylogenetic result (Fig. 1). Therefore, these hyphomycetes are regarded as
associated genera of Melanospora, until the cultural characteristics of Melanospora or molecular
information is available to determine their relationships.
Melanospora zamiae Corda, Icon. fung. (Prague) 1: 24 (1837) Fig. 6
Facesoffungi number: FoF 10056
Saprobic on rotting vegetation or dung. Sexual morph: Ascomata 95–240 µm (x
̄ = 180 µm, n
= 10) diam., perithecial, solitary, superficial, globose venter with a long neck, yellow to reddish
brown, membranaceous, semi-translucent, glabrous or slightly hairy, ostiolate. Necks 200–300 ×
35–50 µm (x
̄ = 245 × 42 µm, n = 10), cylindrical, composed of fasciculate, cylindrical, pale yellow,
septate hyphae 2–3 µm wide, straight or slightly curved, with a jagged apex, ascospores usually
accumulating in translucent necks. Peridium composed of yellow to pale brown cells of textura
angularis to textura globulosa, membranaceous. Asci 4- or 8-spored, unitunicate, obovoid to
broadly clavate, evanescent. Ascospores 20–23 × (12–)14–15(–17) µm (x
̄ = 22 × 14.5 µm, n = 50),
irregular or bi-seriate, hyaline becoming brown or reddish brown, broadly fusiform to ellipsoidal,
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aseptate, smooth-walled, usually laterally collapsing, umbonate and truncate at each end, with germ
pore at both ends, mostly with guttules. Asexual morph: Undetermined.
Figure 6 – Melanospora zamiae: a, g–h, p–q (UPS-UPS:BOT:F-129707); b, d–e, i–m, r–u (PDD
36919); c, f (UPS-UPS:BOT:F-129706); n–o (drawn from Cannon & Hawksworth 1982). a–b Two
types of host. c–d Superficial ascomata on the host. e–g Squashed ascomata with long neck. h Apex
of neck with fasciculate hyphae. i Neck with a jagged apex. j Neck composed of cylindrical cells. k
Cells at venter of ascoma. l–o Asci. p–u Ascospores (t ascospore laterally collapsing). Scale bars:
d–g = 200 µm, c, h = 100 µm, i–j, m = 50 µm, k–l, n–o = 20 µm, p–u = 10 µm.
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Material examined – Sweden, Öland, Resmo par., 4 km E of the church on the Great Alvar,
on cow dung in moist chamber in Uppsala, 2 June 1968, R. Santesson (UPS-UPS:BOT:F-129706);
Sweden, Öland, Räpplinge, Borgholm, at the Castle ruin, on old horse dung in moist chamber, 30
August 1959, N. Lundqvist (UPS-UPS:BOT:F-129707); New Zealand, Auckland, on leaves of
Microlaena stipoides, 14 April 1975, J.M. Dingley (PDD 36919).
Known hosts and distribution – on decayed leaves of Zamia pungens (Zamiaceae) in
Germany (type locality) (Corda 1837); on dung or leaves in Israel, New Zealand and Sweden (Vu
et al. 2019).
Notes – The generic type Melanospora zamiae has been considered as conspecific with M.
damnosa (Cannon & Hawksworth 1982, Réblová et al. 2016). However, these two species exist
independently in Melanospora based on phylogenetic results (Marin-Felix et al. 2018, Vu et al.
2019). Figure 1 reveals that Melanospora species clustered as a distinct clade (100%ML/1.00BY),
except M. kurssanoviana which is basal to Harzia, Melanospora and Microthecium
(57%ML/1.00BY, Fig. 1). We were unable to obtain the type specimen, therefore, we re-examined
authentic materials collected by Lundqvist (UPS:BOT:F-129707) or determined by Samuels
(36919, mentioned in the label of material).
Microthecium Corda, Icon. fung. (Prague) 5: 30, 74 (1842)
Saprobic on wood or leaves or isolated from soil, sometimes fungicolous. Sexual morph:
Ascomata perithecial or without an external opening, superficial or immersed, globose to
subglobose, scattered or gregarious, yellowish to brown, membranaceous, translucent, glabrous,
tomentose or hairy, necks short or absent, with or lacking ostioles. Peridium usually translucent or
semi-translucent, membranaceous, composed of yellow to pale brown cells of textura angularis.
Asci 4- or 8-spored, clavate to pyriform, evanescent. Ascospores irregular or bi-seriate, hyaline
becoming brown to black, ellipsoidal to citriform, usually umbonate and truncate at both ends,
smooth-walled or verrucose or with reticulate ribs on surface, with a germ pore at each end, mostly
with guttules. Sexual morph: Conidiophores unbranched or branched, cylindrical, septate. Conidia
subglobose to oval, hyaline to pale brown, smooth-walled. Bulbils globose to ellipsoidal, pale
brown to reddish brown (adapted from Udagawa & Cain 1969, Marin-Felix et al. 2018).
Notes – Microthecium was introduced by Corda (1842) and typified by M. zobelii which was
found growing in the hymenial tissue of Rhizopogon albus. This genus has translucent ascomata
and ellipsoidal to citriform ascospores with umbonate and truncated ends, simliar to Melanospora
and Sphaerodes (Winter 1887b, Cannon & Hawksworth 1982). Therefore, Microthecium was
regarded as a synonym of Melanospora, until Marin-Felix et al. (2018) sequenced several species
in these two genera, and Microthecium species existed in Ceratostomataceae as a clade independent
of Melanospora. Thus, Microthecium was re-accepted as a genus in Ceratostomataceae (Hyde et al.
2020e, Wijayawardene et al. 2020). In this study, Microthecium species clustered
(86%ML/0.92BY) and are basal to Harzia and Melanospora (57%ML/1.00BY) in
Ceratostomataceae (Fig. 1).
Microthecium zobelii Corda, Icon. fung. (Prague) 5: 74 (1842) Fig. 8l
Facesoffungi number: FoF 10149
Saprobic on dung or isolated from soil, or fungicolous. Sexual morph: Ascomata superficial
or immersed, globose to subglobose, without an external opening, scattered or gregarious,
yellowish to brown, membranaceous, translucent, glabrous or hairy. Peridium membranaceous,
composed of yellow to pale brown cells of textura angularis. Asci 8-spored, clavate, evanescent.
Ascospores irregular or bi-seriate, hyaline becoming brown to black, ellipsoidal to citriform,
usually umbonate and truncate at both ends, smooth-walled, with a germ pore at each end, mostly
with guttules. Sexual morph: Conidia subglobose to oval, hyaline to pale brown, smooth-walled
(adapted from Udagawa & Cain 1969, Marin-Felix et al. 2018).
Known hosts and distribution – on Rhizopogon albus in Germany (type locality) (Corda
1842); on Choiromyces maeandriformis in Czech Republic (Udagawa & Cain 1969); on decayed
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Coriolus flabelliformis in Papua New Guinea (Marin-Felix et al. 2018, Wang et al. 2019a); on roe
dung in Netherlands (Wang et al. 2019a); on soil in Japan (Hawksworth & Udagawa 1977).
Notes – Microthecium zobelii has ascomata without an external opening and ellipsoidal to
citriform ascospores with a terminal apiculate or depressed germ pore at each end (Marin-Felix et
al. 2018). In this study, M. zobelii (CBS 341.73) nests in Microthecium based on the phylogenetic
result (Fig. 1).
Neotrotteria Sacc., Bulletino dell'orto Botanico della R. Universitá di Napoli 6: 45 (1921)
Saprobic on decorticated wood. Sexual morph: Subiculum extensive, forming large and dense
patches, composed of densely clustered, abundant, brown to black, septate, branched, spinulose
hyphae with dichotomously branched, tapering ends. Ascomata superficial, collabent, cupulate,
turbinate, tuberculate, scattered, black, carbonaceous, lacking ostioles, surrounded by rigid,
straight, tapering setae. Peridium centrum with an obconical Quellkörper, comprising three layers,
outer layer composed of carbonaceous, black tissues; middle layer composed of membranaceous,
olivaceous to brown cells of textura angularis, Munk pores present; inner layer composed of
membranaceous, hyaline cells of textura prismatica. Asci multi-spored, unitunicate, cylindrical to
clavate, evanescent. Ascospores irregularly arranged, hyaline, allantoid to oval, slightly curved,
smooth-walled, aseptate, mostly with guttules. Asexual morph: Undetermined.
Notes – The monotypic Neotrotteria was introduced by Saccardo (1918) and typified by N.
pulchella in having carbonaceous ascomata with an obconical Quellkörper and clavate asci with
numerous spores. These characters were considered similar to Acanthonitschkea (Nitschkiaceae),
thus, N. pulchella was transferred as A. pulchella (Nannfeldt 1975, Nannfeldt & Santesson 1975).
Subsequently, Mugambi & Huhndorf (2010) analyzed LSU sequence data and found that this
species is basal to other Scortechiniaceae species. In this study, Neotrotteria is morphologically
similar to genera in Scortechiniaceae that have cupulate ascomata with Quellkörper, however,
phylogenetically, this genus nests in Ceratostomataceae based on LSU sequence (83%ML, not
show) and multi-gene analyses (68%ML, Fig. 1). Therefore, Neotrotteria has an existence between
Ceratostomataceae and Scortechiniaceae. However, we place Neotrotteria in Ceratostomataceae
based on our phylogenetic result (Fig. 1).
Neotrotteria pulchella Sacc., Bulletino dell'orto Botanico della R. Universitá di Napoli 6: 45
(1918) Fig. 7
Facesoffungi number: FoF 10057
Saprobic on decorticated wood. Sexual morph: Subiculum extensive, forming large and dense
patches, composed of densely clustered, abundant, brown to black, septate, branched, spinulose
hyphae 4.5–7 µm thick, with dichotomously branched, tapering ends. Ascomata 300–400 × 325–
400 µm (x
̄ = 340 × 360 µm, n = 5), superficial, collabent, cupulate, turbinate, tuberculate, scattered,
black, carbonaceous, lacking ostioles, with rigid, straight, tapering setae 7–13 µm thick. Peridium
centrum with an obconical Quellkörper 65–110 × 140–200 µm (x
̄ = 85 × 160 µm, n = 10),
comprising three layers 45–70 µm thick, outer layer composed of carbonaceous, black tissues;
middle layer composed of membranaceous, olivaceous to brown cells of textura angularis, Munk
pores present; inner layer composed of membranaceous, hyaline cells of textura prismatica. Asci
multi-spored, unitunicate, cylindrical to clavate, evanescent. Ascospores 3.5–5 × 1–2 µm (x
̄ = 4 ×
1.2 µm, n = 50), irregularly arranged, hyaline, allantoid to oval, slightly curved, smooth-walled,
aseptate, mostly with guttules. Asexual morph: Undetermined.
Material examined – China, Taiwan, Kukuan, on stems of Lycium chinense, 17 July 1990,
C.Y. Chen (IMI 354793).
Known hosts and distribution – on decayed wood of rubber tree (Hevea brasiliensis) in
Singapore (type locality) (Saccardo 1918); on stems of Lycium chinense (Solanaceae) in China; on
dead wood in Kenya (Mugambi & Huhndorf 2010).
Notes – Neotrotteria pulchella is characterized by carbonaceous ascomata that have
obconical Quellkörper and surrounded by tapering setae, multi-spored asci and oval to allantoid
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ascospores (Saccardo 1918). Subsequently, this species was considered as a member of
Nitschkiaceae or Coronophoraceae (Nannfeldt 1975, Nannfeldt & Santesson 1975, Subramanian &
Sekar 1990). Mugambi & Huhndorf (2010) clarified the genera in Coronophorales and found that
N. pulchella groups together with Scortechiniaceae based on LSU sequence analysis. In this study,
N. pulchella nests in Ceratostomataceae based on phylogenetic result (68%ML, Fig. 1). We were
unable to obtain the type material. Therefore, we re-examined an authentic specimen, 354793,
which was determined by Sivanesan (mentioned in the label of material).
Figure 7 – Neotrotteria pulchella: (IMI-354793). a Ascomata on decorticated wood.
b–c Ascomata. d–e Ascomata cross section. f Quellkörper. g Peridium (arrow indicates Munk pores
between cells). h Seta. i Hypha with dichotomously branched end. j–k Asci. l Ascospores. Scale
bars: b = 2 mm, c = 500 µm, d–e = 200 µm, f = 100 µm, g–i = 50 µm, j–k = 20 µm, l = 10 µm.
898
Pseudomicrothecium Y. Marín, Stchigel, Guarro & Cano, in Marin-Felix et al., MycoKeys 44: 114
(2018)
Saprobic on Tuber. Sexual morph: Ascomata immersed, globose to subglobose, without an
external opening, scattered, pale brown to brown, membranaceous, translucent, glabrous or setose.
Peridium membranaceous, composed of pale brown to hyaline cells of textura angularis. Asci 2-
spored, clavate to cylindrical, evanescent. Ascospores hyaline becoming dark brown, ellipsoidal to
citriform, usually umbonate and truncate at both ends, smooth-walled, with a terminal indistinct
germ pore at each end, mostly with guttules. Sexual morph: Undetermined (adapted from Fan et al.
2012).
Notes – The monotypic Pseudomicrothecium was established by Marin-Felix et al. (2018)
and is characterized by ascomata without an external opening and 2-spored asci with ellipsoidal to
citriform ascospores (Fan et al. 2012). In this study, P. subterraneum nests in Ceratostomataceae
(80%ML/1.00BY, Fig. 1) based on phylogenetic result.
Pseudomicrothecium subterraneum (L. Fan et al.) Y. Marín, Stchigel, Guarro & Cano, in Marin-
Felix et al., MycoKeys 44: 115 (2018)
Basionym – Melanospora subterranea L. Fan, C.L. Hou, P.F. Cannon & Yu Li, Mycologia
104(6): 1434 (2012)
Facesoffungi number: FoF 10150
Saprobic on Tuber. Sexual morph: Ascomata immersed, globose to subglobose, without an
external opening, scattered, pale brown to brown, membranaceous, translucent, glabrous or setose.
Peridium membranaceous, composed of pale brown to hyaline cells of textura angularis. Asci 2-
spored, clavate to cylindrical, evanescent. Ascospores hyaline becoming dark brown, ellipsoidal to
citriform, usually umbonate and truncate at both ends, smooth-walled, with a terminal indistinct
germ pore at each end, mostly with guttules. Sexual morph: Undetermined (adapted from Fan et al.
2012).
Known hosts and distribution – on decayed tissues of Tuber huidongense and T. indicum in
China (type locality) (Fan et al. 2012).
Notes – Marin-Felix et al. (2018) transferred Melanospora subterranea as generic type of
Pseudomicrothecium based on phylogenetic result.
Pustulipora P.F. Cannon, Mycotaxon 15: 526 (1982)
Saprobic on wood. Sexual morph: Ascomata perithecial, superficial, globose to subglobose,
gregarious, brown to dark brown, membranaceous, covered by white tomentum, ostiolate. Peridium
membranaceous, composed of yellow to pale brown cells of textura angularis. Asci 4-spored,
clavate to obovoid, evanescent. Ascospores bi-seriate, brown, ellipsoidal to citriform, usually
umbonate and truncate at both ends, smooth-walled, with a germ pore at each end, sometimes
surrounded by hyaline cushion-like structures. Sexual morph: Undetermined (adapted from Cannon
1982).
Notes – The monotypic genus Pustulipora was introduced by Cannon (1982) and typified by
P. corticola. This species was considered as Melanospora theleboloides (Fig. 8a–d). However,
Cannon (1982) stated that P. corticola is different from M. theleboloides with glabrous ascomata,
whereas, the former has tomentose ascomata.
Pustulipora corticola P.F. Cannon, Mycotaxon 15: 526 (1982) Fig. 8e–f
Facesoffungi number: FoF 10151
Saprobic on wood or leaves. Sexual morph: Ascomata perithecial, superficial, globose to
subglobose, gregarious, brown to dark brown, membranaceous, covered by white tomentum,
ostiolate. Peridium membranaceous, composed of yellow to pale brown cells of textura angularis.
Asci 4-spored, clavate to obovoid, evanescent. Ascospores bi-seriate, brown, ellipsoidal to
citriform, usually umbonate and truncate at both ends, smooth-walled, with a germ pore at each
899
end, sometimes surrounded by hyaline cushion-like structures. Sexual morph: Undetermined
(adapted from Cannon 1982).
Known hosts and distribution – on decayed wood in the UK (type locality) (Cannon 1982).
Notes – We re-examined the type of Melanospora theleboloides (Germany, Hessen,
Rheingau, in the castle park near Hattenheim, on decaying leaves, K.W.G. Fuckel, G-326774/1-
G00127866, holotype), but its ascomata are too dry to observe any complete structure. However,
we drew the ascus and ascospores of Pustulipora corticola with reference to Cannon (1982).
Figure 8 – Melanospora theleboloides: a–d (G-326774/1-G00127866, holotype). a Material.
b Crushed ascomata. c Immature asci. d ascospores; Pustulipora corticola: e–f (redrawn from
Cannon 1982). e Ascus. f Ascospores; Erythrocarpon microstomum: g–k (redrawn from Zukal
1885). g Ascoma cross section. h Ascus. i Ascospores. j–k Two types of conidiophores with
conidia; Microthecium zobelii: l (redrawn from Corda 1842, Udagawa & Cain 1969). l Ascospores.
Microthecium foveolatum: m–n (redrawn from Hawksworth & Udagawa 1977). m Ascospores with
irregular ridges. n Conidiophores with conidia. Scale bars: g = 200 µm, b = 100 µm, e, k = 50 µm,
c–d, f, h, j, l = 20 µm, i, m–n = 10 µm.
Rhytidospora Jeng & Cain, Mycotaxon 5(1): 278 (1977)
Coprophilous. Sexual morph: Ascomata solitary, superficial, globose, without an external
opening, yellow to pale orange, membranaceous, translucent, glabrous or hairy. Peridium
cephalothecoid, irregularly polygonous pattern on surface view, membranaceous, outer layer
900
composed of yellow to pale orange cells of textura angularis; inner layer composed of hyaline cells
of textura prismatica. Paraphyses absent. Asci 2-, 4- or 8-spored, unitunicate, subglobose to
clavate, evanescent. Ascospores hyaline becoming dark brown or olivaceous brown, broadly
fusiform to ellipsoidal, aseptate, verrucose to tuberculate to wrinkled, sometimes with pits on
surface, with germ pore at each end and a large guttule. Asexual morph: Undetermined (adapted
from Jeng & Cain 1977, Krug & Jeng 1979, Valldosera & Guarro 1994).
Notes – Rhytidospora was established by Jeng & Cain (1977) and typified by Rhytidospora
tetraspora. It is characterized by translucent, glabrous ascomata and broadly fusiform ascospores
with wrinkled or pitted surface (Jeng & Cain 1977, Krug & Jeng 1979, Valldosera & Guarro 1994).
These characters were considered similar to Melanospora (Jeng & Cain 1977), and we found that
they are also similar to Neurospora and Sordaria. However, molecular data for this genus is
lacking.
Rhytidospora tetraspora Jeng & Cain, Mycotaxon 5(1): 279 (1977) Fig. 9
Facesoffungi number: FoF 10058
Coprophilous. Sexual morph: Ascomata 130–250 µm (x
̄ =185 µm, n = 10) diam., solitary,
superficial, globose, without an external opening, yellow to pale orange, membranaceous,
translucent, glabrous. Peridium 45–80 µm (x
̄ = 60 µm, n = 20), cephalothecoid, irregularly
polygonous pattern on surface view, membranaceous, comprising three layers, outer layer
composed of yellow to pale orange cells of textura globulosa; middle layer composed of yellow to
pale orange cells of textura angularis; inner layer composed of hyaline cells of textura prismatica.
Asci (21–)25–32(–35) × (14–)18–20(–23) µm (x
̄ = 28 × 19 µm, n = 10) ascospore-bearing part, 4-
spored, unitunicate, obovoid to clavate, evanescent. Ascospores 9.5–14(–18) × (7–)8–10(–12.5) µm
(x
̄ = 13 × 9 µm, n = 50), bi-seriate, hyaline becoming dark brown or olivaceous brown, broadly
fusiform to ellipsoidal, aseptate, verrucose to tuberculate, with germ pore at each end and a large
guttule. Asexual morph: Undetermined.
Material examined – Venezuela, Edo. Scure, trail between Manacal and Los Pocitos, NW of
Irapa, 9°05’00’’N/62°41’00’’W, on burro dung, 13 July 1972, K.P. Dumont, R.F. Cain, G.J.
Samuels, G. Morillo and J. Farfan (TRTC-052086, holotype); Sweden, Uppland, Häverö,
Prästăngen, in a grove of deciduous trees, on old horse dung in moist chamber, 17 June 1976, N.
Lundqvist (UPS-UPS:BOT:F-132032).
Known hosts and distribution – on burro dung in Venezuela (type locality) (Jeng & Cain
1977); on horse dung in Sweden.
Notes – We re-examined the holotype (052086) and an authentic specimen (F-132032) which
was collected by Lundqvist, and provide their illustration.
Scopinella Lév., in Orbigny, Dict. Univ. Hist. Nat. 8: 493 (1846)
Saprobic on wood, leaves and/or isolated from soil, or coprophilous or fungicolous. Sexual
morph: Ascomata perithecial, solitary to gregarious, superficial to semi-immersed, sometimes
sitting on a dense mass of brown hyphae, globose to subglobose venter, usually with long necks,
brown to dark brown, semi-translucent, membranaceous, glabrous or hairy, ostiolate. Peridium
membranaceous, composed of brown cells of textura angularis. Paraphyses absent or filiform,
septate. Asci 2-, 8- to multi-spored, clavate to cylindrical. Ascospores olivaceous brown to dark
brown, ellipsoidal, cuboid-ellipsoidal to Z-shaped, verrucose or smooth-walled, usually with two
prominent longitudinal germ slits. Sexual morph: Undetermined (adapted from Hawksworth 1975,
van Geel & Aptroot 2006, Pastirčák & Pastirčáková 2007, Sanchez et al. 2012).
Notes – Scopinella was introduced by Léveillé (1847) based on S. barbata having ascomata
with long necks, 8-spored asci and cuboid-ellipsoidal ascospores. Saccardo (1891) proposed a
second species based on Melanospora pleiospora, which has multi-spored asci and ellipsoidal
ascospores. However, Hawksworth (1975) clarified Scopinella and did not include S. pleiospora.
Seven Scopinella species with cuboid-ellipsoidal ascospores were subsequently introduced
(Hawksworth 1975, Cannon & Hawksworth 1982, van Geel & Aptroot 2006, Pastirčák &
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Pastirčáková 2007, Sanchez et al. 2012, Marin-Felix et al. 2018, Index Fungorum 2021). Zhang &
Blackwell (2002) sequenced S. solani (CBS 770.84) which nests in Stachybotryaceae
(Hypocreales) in our phylogenetic study (Fig. 1). This genus was placed in Ceratostomataceae
because of its ascomata with long necks (Pastirčák & Pastirčáková 2007, Wijayawardene et al.
2020, Hyde et al. 2020e). Lumbsch & Huhndorf (2010) placed this genus in Hypocreales. In this
study, we propose to place Scopinella in Ceratostomataceae pending molecular data of type
species.
Figure 9 – Rhytidospora tetraspora: a–b, d–g, i–o (TRTC-052086, holotype); c, h, p–q (UPS-
UPS:BOT:F-132032). a–c Superficial ascomata on the host. d Ascoma cross section. e Peridium
902
surface view. f–g Asci. h–q Ascospores (p arrow indicates germ pore). Scale bars: a–b = 200 µm,
c–e = 100 µm, f = 20 µm, g–h, p–q = 10 µm, i–o = 5 µm.
Scopinella barbata (Pers.) Lév. ex Sacc., Michelia 1(no. 3): 284 (1878) Fig. 15g–h
Facesoffungi number: FoF 10152
Saprobic on leaves. Sexual morph: Ascomata perithecial, solitary to scattered, superficial to
semi-immersed, globose to subglobose venter, with a long neck, dark brown to black, semi-
translucent, membranaceous, hairy, ostiolate. Necks cylindrical to conical, composed of fasciculate,
cylindrical, brown to dark brown, septate hyphae, straight or slightly curved, with a jagged apex,
ascospores usually accumulating in necks. Peridium membranaceous, composed of brown cells of
textura angularis. Paraphyses absent. Asci 8-spored, unitunicate, clavate, with short pedicel.
Ascospores irregular to bi-seriate, brown to dark brown, cuboid-ellipsoidal, smooth-walled, with
two prominent longitudinal germ slits. Sexual morph: Undetermined (adapted from Hawksworth
1975)
Known hosts and distribution – on fallen leaves of Quercus, Castanea, Potentilla and
Rhododendron in Algeria, Canada, France, Germany (type locality), Italy, Switzerland, the UK and
Pakistan (Hawksworth 1975, van Geel & Aptroot 2006).
Notes – Scopinella barbata was introduced as type species (Léveillé 1847, Saccardo 1878).
This genus was ignored until Hawksworth (1975) clarified Chaetoceratostoma hispidum as a
synonym of S. barbata based on their authentic materials. However, molecular data for this species
is unavailable. We were unable to obtain authentic material; therefore, a hand-drawing of ascus and
ascospores are provided (Fig. 15g–h) based on Hawksworth (1975).
Setiferotheca Matsush., Matsush. Mycol. Mem. 8: 34 (1995)
Saprobic on litter in forest. Sexual morph: Subiculum white, composed of hyaline, septate
hyphae. Ascomata perithecial, superficial, gregarious to solitary, globose to subglobose, pale brown
to brown, translucent, membranaceous, with inconspicuous ostioles, surrounded by straight, rigid,
tapering, dark brown to black setae. Peridium membranaceous, composed of pale brown cells of
textura angularis. Asci 4- to 8-spored, clavate to cylindrical, evanescent. Ascospores usually bi-
seriate, brown, ellipsoidal, smooth-walled, rounded apex, terminal apiculate base with germ pore.
Sexual morph: Undetermined (adapted from Matsushima 1995).
Notes – Setiferotheca was established as a genus related to Sphaeroderma (= Melanospora),
although the former has ascospores with a germ pore at one end, and the latter has one germ pore at
each end (Matsushima 1995). However, no molecular data is available for the Setiferotheca.
Setiferotheca nipponica Matsush., Matsush. Mycol. Mem. 8: 35 (1995) Fig. 15k–m
Facesoffungi number: FoF 10153
Saprobic on litter in forest. Sexual morph: Subiculum white, composed of hyaline, septate
hyphae. Ascomata perithecial, superficial, gregarious to solitary, globose to subglobose, pale brown
to brown, translucent, membranaceous, with inconspicuous ostioles, surrounded by straight, rigid,
tapering, dark brown to black setae. Peridium membranaceous, composed of pale brown cells of
textura angularis. Asci 4- to 8-spored, clavate to cylindrical, evanescent. Ascospores usually bi-
seriate, brown, ellipsoidal, smooth-walled, rounded apex, terminal apiculate base with germ pore.
Sexual morph: Undetermined (adapted from Matsushima 1995).
Known hosts and distribution – on litter in forest in Japan (type locality) (Matsushima 1995).
Notes – The monotypic Setiferotheca is typified by S. nipponica, and this species was found
in the litter in forests of Ise-shi, Amami-ohshima and Minakuchi-choh in Japan from 1993 to 1995
(Matsushima 1995).
Syspastospora P.F. Cannon & D. Hawksw., J. Linn. Soc., Bot. 84(2): 152 (1982)
Parasitic on various moniliaceous hyphomycetes, or isolated from soil, or lichenicolous.
Sexual morph: Ascomata perithecial or without an external opening, solitary, superficial to
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immersed, globose to subglobose, sometimes with long necks, pale brown to brown,
membranaceous, glabrous or setose or hairy, with or lacking ostioles. Necks absent or cylindrical,
composed of fasciculate, cylindrical, septate hyphae, straight or slightly curved. Peridium
membranaceous, composed of pale brown to brown cells of textura angularis. Asci 8-spored,
unitunicate, clavate, evanescent. Ascospores irregular or bi-seriate, hyaline becoming brown or
olivaceous brown, broadly fusiform to cylindrical to doliiform, truncate at both ends, aseptate,
smooth-walled or with reticulate ribs on surface, with a large depressed germ pore at each end,
mostly with guttules. Asexual morph: Hyphomycetous. Mycelium consisting of hyaline to
yellowish brown, branched, septate, smooth-walled hyphae. Chlamydospores terminal to
intercalary, globose to ovoid, hyaline to brown (adapted from Cannon & Hawksworth 1982, Horie
et al. 1986, García et al. 2002, Etayo & Sancho 2008).
Notes – Syspastospora was established as a genus similar to Melanospora, and its generic
type, S. parasitica, has cylindrical to doliiform ascospores with large terminal depressed germ
pores at each end, whereas Melanospora has ellipsoidal to citriform ascospores (Cannon &
Hawksworth 1982). The strain of S. parasitica nested in Hypocreales (Zhang & Blackwell 2002).
This genus was accepted as a member of Ceratostomataceae based on multi-gene analyses (Marin-
Felix et al. 2018, Hyde et al. 2020e, Wijayawardene et al. 2020). In this study, Syspastospora is
sister to Gonatobotrys and Vittatispora in Ceratostomataceae (54%ML/0.99BY, Fig. 1).
Syspastospora parasitica (Tul.) P.F. Cannon & D. Hawksw., J. Linn. Soc., Bot. 84(2): 152 (1982)
Fig. 15f
Basionym – Sphaeronaema parasiticum Tul. [as 'parasitica'], Annls Sci. Nat., Bot., sér. 4 8:
40 (1857)
Facesoffungi number: FoF 10154
Parasitic on various moniliaceous hyphomycetes. Sexual morph: Ascomata perithecial,
solitary, superficial to immersed, globose to subglobose, with long necks, pale brown to brown,
membranaceous, glabrous or hairy, ostiolate. Necks cylindrical, composed of fasciculate,
cylindrical, septate hyphae, straight or slightly curved. Peridium membranaceous, composed of
pale brown to brown cells of textura angularis. Asci 8-spored, unitunicate, clavate, evanescent.
Ascospores irregular or bi-seriate, hyaline becoming brown or olivaceous brown, cylindrical to
doliiform, truncate at both ends, aseptate, smooth-walled or with reticulate ribs on surface, with a
depressed germ pore at each end, mostly with guttules. Asexual morph: Undetermined (adapted
from Cannon & Hawksworth 1982).
Known hosts and distribution – on various moniliaceous hyphomycetes, e.g. Beauveria,
Cephalosporium, Paecilomyce, Hirsutella and Verticillium in Canada, Europe, Kenya, South
Africa, Sri Lanka, Trinidad and the USA (Cannon & Hawksworth 1982).
Notes – Syspastospora parasitica was initially introduced as Sphaeronema parasiticum, and
then transferred as Melanospora parasitica based on its ascomata with long hairy necks and
ascospores with truncate ends being similar to Melanospora (Tulasne 1857, Tulasne & Tulasne
1865). However, Sys. parasitica with cylindrical to doliiform ascospores differs from Melanospora
species with ellipsoidal to citriform ascospores (Cannon & Hawksworth 1982). In this study, Sys.
parasitica is basal to Gonatobotrys simmplex and Vittatispora coorgii (54%ML/0.99BY, Fig. 1) in
Ceratostomataceae.
Vittatispora P. Chaudhary, J. Campb., D. Hawksw. & K.N. Sastry, Mycologia 98(3): 461 (2006)
Isolated from soil. Sexual morph: Ascomata perithecial, solitary or gregarious, superficial to
semi-immersed, globose, with long necks, pale brown to brown, membranaceous, translucent,
glabrous or hairy, ostiolate. Necks cylindrical, composed of fasciculate, cylindrical, pale brown
hyphae, straight or slightly curved, ascospores usually accumulating in translucent necks. Peridium
membranaceous, composed of brown cells of textura angularis. Asci 8-spored, unitunicate, clavate,
evanescent. Ascospores hyaline becoming brown or dark brown, ellipsoidal to citriform, apiculate,
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aseptate, smooth-walled, with germ pore surrounded by a raised rim and connected by a prominent
longitudinal ridge. Asexual morph: Undetermined (adapted from Chaudhary et al. 2016).
Notes – The monotypic genus Vittatispora (type V. coorgii) has translucent ascomata with
ellipsoidal to citriform ascospores with germ pores surrounded by a raised rim and connected by a
prominent longitudinal ridge (Chaudhary et al. 2006). These characters are similar to Melanospora.
However, Chaudhary et al. (2006) analyzed LSU sequence data and proved that Vittatispora is
independent of Melanospora. In this study, Vittatispora is sister to Gonatobotrys in
Ceratostomataceae (56%ML/0.97BY, Fig. 1).
Vittatispora coorgii P. Chaudhary, J. Campb., D. Hawksw. & K.N. Sastry, Mycologia 98(3): 463
(2006) Fig. 15o–q
Facesoffungi number: FoF 10155
Isolated from soil. Sexual morph: Ascomata perithecial, solitary or gregarious, superficial to
semi-immersed, globose, with long necks, pale brown to brown, membranaceous, translucent,
glabrous or hairy, ostiolate. Necks cylindrical, composed of fasciculate, cylindrical, pale brown
hyphae, straight or slightly curved, ascospores usually accumulating in translucent necks. Peridium
membranaceous, composed of brown cells of textura angularis. Asci 8-spored, unitunicate, clavate,
evanescent. Ascospores hyaline becoming brown or dark brown, ellipsoidal to citriform, apiculate,
aseptate, smooth-walled, with germ pore surrounded by a raised rim and connected by a prominent
longitudinal ridge. Asexual morph: Undetermined (adapted from Chaudhary et al. 2016).
Known hosts and distribution – on soil in India (type locality) (Chaudhary et al. 2006)
Notes – Vittatispora coorgii is similar to Syspastospora tropicalis has translucent ascomata,
clavate asci and ascospores with germ pore surrounded by a raised rim and connected by a
prominent longitudinal ridge, but S. tropicalis has cylindrical to doliiform ascospores (García et al.
2002, Chaudhary et al. 2006). In this study, Syspastospora is basal to Gonatobotrys and
Vittatispora based on phylogenetic result (54%ML/0.99BY, Fig. 1).
Chaetosphaerellaceae Huhndorf, A.N. Mill. & F.A. Fernández, in Huhndorf, Miller & Fernández,
Mycol. Res. 108(12): 1387 (2004)
Saprobic on wood. Sexual morph: Subiculum absent or extensive, composed of densely
clustered, abundant, dark brown to black, septate, rigid, branched, spinulose hyphae with
dichotomously branched, tapering ends. Ascomata perithecial or without an external opening,
scattered or densely gregarious, superficial or sitting on or in a subiculum, turbinate, globose to
subglobose, carbonaceous or coriaceous, tuberculate, black, glabrous or with brown, cylindrical,
spinous setae or black tooth-like spines, ostiolate with papilla or lacking ostioles with Quellkörper,
collabent or not. Peridium carbonaceous to coriaceous to membranaceous, composed of brown to
hyaline cells of textura angularis to textura prismatica, sometimes Munk pore present. Paraphyses
numerous, filiform or broadly cylindrical, septate, inflated, sometimes present only in young
ascomata. Asci 8-spored, unitunicate, clavate to cylindrical, with long or short pedicel, apical ring
or inconspicuous, evanescent. Ascospores uni- or bi-seriate or overlapping, hyaline, becoming
brown or brown median cells and hyaline end cells, ellipsoidal to cylindrical, 1–3-septate, smooth-
walled, mostly with guttules. Asexual morph: Hyphomycetous. Conidiophores macronematous,
mononematous, brown to dark brown, pale brown towards the apex, occasionally pale towards the
base, branched, septate. Conidiogenous cells terminal and intercalary, polyblastic, holoblastic,
brown, swollen, globose to subglobose. Conidia solitary, multiple conidia are produced serially
from a retrogressive locus, usually central cell dark brown and end cell subhyaline to hyaline, 1–3-
septate, oval to ellipsoidal (adapted from Müller & Booth 1972, Sivanesan 1974, Réblová 1999a,
b). Type genus – Chaetosphaerella E. Müll. & C. Booth
Notes – Chaetosphaerellaceae was established by Huhndorf et al. (2004b) and initially
accommodated Chaetosphaerella and Crassochaeta based on phylogenetic results. Subsequently,
Mugambi & Huhndorf (2010) found that Spinulosphaeria exists independently in Coronophorales
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based on LSU sequence data analysis. These three genera were accepted in Chaetosphaerellaceae,
because they have similar characteristics of turbinate, tuberculate, black ascomata sitting on hyphal
subiculum, clavate asci and ellipsoidal ascospores, and they clustered based on multi-gene analyses
(Maharachchikumbura et al. 2015, 2016, Hyde et al. 2017). In this study, Chaetosphaerellaceae
clade, which accommodates Chaetosphaerella, Crassochaeta and Spinulosphaeria, nests in
Coronophorales (99%ML/1.00BY, Fig. 1).
Chaetosphaerella E. Müll. & C. Booth, Trans. Br. mycol. Soc. 58(1): 76 (1972)
Saprobic on wood. Sexual morph: Subiculum extensive, composed of densely clustered,
abundant, dark brown to black, septate, rigid, branched, spinulose hyphae with dichotomously
branched, tapering ends. Ascomata perithecial, scattered or densely gregarious, superficial or sitting
on or in a subiculum, turbinate, globose to subglobose, carbonaceous, tuberculate, black, glabrous
or with brown, cylindrical, spinous setae, ostiolate with papilla, periphysate. Peridium
carbonaceous to membranaceous, composed of dark brown to brown to hyaline cells of textura
angularis to textura prismatica, Munk pore present. Paraphyses broadly cylindrical, septate,
inflated, sometimes present only in young ascomata. Asci 8-spored, unitunicate, cylindrical, apical
ring indistinct, with short pedicel. Ascospores bi-seriate, hyaline, aseptate, becoming brown median
cells and hyaline end cells, 3-septate, slightly constricted at the middle septum, ellipsoidal to
cylindrical, slightly curved, smooth-walled, mostly with guttules. Asexual morph: Hyphomycetous.
Conidiophores macronematous, mononematous, brown to dark brown, pale brown towards the
apex, occasionally pale towards the base, branched, septate. Conidiogenous cells terminal and
intercalary, polyblastic, holoblastic, brown, swollen, globose to subglobose. Conidia solitary,
multiple conidia are produced serially from a retrogressive locus, usually central cell dark brown
and end cell subhyaline to hyaline, 1–3-septate, oval to ellipsoidal (adapted from Müller & Booth
1972, Réblová 1999a).
Notes – Chaetosphaerella was introduced as a genus similar to Chaetosphaeria in having
turbinate ascomata with hyphal subiculum and septate ascospores. However, the latter has
enteroblastic conidiogenesis, while the former is holoblastic (Müller & Booth 1972). Réblová
(1999a) clarified two Chaetosphaerella species, C. fusca and C. phaeostroma (type), based on their
characteristics. Subsequently, the molecular data showed that these two species clustered in
Chaetosphaerellaceae, which is distinct from Chaetosphaeria (Chaetosphaeriaceae) (Huhndorf et
al. 2004b, Mugambi & Huhndorf 2010). Its asexual morphs Oedemium and Veramycina were
synonymised as Chaetosphaerella (Réblová et al. 2016). We found that the sexual morphs of C.
fusca and C. phaeostroma are similar, but their conidiogenesis differs. Chaetosphaerella
phaeostroma has polyblastic conidiogenous cells (Fig. 10 t) whereas C. fusca has the multiple
conidia produced serially from a retrogressive locus of hyphae (Fig. 10 u–y). Phylogenetically,
these two species form Chaetosphaerella clade, sister to Crassochaeta (100%ML/1.00BY, Fig. 1)
in Chaetosphaerellaceae.
Chaetosphaerella phaeostroma (Durieu & Mont.) E. Müll. & C. Booth, Trans. Br. mycol. Soc.
58(1): 77 (1972) Fig. 10a–t
Basionym – Sphaeria phaeostroma Durieu & Mont., in Durieu, Expl. Sci. Alg., Fl. Algér.
1(livr. 13): 491 (1846)
Facesoffungi number: FoF 01116
Saprobic on wood. Sexual morph: Subiculum extensive, composed of densely clustered,
abundant, dark brown to black, septate, rigid, branched, spinulose hyphae (3–)5–7(–10) µm wide,
with dichotomously branched, tapering ends. Ascomata 400–460 × 200–335 µm (x
̄ = 435 × 275
µm, n = 10), perithecial, scattered or densely gregarious, superficial or sitting on or in subiculum,
turbinate, globose to subglobose, carbonaceous, tuberculate, black, glabrous or with brown,
cylindrical, spinous setae 7–12 µm, ostiolate with papilla, periphysate. Peridium 25–65 µm thick,
comprising three layers, outer layer composed of carbonaceous, dark tissues; middle layer
composed of membranaceous, dark brown to brown cells of textura angularis, Munk pores present;
906
inner layer composed of membranaceous, hyaline cells of textura prismatica. Paraphyses broadly
cylindrical, septate, inflated, present only in young ascomata. Asci (90–)115–135(–145) × (16–)17–
20(–21) µm (x
̄ = 121 × 19 µm, n = 30), 8-spored, unitunicate, cylindrical, apical ring indistinct,
with short pedicel. Ascospores (27–)29–32(–36) × (5–)7–8(–9.5) µm (x
̄ = 30 × 7.5 µm, n = 50), bi-
seriate, hyaline, aseptate, becoming brown median cells and hyaline end cells, 3-septate, slightly
constricted at the middle septum, ellipsoidal to cylindrical, slightly curved, smooth-walled, mostly
with guttules. Asexual morph: Hyphomycetous. Conidiophores macronematous, mononematous,
brown to dark brown, pale brown towards the apex, occasionally pale towards the base, branched,
septate. Conidiogenous cells terminal and intercalary, holoblastic, polyblastic, brown, globose to
subglobose, with swollen apex. Conidia solitary, brown to dark brown, pale towards the ends,
subglobose to ellipsoidal, aseptate when young, becoming 1–3-septate, constricted at the septum,
conidial secession with conspicuous denticles on the conidia and conidiogenous cells (adapted from
Réblová 1999a).
Figure 10 – Chaetosphaerella phaeostroma: a–b, e, g, m–n, p–s (PC-MNHN-PC-PC0167645); c, i
(S-F133811); d, o (PC-MNHN-PC-PC0167641); f, h, j (S-F133805); k (S-F133806); l, t (redrawn
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from Réblová 1999a). a Material. b–d Ascomata on the host. e–f Ascoma with setae. g–h Ascoma
cross section. i Ostiole with periphyses. j Branched hypha. k–l Setae. m–o Asci. p–r Ascospores. s
Munk pores (arrow). t Conidiophores with polyblastic conidiogenous cells; Chaetosphaerella
fusca: u–aa (UPS-UPS:BOT:F-671626; Russia, Saint Petersburg, Botanical Garden of the Komarov
Botanical Institute (lat. 59.96778 long. 30.33167), on dead branches of unknown deciduous tree
lying on the surface of ground, 27 August 2009, V. Mel’nik). u–y Conidiophores and conidia
produced serially from a retrogressive locus. z–aa Conidia. Scale bars: b–c, e–f = 500 µm, d, g =
200 µm, h, j = 100 µm, i, m–o, s–v = 50 µm, k–l, p–r, w–aa = 20 µm.
Material examined – France, Paris, on dead wood (PC-MNHN-PC-PC0167641); Algeria, on
dead wood (PC-MNHN-PC-PC0167645); on dead wood (PC-MNHN-PC-PC0167647); Sweden,
Uppland, Gamla Uppsala parish, Fullerö, in a grove by R. Fyris, on decaying fallen branches of
ash, associated with Eutypa flavovirens, 9 May 1982, N. Lundqvist (S-F133805); Sweden,
Uppland, Stockholm, in the experimental field, on dead wood, 3 May 1920, T. Vestergren (S-
F133806); Denmark, Sydjylland, Jutland, Haderslev prov., TBU 48, Pamhule forest near
Christiansdal (= 7 km SW of Haderslev), on Eutypa stroma on Fagus in beech forest, 22 September
1988, N. Lundqvist (S-F133811); Czech Republic, Praha, Cholupice, Bažantnice, on dead wood of
Acer negundo, 24 November 1978, F. Kotlaba (PRM-815939); Czech Republic, Mährisch-
Weißkirchen (Hranice), Militär-Oberrealschule, on dead wood, November 1913, J. Petrak (PRM-
875781).
Known hosts and distribution – on decayed wood of Pistacia lentiscus in Algeria (type
locality) (Bory de Saint-Vincent & Durieu de Maisonneuve 1849); on deaded wood of Acer,
Broussonetia, Carpinus, Corylus, Fagus, Fraxinus, Hedera, Ilex, Lonnicera, Picea, Pistacia,
Prunus, Quercus, Ribes, Salix, Tilia, Ulmus in Austria, China, Czech Republic, Denmark, England,
France, Germany, Guyana, Italy, Sweden and the USA (Fuckel 1870a, Saccardo 1883, Winter
1887a, Ellis & Everhart 1892, Strasser 1911, Teng 1934, Booth 1958, Eriksson 1992, Courtecuisse
et al. 1996, Farr et al. 1989, Réblová 1999a).
Notes – Chaetosphaerella phaeostroma was introduced with two types of asexual morphs,
Oedemium minus and Veramycina elegans (synanamorph) on natural substrate (Réblová 1999a).
Initially, V. elegans was described on cultural studies of O. minus (Subramanian 1993), and
Réblová (1999a) illustrated that O. minus has polyblastic conidiogenous cells and 1–3-septate,
ellipsoidal conidia (Fig. 10 t), and V. elegans has single or clustered phialides and hyaline, aseptate,
cylindrical conidia. However, more culture characteristics and sequence data are needed to
determine their affinities. Figure 1 reveals that C. phaeostroma is sister to C. fusca (59%/ML) in
Chaetosphaerellaceae.
Crassochaeta Réblová, Mycotaxon 71: 46 (1999)
Saprobic on decayed wood. Sexual morph: Subiculum lacking or extensive, composed of
densely clustered, dark brown to black, septate, rigid, branched, spinulose hyphae with tapering
ends. Ascomata perithecial, superficial, gregarious, dark brown to black, turbinate, conical to
ovoidal, tuberculate, collapsing or laterally collapsing when dry, surrounded with dark brown
spinous setae, ostiolate with papilla, periphysate. Peridium membranaceous, composed of brown to
hyaline cells of textura angularis to textura prismatica, Munk pore absent or present. Paraphyses
cylindrical, branched, septate, sometimes constricted at the septa, evanescent. Asci 8-spored,
unitunicate, cylindrical, with short pedicel, evanescent. Ascospores fusiform to ellipsoidal, smooth-
walled, slightly curved, aseptate, hyaline when young, becoming 3-septate, slightly constricted at
the septa, two middle cells brown to dark brown and two end cells hyaline, mostly with guttules.
Asexual morph (associated): Conidia solitary, scattered among the hyphae of the subiculum or
attached to the surface of the ascomata, ellipsoidal to citriform, slightly apiculate, brown to black,
aseptate, with a longitudinal germ slit that nearly connects the two ends (adapted from Réblová
1999b).
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Notes – Réblová (1999b) established Crassochaeta in Trichosphaeriaceae and segregated two
Chaetosphaeria species as its members, Crassochaeta fusispora and Cra. nigrita (type), based on
their ascomata being larger than Chaetosphaeria. Subsequently, Huhndorf et al. (2004b) analyzed
LSU sequence data of Cra. nigrita and found that Crassochaeta is sister to Chaetosphaerella in
Coronophorales. Morphologically, Chaetosphaerella and Crassochaeta have similar setose
ascomata, cylindrical asci and versicoloured ascospores, but are associated with different asexual
morphs. Chaetosphaerella has septate, ellipsoidal conidia, while Crassochaeta has citriform,
aseptate conidia with a longitudinal germ slit (Réblová 1999a, b). Thus, Crassochaeta is
accommodated in Chaetosphaerellaceae based on morphology and phylogenetic results
(Maharachchikumbura et al. 2015, 2016, Hyde et al. 2020e, Wijayawardene et al. 2020). In this
study, Crassochaeta is sister to Chaetosphaerella (100%ML/1.00BY, Fig. 1).
Crassochaeta nigrita (Berk. & Broome) Réblová, Mycotaxon 71: 48 (1999) Fig. 11
Basionym – Sphaeria nigrita Berk. & Broome, J. Linn. Soc., Bot. 14(no. 74): 125 (1873)
Facesoffungi number: FoF 10059
Saprobic on decayed wood. Sexual morph: Subiculum extensive, composed of densely
clustered, dark brown to black, septate, rigid, branched, spinulose hyphae 3–7 µm wide, with
dichotomously branched or three to multi-branched, tapering ends. Ascomata 330–500 × 265–365
µm (x
̄ = 400 × 300 µm, n = 10), perithecial, superficial, gregarious, dark brown to black, turbinate,
conical to ovoidal, tuberculate, collapsing or laterally collapsing when dry, surrounded with dark
brown spinous setae 5–11 µm wide at base, ostiolate with papilla, periphysate. Peridium
membranaceous, comprising two layers, outer layer composed of dark brown to brown cells of
textura angularis, Munk pore present; inner layer composed of hyaline cells of textura prismatica.
Paraphyses cylindrical, branched, septate, evanescent. Asci (110–)120–145(–160) × (8–)10–13(–
15) µm (x
̄ = 140 × 11.5 µm, n = 30), 8-spored, unitunicate, cylindrical, apical ring distinct in empty
asci, with short pedicel, evanescent. Ascospores 18–23 × 7–10 µm (x
̄ = 20 × 8 µm, n = 50),
ellipsoidal, smooth-walled, slightly curved, aseptate, hyaline when young, becoming 3-septate,
slightly constricted at the septa, two middle cells brown to dark brown, and two end cells hyaline,
mostly with guttules. Asexual morph (associated): Conidia solitary, scattered among the hyphae of
the subiculum or attached to the surface of the ascomata, ellipsoidal to citriform, slightly apiculate,
brown to black, aseptate, with a longitudinal germ slit nearly connects the two ends (adapted from
Réblová 1999b).
Material examined – Sri Lanka, Hakgala, Nuwara Eliya Distr., on dead wood, December
1864, G.H.K. Thwaites (IMI-K(M):37405, isotype); China, Hainan, Tan-hsien, on dead wood, 5
December 1934, S.Q. Deng (CUP-CUP-CH 661, paratype).
Known hosts and distribution – on decayed wood in Brazil, Costa Rica, China, Panama, Sri
Lanka (type locality) (Réblová 1999b, Huhndorf et al. 2004b).
Notes – Crassochaeta nigrita was initially introduced as Sphaeria nigrita Berk. & Broome
from Sri Lanka (Berkeley & Broome 1873). Réblová (1999b) proposed that Chaetosphaeria
hainanensis is a synonym of Cra. nigrita based on similar characteristics. Subsequently, Huhndorf
et al. (2004b) analyzed LSU sequence data of Cra. nigrita and placed Crassochaeta in
Chaetosphaerellaceae. We re-examined two well preserved specimens, one is isotype of S. nigrita
from Sri Lanka identified by Berkeley, and the other is Cha. hainanensis from China was
determined as paratype of C. nigrita by Huhndorf (mentioned in the label of material). The strains
of Cra. nigrita are sister to Chaetosphaerella (100%ML/1.00BY, Fig. 1).
Neochaetosphaerella Lar.N. Vassiljeva, S.L. Stephenson & Chernyshev, in Vasilyeva, Chernyshev
& Stephenson, Fungal Diversity 52(1): 192 (2012)
Saprobic on wood. Sexual morph: Subiculum composed of brown, septate, branched, terminal
acuminate hyphae. Ascomata perithecial, gregarious, superficial, sitting in a subiculum, oval to
subglobose, black, tuberculate, with spiny, septate setae, ostiolate. Peridium composed of brown
cells of textura angularis. Paraphyses filiform, evanescent. Asci 8-spored, unitunicate, clavate,
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apex rounded, apical ring inconspicuous, with a short pedicel or sessile, evanescent. Ascospores bi-
seriate to overlapping, broadly allantoid, brown, 3-septate, smooth-walled, guttulate. Asexual
morph: Undetermined (adapted from Vasilyeva et al. 2012).
Figure 11 – Crassochaeta nigrita: a–c, h, k–m (IMI-K(M):37405, isotype); d–g, i–j, n–o (CUP-
CUP-CH 661, paratype). a Material. b Gregarious ascomata. c–d Ascomata on the host. e Peridium
with Munk pores. f Periphyses. g Setae. h Hyphae. i–k Asci (j apical ring in empty ascus). l–o
Ascospores. Scale bars: b = 1 mm, c = 500 µm, d = 200 µm, i = 100 µm, f–h, k = 50 µm, e = 20
µm, l–o = 10 µm.
Notes – The monotypic Neochaetosphaerella is similar to Chaetosphaerella and
Crassochaeta in having tuberculate, setose ascomata and its subiculum is similar to Scortechinia
(Scortechiniaceae) with acuminate hyphae. However, Neochaetosphaerella has septate, brown
ascospores whereas Chaetosphaerella and Crassochaeta have versicoloured ascospores and
Scortechinia has aseptate hyaline ascospores (Vasilyeva et al. 2012). Neochaetosphaerella was
initially placed in Coronophorales (Vasilyeva et al. 2012), and then Nitschkiaceae
(Maharachchikumbura et al. 2015, Wijayawardene et al. 2020). However, Neochaetosphaerella
does not have the hyaline ascospores of Nitschkiaceae. We transfer Neochaetosphaerella to
Chaetosphaerellaceae based on the similar tuberculate ascomata, clavate asci and 3-septate
ascospores pending molecular data.
910
Neochaetosphaerella thaxteriospora Lar.N. Vassiljeva, S.L. Stephenson & Chernyshev, in
Vasilyeva, Chernyshev & Stephenson, Fungal Diversity 52(1): 192 (2012) Fig. 15t
Facesoffungi number: FoF 10156
Saprobic on wood. Sexual morph: Subiculum composed of brown, septate, branched, terminal
acuminate hyphae. Ascomata perithecial, gregarious, superficial, sitting in a subiculum, oval to
subglobose, black, tuberculate, with spiny, septate setae, ostiolate. Peridium composed of brown
cells of textura angularis. Paraphyses filiform, evanescent. Asci 8-spored, unitunicate, clavate,
apex rounded, apical ring inconspicuous, with a short pedicel or sessile, evanescent. Ascospores bi-
seriate to overlapping, broadly allantoid, brown, 3-septate, smooth-walled, guttulate. Asexual
morph: Undetermined (adapted from Vasilyeva et al. 2012).
Known hosts and distribution – on decayed wood in Russia (type locality) (Vasilyeva et al.
2012).
Notes – Neochaetosphaerella thaxteriospora was initially collected from Russia and is
similar to Chaetosphaerella phaeostroma in having setose ascomata and septate, brown ascospores
(Vasilyeva et al. 2012).
Spinulosphaeria Sivan., Trans. Br. mycol. Soc. 62(1): 5 (1974)
Saprobic on decayed wood. Sexual morph: Subiculum lacking or extensive, composed of
densely clustered, brown to dark brown, septate, branched hyphae. Ascomata superficial, usually
sitting on a subiculum, scattered to gregarious, black, with metallic iridescence, turbinate, conical
to ovoidal, tuberculate, collapsing when dry, glabrous or surrounded with black, short, tooth-like,
usually furcate, spinose ornaments, lacking ostioles, with Quellkörper or not. Peridium
carbonaceous or membranaceous, composed of brown to hyaline cells of textura angularis to
textura prismatica, Munk pore absent or present. Paraphyses septate, longer than asci, filiform or
broad and constricted at the septa, evanescent. Asci 8-spored, unitunicate, clavate, apical ring
distinct or indistinct, long pedicellate, evanescent. Ascospores oblong to ellipsoidal, smooth-walled
or verrucose, hyaline when young, becoming brown, 1-septate, with large mucilaginous sheath or
not. Asexual morph: Undetermined (adapted from Sivanesan 1974, Mugambi & Huhndorf 2010).
Notes – Spinulosphaeria was placed in Coronophorales and typified by S. thaxteri in having
turbinate ascomata covered by tooth-like spines, Quellkörper present and ellipsoidal ascospores
without sheath (Sivanesan 1974). Subsequently, Mugambi & Huhndorf (2010) introduced a second
species, S. nuda, and analyzed its LSU sequence data. However, S. nuda is characterized by
glabrous ascomata without Quellkörper and ellipsoidal ascospores with large glutinous sheath,
which are different from S. thaxteri. Spinulosphaeria is accepted as a member of
Chaetosphaerellaceae based on molecular information of S. nuda (Maharachchikumbura et al.
2015, 2016, Hyde et al. 2017). In this study, Spinulosphaeria is basal to Chaetosphaerella and
Crassochaeta in Chaetosphaerellaceae (0.92BY, Fig. 1).
Type species – Spinulosphaeria thaxteri (Pat.) Sivan., Trans. Br. mycol. Soc. 62(1): 36 (1974)Fig 12
Basionym – Neopeckia thaxteri Pat., Bull. Soc. mycol. Fr. 38(2): 83 (1922)
Facesoffungi number: FoF 10060
Saprobic on decayed wood. Sexual morph: Subiculum extensive, composed of densely
clustered, brown to dark brown, septate, branched hyphae. Ascomata 860–910 × 230–440 µm (x
̄ =
885 × 280 µm, n = 5), superficial, scattered to gregarious, black, with metallic iridescence,
carbonaceous, turbinate, conical to ovoidal, tuberculate, collapsing when dry, surrounded with
black, short, tooth-like, 1–4 furcate spines, lacking ostioles, with thin Quellkörper composed of
elongated cells at the inner wall of apical region. Peridium 30–70 µm wide, outer layer composed
of carbonaceous, dark tissues; middle layer composed of membranaceous, brown to hyaline cells of
textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses septate,
longer than asci, filiform, evanescent. Asci (30–)45–50(–68) × (8–)10–13(–15) µm (x
̄ = 48 × 12
µm, n = 20) at ascospore-bearing part, 8-spored, unitunicate, clavate, apical ring, long pedicellate,
evanescent. Ascospores 9.5–13 × 3.5–5.5 µm (x
̄ = 11 × 4.5 µm, n = 30), oblong to ellipsoidal,
911
smooth-walled, hyaline, aseptate when young, becoming brown to olivaceous, 1-septate, guttulate.
Asexual morph: Undetermined (adapted from Sivanesan 1974).
Material examined – USA, Florida, Dayton, on decayed wood, January 1898, R. Thaxter
(FH-16-10965, holotype).
Known hosts and distribution – on decayed wood in Liberia, Sierra Leone, the USA (type
locality).
Notes – Spinulosphaeria thaxteri was introduced based on Neopeckia thaxteri from the USA
and is characterized by subiculum composed of branched hyphae, ascomata surrounded by special
tooth-like spines and with thin Quellkörper. There is no molecular data available for this species. In
this study, we re-examined the holotype and found that its ascomata are brittle; paraphyses and asci
are evanescent.
Figure 12 – Spinulosphaeria thaxteri: a–f, h–n (FH-16-10965, holotype); g (redrawn from
Sivanesan 1974). a Material label. b–c Turbinate ascomata on wood. d Tuberculate ascoma (arrow
912
indicates the enlarged tooth-like spine). e Ascoma cross section. f Peridium. g Sectional view of
ascoma with spines. h–j Asci. k–n Ascospores. Scale bars: b–c = 500 µm, d–e, g = 200 µm, f = 100
µm, h–j = 20 µm, k = 10 µm, l–n = 5 µm.
Coronophoraceae Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 116: 624 (1907)
Saprobic on wood, sometimes fungicolous. Sexual morph: Ascomata perithecial or without
an external opening, solitary or gregarious, semi-immersed or erumpent or superficial, black,
usually carbonaceous, globose to subglobose, sometimes with necks, tuberculate, collapsing or
laterally collapsing when dry, glabrous or with hairs, with or lacking ostioles, usually with a central
column of mucilaginous cells (Quellkörper). Peridium outer layer composed of carbonaceous,
black tissue; inner layer composed of membranaceous, brown to hyaline cells of textura angularis
to textura prismatica. Paraphyses absent or numerous, filiform, septate, evanescent. Asci numerous
or rare, polysporous, unitunicate, clavate to cylindrical, long pedicellate, apex blunt or rounded,
apical ring inconspicuous, evanescent. Ascospores crowded, numerous, hyaline, allantoid to
cylindrical, sometimes curved, 0–1-septate, smooth-walled, mostly with guttules. Asexual morph:
Undetermined (adapted from Fuckel 1864, Saccardo 1882a).
Type genus – Coronophora Fuckel
Notes – Coronophoraceae was introduced by von Höhnel (1907) to accommodate
Cryptosphaerella and Coronophora, and it is characterized by polysporous asci and hyaline,
allantoid ascospores. Subsequently, Cryptosphaerella was clarified as a genus in Scortechiniaceae
based on phylogenetic result (Mugambi & Huhndorf 2010). Coronophora was placed as an
independent clade in Coronophorales based on the sequences data of Cor. gregaria (Mugambi &
Huhndorf 2010). Therefore, Coronophora is the only member in Coronophoraceae, and this genus
is similar to Fracchiaea (Nitschkiaceae) and Pleurostoma (Pleurostomataceae, Calosphaeriales) in
having black ascomata and clavate, polysporous asci (Petch 1916, Barr 1985, Mugambi &
Huhndorf 2010, Maharachchikumbura et al. 2015, 2016, Hyde et al. 2020e).
Coronophora Fuckel, Fungi rhenani exsic., fasc. 10: no. 961 (1864)
Saprobic on wood, sometimes fungicolous. Sexual morph: Ascomata perithecial or without
an external opening, solitary or gregarious, semi-immersed or erumpent or superficial, black,
usually carbonaceous, globose to subglobose, sometimes with necks, tuberculate, collapsing or
laterally collapsing when dry, glabrous or with hairs, with or lacking ostioles, usually with
Quellkörper. Peridium usually outer layer composed of carbonaceous, black tissue; inner layer
composed of membranaceous, brown to hyaline cells of textura angularis to textura prismatica.
Paraphyses absent or numerous, filiform, septate, evanescent. Asci numerous or rare, polysporous,
unitunicate, clavate to cylindrical, long pedicellate, apex blunt or rounded, apical ring
inconspicuous, evanescent. Ascospores crowded, numerous, hyaline, allantoid to cylindrical,
sometimes curved, 0–1-septate, smooth-walled, mostly with guttules. Asexual morph:
Undetermined (adapted from Fuckel 1864, Saccardo 1882a).
Notes – Coronophora is typified by C. gregaria which has polysporous asci and hyaline,
cylindrical, aseptate ascospores (Fuckel 1864, Saccardo 1882b). There are 27 species in this genus,
two of which have available molecular data, C. gregaria and C. myricoides (Mugambi & Huhndorf
2010, Hyde et al. 2016, Index Fungorum 2021). The characteristics of these two species are similar
except for the ascospore size (Hyde et al. 2016). In this study, C. gregaria is sister to
Scortechiniaceae (100%ML/1.00BY), whereas C. myricoides is sister to Fracchiaea
(100%ML/1.00BY) in Nitschkiaceae (Fig. 1). Thus, we transferred C. myricoides as Fracchiaea
species (see notes for Fracchiaea myricoides).
Coronophora gregaria Fuckel, Jb. nassau. Ver. Naturk. 23-24: 229 (1870) [1869-70]
Fig. 13
Basionym – Sphaeria gregaria Lib., Pl. crypt. Arduenna, fasc. (Liège) 2(nos 101-200): no.
145 (1832)
913
Facesoffungi number: FoF 01119
Saprobic on wood. Sexual morph: Ascomata 0.9–1.4 × 0.3–1.2 mm (x
̄ = 1.2 × 0.6 mm, n =
10), gregarious in oval groups, in valsoid configuration with necks converging, immersed to
erumpent through the bark of host, black, carbonaceous, oblong to subglobose, tuberculate,
laterally collapsing when dry, glabrous, lacking ostioles, with a central column of hyaline,
mucilaginous cells (Quellkörper). Peridium 70–170 µm wide, comprising three layers, outer layer
composed of carbonaceous, black tissue, 14–95 µm wide; middle layer composed of
membranaceous, dark brown to brown cells of textura angularis, 30–85 µm wide, Munk pores
rarely present; inner layer composed of membranaceous, hyaline cells of textura prismatica, 12–40
µm wide. Paraphyses absent. Asci (57–)75–90(–120) × (10–)12–15(–20) µm (x
̄ = 85 × 13.5 µm, n
= 30), numerous, polysporous, unitunicate, clavate, long pedicellate, apex rounded, apical ring
inconspicuous. Ascospores (5.5–)7–8(–9.5) × 1.2–2.5 µm (x
̄ = 7.5 × 1.8 µm, n = 50), crowded,
numerous, hyaline, allantoid to cylindrical, slightly curved, aseptate, smooth-walled, with rare
guttules. Asexual morph: Undetermined (adapted from Saccardo 1882a).
Figure 13 – Coronophora gregaria: (BR-BR5020094489111, isotype). a Material. b Ascomata
erumpent through the bark of host. c–d Ascomata on decorticated wood. e Ascoma cross section.
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f Quellkörper. g Peridium. h Munk pore. i–k Asci. l–q Ascospores. Scale bars: b–d = 2 mm,
e–f = 200 µm, g, i = 100 µm, h, j–k = 20 µm, l–q = 5 µm.
Material examined – Belgium, on decayed wood of Prunus cerasus or Sorbus aucuparia,
Autumn, M.A. Libert (BR-BR5020094489111, isotype).
Known hosts and distribution – on decayed branch of Betula, Prunus, Sorbus and Tilia in
Belgium (type locality), France, Germany and Italy (Saccardo 1882b).
Notes – Coronophora gregaria was initially introduced as Sphaeria gregaria Lib. from
Belgium. Mugambi & Huhndorf (2010) placed this species in Coronophoraceae based on TEF and
RPB2 sequences data. Morphologically, Coronophora was reported to be similar to Fracchiaea and
Pleurostoma (Petch 1916, Barr 1985). We re-examined the isotype and found that it has gregarious
ascomata in oval groups as valsoid configuration with necks converging, Quellkörper present and
polysporous asci with inconspicuous discharge mechanism (ascospores accumulate on the ascus
apex, causing the ascus to expand to rupture and release ascospores, Fig. 13i). These characteristics
are different from Fracchiaea and Pleurostoma. The strain of C. gregaria (ANM1555) is basal to
Scortechiniaceae in Coronophorales (100%ML/1.00BY, Fig. 1).
Nitschkiaceae Nannf., Nova Acta R. Soc. Scient. upsal., Ser. 4 8(no. 2): 56 (1932)
Saprobic or parasitic on wood, leaves or lichen. Sexual morph: Subiculum absent or
composed of numerous or rare, brown, branched, septate hyphae. Ascomata perithecial or without
an external opening, solitary or gregarious, semi-immersed or erumpent or superficial, sitting on or
in a subiculum, usually carbonaceous, brown to black, cupuliform, globose to subglobose, glabrous
or with spines, the apex collapsing when dry or shallowly cupulate when moist, with periphyses
inside of the ostioles or inconspicuous ostioles or lacking ostioles, with or Quellkörper absent.
Peridium usually outer layer composed of carbonaceous, black tissue; inner layer composed of
membranaceous, brown to hyaline cells of textura angularis to textura prismatica, sometimes
Munk pores present. Paraphyses absent or numerous, filiform, branched or unbranched, septate,
evanescent. Asci 4-, 8- or multi-spored, unitunicate, clavate to cylindrical, pedicellate or sessile,
apex blunt or rounded, apical ring inconspicuous, evanescent. Ascospores hyaline to brown,
allantoid, ellipsoidal to cylindrical, sometimes curved, smooth-walled, sometimes with
mucilaginous sheath or appendages, mostly with guttules. Asexual morph: Undetermined (adapted
from Subramanian & Sekar 1990, Hyde et al. 2020e).
Type genus – Nitschkia G.H. Otth ex P. Karst.
Notes – Nitschkiaceae was initially introduced as subfamily ‘Nitschkieae’, which has affinity
with morphology of Coronophoraceae (Fitzpatrick 1923). Nannfeldt (1932) clarified their
relationship and established Nitschkiaceae in Coronophorales. Subsequently, this family
accommodated 14 genera and, is characterized by black, turbinate, cupuliform ascomata, usually
with Quellkörper and clavate to subcylindrical asci (Nannfeldt 1932, 1975, Subramanian & Sekar
1990, Maharachchikumbura et al. 2016, Hyde et al. 2020e). Four genera, Acanthonitschkea,
Fracchiaea, Neotrotteria and Nitschkia, have sequences data (Huhndorf et al. 2004b, Mugambi &
Huhndorf 2010). We found that Acanthonitschkea, Fracchiaea and Nitschkia (except N.
pezizoidea) are well clustered in Nitschkiaceae (96%ML/1.00BY) and closely related to Bertiaceae,
but Neotrotteria nests in Ceratostomataceae (68%ML, Fig. 1). We place Neotrotteria in
Ceratostomataceae based on our phylogenetic result (see notes for Neotrotteria).
Acanthonitschkea Speg., Anal. Mus. nac. B. Aires, Ser. 3 17(10): 116 (1908)
Saprobic on wood or lichen. Sexual morph: Subiculum composed of numerous, brown to
black, branched, septate, sinuous hyphae with metallic iridescence; or rigid, straight or slightly
sinuous, unbranched, aseptate, brown to black spines with metallic iridescence form a dense
barricade. Ascomata perithecial or without an external opening, scattered or gregarious, semi-
immersed or erumpent or superficial, sitting on or in a subiculum, carbonaceous, coriaceous or
membranaceous, brown to black, turbinate, cupuliform to subglobose, tuberculate, with spines
915
similar to those on the mycelium, the apex collapsing, with periphyses inside of the inconspicuous
ostioles or lacking ostioles, with or Quellkörper absent. Peridium outer layer composed of
carbonaceous, black tissue; inner layer composed of membranaceous, brown to hyaline cells of
textura angularis to textura prismatica, sometimes Munk pores present. Paraphyses absent. Asci 8-
spored, unitunicate, clavate, apex blunt or rounded, apical ring inconspicuous, with long pedicel,
evanescent. Ascospores bi-seriate to overlapping, hyaline, 0–1-septate, ellipsoidal to cylindrical,
sometimes curved, smooth-walled, mostly with guttules. Asexual morph: Undetermined (adapted
from Fitzpatrick 1923).
Notes – Acanthonitschkea was introduced as a genus with characteristics similar to Nitschkia,
but with spiny ascomata (Spegazzini 1908, Fitzpatrick 1923). This genus includes 10 species and is
typified by A. argentinensis in having spiny, tuberculate ascomata, clavate asci and allantoid
ascospores (Fitzpatrick 1923, Index Fungorum 2021). Two species have available molecular data,
A. argentinensis and A. tristis (Mugambi & Huhndorf 2010). In this study, these two
Acanthonitschkea species are sister to Nitschkia calyculus, N. grevillii and N. tetraspora
(73%ML/0.95BY, Fig. 1).
Acanthonitschkea argentinensis Speg., Anal. Mus. nac. B. Aires, Ser. 3 17(10): 116 (1908)
Fig. 14p
Facesoffungi number: FoF 10157
Saprobic on wood. Sexual morph: Subiculum composed of sparse to numerous, brown to
black, branched, septate, sinuous hyphae with metallic iridescence; or erect, rigid, straight or
slightly sinuous, unbranched, aseptate, brown to black spines with metallic iridescence form a
dense barricade. Ascomata scattered or densely gregarious, superficial, sitting on a subiculum,
membranaceous to coriaceous, brown to black, turbinate, cupuliform, tuberculate, with spines
similar to those on the mycelium, the apex collapsing to deeply cupulate, lacking ostioles. Peridium
composed of brown to hyaline cells of textura angularis to textura prismatica, Munk pores present.
Paraphyses absent. Asci 8-spored, unitunicate, clavate, apex blunt or rounded, apical ring
inconspicuous, with long pedicel, evanescent. Ascospores bi-seriate to overlapping, hyaline, 1-
septate, allantoid to horseshoe-shaped, smooth-walled, with two guttules in each cell. Asexual
morph: Undetermined (adapted from Fitzpatrick 1923, Subramanian & Sekar 1990).
Known hosts and distribution – on decayed wood in Argentina (type locality), India and
Puerto Rico (Spegazzini 1908, Subramanian & Sekar 1990, Mugambi & Huhndorf 2010).
Notes – Acanthonitschkea argentinensis was collected from a decayed wood in Argentina in
February (Spegazzini 1908). This species was considered to have affinity with Nitschkia based on
similar morphology (Fitzpatrick 1923, Subramanian & Sekar 1990). A. argentinensis (SMH1395)
was sequenced and shown to be related to Nitschkia species based on LSU, TEF and RPB2
sequence analyses (Mugambi & Huhndorf 2010). In this study, A. argentinensis is sister to A. tristis
(99%ML/1.00BY, Fig. 1) in Nitschkiaceae.
Acanthonitschkea tristis (J. Kickx f.) Nannf., Svensk bot. Tidskr. 69(1): 57 (1975) Fig. 14a–o
Basionym – Hyphasma triste J. Kickx f., FI. crypt. Louvain (Bruxelles): 118 (1835)
Facesoffungi number: FoF 10061
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to black,
branched, septate, sinuous hyphae, 4–7 µm wide, with metallic iridescence. Ascomata 380–600 ×
375–460 µm (x
̄ = 478 × 420 µm, n = 10), scattered or densely gregarious, superficial on bark or
decorticated wood, sometimes erumpent through the bark of substrate, sitting in a subiculum,
carbonaceous, black, turbinate, cupuliform, tuberculate, lacking ostioles, with Quellkörper.
Peridium 26–80 µm wide, comprising three layers, outer layer composed of carbonaceous, black
tissue; middle layer composed of membranaceous, dark brown to brown cells of textura angularis,
Munk pores present; inner layer composed of membranaceous, hyaline cells of textura prismatica.
Paraphyses absent. Asci 8-spored, unitunicate, clavate, with a blunt, thick apex and long pedicel,
evanescent. Ascospores 5.5–8(–10) × 1.5–2.5 µm (x
̄ = 6.5 × 2 µm, n = 50), bi-seriate to
916
overlapping, hyaline, ellipsoidal to fusiform, straight, aseptate when young, becoming 1-septate,
smooth-walled, with guttules. Asexual morph: Undetermined (adapted from Nannfeldt 1975,
Subramanian & Sekar 1990).
Figure 14 – Acanthonitschkea tristis: a–o (PRM-804846). a Material label. b Gregarious ascomata
on decorticated wood. c–d Ascomata. e Ascoma cross section. f Peridium. g Munk pores (arrow).
h Septate hyphae. i Asci with thick apex. j–o Ascospores; Acanthonitschkea argentinensis:
917
p (redrawn from Subramanian & Sekar 1990). p Asci and ascospores. Scale bars: c = 2 mm, d =
500 µm, e = 200 µm, f = 50 µm, h = 20 µm, g, i, p = 10 µm, j–o = 5 µm.
Material examined – USA, Florida, Rock Spa, on decayed, decorticated wood, 20 March
1942, C.L. Shear (PRM-804846).
Known hosts and distribution – on decayed wood in the USA (type locality) and Ecuador
(Fitzpatrick 1923, Mugambi & Huhndorf 2010).
Notes – Nannfeldt (1975) clarified materials related to ‘Sphaeria tristis’ and introduced
Acanthonitschkea tristis. Afterwards, A. tristis (SMH4723) from Ecuador was sequenced and it
nested in Nitschkiaceae (Mugambi & Huhndorf 2010). In this study, A. tristis is sister to A.
argentinensis (99%ML/1.00BY, Fig. 1) in Nitschkiaceae based on multi-gene analysis.
Biciliosporina Subram. & Sekar, Kavaka 18(1-2): 69 (1993) [1990]
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate, coriaceous, globose to subglobose, black, with metallic iridescence,
the apex collapsing when dry, lacking ostioles, with Quellkörper. Peridium composed of brown to
hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses absent.
Asci 8-spored, arising from a hyaline placenta-like tissue, radially extending into the locule around
the Quellkörper from the base, unitunicate, clavate to cylindrical, apex rounded, apical ring
inconspicuous, with long pedicel, evanescent. Ascospores uni- or bi-seriate, hyaline, aseptate,
ellipsoidal to reniform, smooth-walled, with a filiform, flagella-like appendage at each end,
guttulate. Asexual morph: Undetermined (adapted from Subramanian & Sekar 1990).
Notes – The monotypic Biciliosporina was introduced as a genus in Coronophoraceae and
typified by B. karwarensis in having subglobose ascomata, clavate to cylindrical asci and reniform
ascospores with a flagella-like appendage at each end (Subramanian & Sekar 1990). Later, this
genus was clarified as a member of Nitschkiaceae based on morphology (Eriksson & Hawksworth
1987, Lumbsch & Huhndorf 2010, Maharachchikumbura et al. 2016, Hyde et al. 2020e, Index
Fungorum 2021). We found that the ascospores of Biciliosporina are similar to Janannfeldtia,
although the latter has polysporous asci.
Biciliosporina karwarensis Subram. & Sekar, Kavaka 18(1-2): 69 (1993) [1990] Fig. 15a–e
Facesoffungi number: FoF 10158
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate, coriaceous, globose to subglobose, black, with metallic iridescence,
the apex collapsing when dry, lacking ostioles, with Quellkörper. Peridium composed of brown to
hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses absent.
Asci 8-spored, arising from a hyaline placenta-like tissue, radially extending into the locule around
the Quellkörper from the base, unitunicate, clavate to cylindrical, apex rounded, apical ring
inconspicuous, with long pedicel, evanescent. Ascospores uni- or bi-seriate, hyaline, aseptate,
ellipsoidal to reniform, smooth-walled, with a filiform, flagella-like appendage at each end,
guttulate. Asexual morph: Undetermined (adapted from Subramanian & Sekar 1990).
Material examined – India, Karnataka, Karwar, upper Anmod, on dead twig, 22 November
1980, G. Sekar (IMI-324681, holotype).
Known hosts and distribution – on decayed wood in India (type locality) (Subramanian &
Sekar 1990).
Notes – Biciliosporina karwarensis was introduced based on the specimen IMI324681
(Subramanian & Sekar 1990). We re-examined this sample and found that most of the ascomata
were shrivelled and collapsed, and no other structures could be found.
Botryola Bat. & J.L. Bezerra, in Batista et al., Publicações Inst. Micol. Recife 431: 11 (1964)
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate, carbonaceous, cupuliform, globose to subglobose, black, with spinous
918
setae, lacking ostioles, with Quellkörper. Peridium composed of brown to hyaline cells of textura
angularis to textura prismatica. Paraphyses absent. Asci 4-spored, unitunicate, clavate to
cylindrical, apex rounded, with long pedicel, evanescent. Ascospores uni- or bi-seriate, hyaline, 1-
septate, oblong to cylindrical, smooth-walled, guttulate. Asexual morph: Undetermined (adapted
from Batista et al. 1964).
Figure 15 – Biciliosporina karwarensis: a–b (IMI-IMI324681, holotype); c–e (redrawn from
Subramanian & Sekar 1990). a Material. b Collapsed ascomata on the host. c Sectional view of
ascoma. d Asci. e Ascospores; Syspastospora parasitica: f (redrawn from Cannon & Hawksworth
1982). f Ascospores; Scopinella barbata: g–h (redrawn from Hawksworth 1975). g Ascus.
h Ascospores; Loranitschkia viticola: i (redrawn from Vasilyeva et al. 2010). i Asci and ascospores;
Botryola tetrasperma: j (redrawn from Batista et al. 1964). j Asci and ascospores; Setiferotheca
nipponica: k–m (redrawn from Matsushima 1995). k Peridium with setae. l Asci. m Ascospores;
Janannfeldtia karwarensis: n (redrawn from Subramanian & Sekar 1990). n Asci and ascospores;
Vittatispora coorgii: o–q (redrawn from Chaudhary et al. 2006). o Ascoma. p Ascus. q Ascospores;
Rhagadostomella gregaria: r–s (redrawn from Etayo 2002). r Ascospores. s Asci;
Neochaetosphaerella thaxteriospora: t (redrawn from Vasilyeva et al. 2012). t Ascus and
ascospores; Tortulomyces thailandicus: u (redrawn from Vasilyeva et al. 2013). u Ascus and
919
ascospores. Scale bars: b = 500 µm, c, k, o = 200 µm, i = 50 µm, d, j, n, s–u = 20 µm, e, g, l, r = 10
µm, h, m, p–q = 5 µm, f = 2 µm.
Notes – The monotypic Botryola was initially placed in Coronophoraceae, but was
subsequently clarified as a genus in Nitschkiaceae based on its ascomata similar to Fracchiaea
(Batista et al. 1964, Eriksson & Hawksworth 1987, Lumbsch & Huhndorf 2010,
Maharachchikumbura et al. 2016, Hyde et al. 2020e).
Botryola tetrasperma Bat. & J.L. Bezerra, in Batista, Bezerra, Castrillón & Matta, Publicações Inst.
Micol. Recife 431: 13 (1964) Fig. 15j
Facesoffungi number: FoF 10159
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate, carbonaceous, cupuliform, globose to subglobose, black, with spinous
setae, lacking ostioles, with Quellkörper. Peridium composed of brown to hyaline cells of textura
angularis to textura prismatica. Paraphyses absent. Asci 4-spored, unitunicate, clavate to
cylindrical, apex rounded, with long pedicel, evanescent. Ascospores uni- to bi-seriate, hyaline, 1-
septate, oblong to cylindrical, smooth-walled, guttulate. Asexual morph: Undetermined (adapted
from Batista et al. 1964).
Known hosts and distribution – on decayed wood in Brazil (type locality) (Batista et al.
1964).
Notes – Botryola tetrasperma was collected from Manaus, Brazil on 20 February 1961
(Batista et al. 1964). We found that it is similar to Setiferotheca which has spiny ascomata and 4-
spored asci, although the latter has perithecial ascomata without Quellkörper.
Fracchiaea Sacc., Atti Soc. Veneto-Trent. Sci. Nat. 2(1): 163 (1873)
Saprobic on wood. Sexual morph: Ascomata scattered to gregarious, immersed to erumpent
through the bark of substrate or superficial, carbonaceous to coriaceous, black, turbinate, globose to
subglobose, with brown to black, spinous setae, lacking ostioles, sometimes with a small papillate
ostiole-like structure without an external opening, with Quellkörper incipient. Peridium composed
of brown to hyaline cells of textura angularis to textura prismatica, Munk pores present.
Paraphyses absent. Asci polysporous, unitunicate, clavate to cylindrical, pedicellate or sessile, apex
rounded, apical ring inconspicuous, evanescent. Ascospores arranged obliquely in the ascus in
several longitudinal rows, hyaline to yellowish, allantoid, ellipsoidal to cylindrical, slightly curved,
0–1-septate, smooth-walled, mostly with guttules. Asexual morph: Undetermined (adapted from
Fitzpatrick 1924, Subramanian & Sekar 1990).
Notes – Fracchiaea is typified by F. heterogenea and was clarified as a genus in
Nitschkiaceae based on ascomata with Quellkörper (Fitzpatrick 1924, Subramanian & Sekar 1990).
Subsequently, F. broomeana and F. lunata were sequenced and these strains nested in
Nitschkiaceae (Huhndorf et al. 2004b, Mugambi & Huhndorf 2010). In this study, Fracchiaea is
basal to the clade of Acanthonitschkea and Nitschkia (96%ML/1.00BY, Fig. 1). Coronophora
myricoides is transferred as F. myricoides based on morphology and phylogenetic result (see notes
for F. myricoides).
Fracchiaea heterogenea Sacc., Atti Soc. Veneto-Trent. Sci. Nat. 2(1): 163 (1873) Fig. 16
Facesoffungi number: FoF 10062
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to black,
branched, septate, sinuous hyphae, 4–8 µm wide. Ascomata 350–600 µm (x
̄ = 470 µm, n = 5)
diam., scattered to densely gregarious, immersed to erumpent through the bark of substrate or
superficial, sitting on a subiculum, carbonaceous, black, turbinate, globose to subglobose, the apex
collapsing when dry, tuberculate, surrounded by dark brown to black, tiny spines 13–22 µm high,
with a small papillate ostiole-like structure without an external opening, with Quellkörper incipient.
Peridium 36–95 µm wide, comprising three layers, outer layer composed of carbonaceous, black
920
tissue; middle layer composed of membranaceous, dark brown to brown cells of textura angularis,
rare Munk pore present; inner layer membranaceous, composed of hyaline cells of textura
prismatica. Paraphyses absent. Asci (73–)85–90(–100) × 10–15(–20) µm (x
̄ = 87 × 12 µm, n = 20),
polysporous, unitunicate, clavate to cylindrical, apex rounded, apical ring inconspicuous, with a
short pedicel, evanescent. Ascospores 8–12 × 1.5–2.5 µm (x
̄ = 9.5 × 2 µm, n = 50), arranged
obliquely in the ascus in several longitudinal rows, hyaline, allantoid to cylindrical, slightly curved,
aseptate, rarely 1-septate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from
Fitzpatrick 1924, Subramanian & Sekar 1990).
Material examined – Italy, Veneto, Treviso, on decayed branch of Aesculus, Corylus,
Rhamnus etc., 1873, P.A. Saccardo (UPS-UPS:BOT:F-736393, isotype).
Known hosts and distribution – on decayed wood of Aesculus, Alnus, Buxus, Corylus,
Eucalyptus, Lonicera, Morus, Myrica, Pinus, Quercus, Rhamnus, Rubus etc. in Brazil, France, Italy
(type locality), India, the USA (Saccardo 1873a, Fitzpatrick 1924, Subramanian & Sekar 1990).
Notes – Fracchiaea heterogenea was introduced by Saccardo (1873b) from Italy and is
characterized by tuberculate ascomata, polysporous asci and allantoid ascospores (Saccardo 1873a,
Fitzpatrick 1924, Subramanian & Sekar 1990). Specimen F-736393 collected by Saccardo from
Italy, in 1873 is consistent with the specimen information described by Saccardo (1837a, b).
Therefore, we state that UPS:BOT:F-736393 is isotype of F. heterogenea.
Figure 16 – Fracchiaea heterogenea: (UPS-UPS:BOT:F-736393, isotype). a Material label.
b Material. c Gregarious ascomata. d Ascoma erumpent through the bark of host. e Ascoma cross
section. f Septate hyphae. g Munk pores (arrow). h Peridium. i Spine on ascoma. j–l Asci. m
Ascospores. Scale bars: c–d = 500 µm, e = 200 µm, h = 100 µm, f, j–l = 50 µm, i = 20 µm, g, m =
10 µm.
Fracchiaea myricoides (H.X. Wu & K.D. Hyde) S.K. Huang & K.D. Hyde, comb. nov.
Basionym – Coronophora myricoides H.X. Wu & K.D. Hyde, in Hyde et al., Fungal
Diversity 80: 164 (2016)
921
Index Fungorum number: IF558200
Notes – Coronophora myricoides is characterized by tuberculate ascomata and polysporous
asci, which are similar to the type species of Coronophora, C. gregaria (Hyde et al. 2016). Hyde et
al. (2016) sequenced C. myricoides (IFRD 9201), but lacked discussion on its phylogenetic
analysis. In this study, strain of C. myricoides (IFRD 9201) is sister to Fracchiaea
(100%ML/1.00BY) in Nitschkiaceae (Fig. 1). In addition, C. myricoides is also similar to
Fracchiaea with tuberculate ascomata and polysporous asci. We therefore place C. myricoides in
Fracchiaea based on morphology and phylogenetic result.
Groenhiella Jørg. Koch, E.B.G. Jones & S.T. Moss, Bot. Mar. 26(6): 265 (1983)
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, superficial, sitting on
rarely developed subiculum, carbonaceous to coriaceous, brown to black, reniform to subglobose,
collapsing or laterally collapsing when dry, lacking ostioles. Peridium composed of brown to
hyaline cells of textura angularis to textura prismatica. Pseudoparaphyses numerous, septate,
constricted at the septa, unbranched or branched, evanescent. Asci 8-spored, unitunicate, arising
from a hyaline placenta-like tissue, radially extending into the locule from the base, broadly
clavate, with long pedicel, evanescent. Ascospores hyaline when young, becoming pale brown to
brown, broadly fusiform to ellipsoidal, with papillate at each end, 1-septate, smooth-walled,
guttulate. Appendages initially mucilaginous sheath-like and encapsulate the whole ascospore,
becoming fragment to give rise at each end and middle septum. Asexual morph: Undetermined
(adapted from Koch et al. 1983).
Notes – The monotypic Groenhiella was introduced as a lignicolous marine fungus and
initially placed in Coronophoraceae (Koch et al. 1983). It is characterized by black ascomata,
clavate asci and 1-septate ascospores with fragmented sheath (Koch et al. 1983). Later, this genus
was transferred as a member of Nitschkiaceae (Eriksson & Hawksworth 1987, Lumbsch &
Huhndorf 2010, Maharachchikumbura et al. 2016, Hyde et al. 2020e, Index Fungorum 2021).
Groenhiella bivestia Jørg. Koch, E.B.G. Jones & S.T. Moss, Bot. Mar. 26(6): 265 (1983) Fig. 17
Facesoffungi number: FoF 10063
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, superficial, sitting on
rarely developed subiculum, carbonaceous to coriaceous, brown to black, reniform to subglobose,
collapsing or laterally collapsing when dry, lacking ostioles. Peridium 27–60 µm wide, composed
of brown to hyaline cells of textura angularis to textura prismatica. Pseudoparaphyses numerous,
septate, constricted at the septa, unbranched or branched, evanescent. Asci (25–)35–40 × 13–20 µm
(x
̄ = 38 × 16 µm, n = 10) at ascospore-bearing part, 8-spored, unitunicate, arising from a hyaline
placenta-like tissue, radially extending into the locule from the base, broadly clavate, with long
pedicel, evanescent. Ascospores 12–15 × 6–9 µm (x
̄ = 14 × 8 µm, n = 30), hyaline when young,
becoming pale brown to brown, broadly fusiform to ellipsoidal, with papillate at each end, 1-
septate, smooth-walled, guttulate. Appendages initially mucilaginous sheath-like and encapsulate
the whole ascospore, becoming a fragment at each end and at middle septum. Asexual morph:
Undetermined (adapted from Koch et al. 1983).
Material examined – Denmark, Grønhøj, west coast of Jutland, on strongly decayed drift
wood deposited at base of white dune, partly buried in sand, 16 November 1975, J. Koch (NY-
1315723, microslide of holotype, from the holotype IMI260599 and the slides made by J.J.
Kohlmeyer).
Known hosts and distribution – on decayed wood in Denmark (type locality) (Koch et al.
1983).
Notes – Groenhiella bivestia was introduced by Koch et al. (1983) from Denmark. We were
unable to obtain the holotype IMI260599. Therefore, we examined the permanent slides of
IMI260599 made by Kohlmeyer, which were deposited as a microslide of holotype, NY-1315723.
922
Figure 17 – Groenhiella bivestia: a–n (NY-1315723, microslide of holotype); o (redrawn from
Koch et al. 1983). a–b Ascoma cross section (b in blue colouring agent). c–d Peridium. e–f Asci.
g–o Ascospores. Scale bars: a–b = 200 µm, c–d = 50 µm, f, n = 20 µm, e, g–m = 10 µm, o = 5 µm.
Janannfeldtia Subram. & Sekar, Kavaka 18(1-2): 69 (1993) [1990]
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate or superficial on decorticated wood, coriaceous to membranaceous,
globose to subglobose, dark brown to black, the apex collapsing when dry, lacking ostioles, with
Quellkörper, sitting on thin subiculum composed of brown, septate hyphae. Peridium composed of
brown to hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses
absent. Asci polysporous, arising from a hyaline placenta-like tissue, radially extending into the
locule around the Quellkörper from the base, unitunicate, clavate, apex rounded, apical ring
inconspicuous, pedicellate, evanescent. Ascospores overlapping, hyaline, aseptate, boat-shaped to
reniform, smooth-walled, with a filiform, flagella-like appendage at each end, guttulate. Asexual
morph: Undetermined (adapted from Subramanian & Sekar 1990).
Notes – The monotypic Janannfeldtia is typified by J. karwarensis, and has clavate asci and
reniform ascospores with filiform appendage at each end (Subramanian & Sekar 1990). This genus
923
was initially placed in Coronophorales (Subramanian & Sekar 1990), but assigned to Nitschkiaceae
(Eriksson & Hawksworth 1987, Lumbsch & Huhndorf 2010, Maharachchikumbura et al. 2016,
Hyde et al. 2020e, Index Fungorum 2021). We found that Janannfeldtia is similar to Biciliosporina
in having subglobose ascomata with Quellkörper, clavate asci and ascospores with filiform
appendage at each end, although the latter has 8-spored asci (Subramanian & Sekar 1990).
Janannfeldtia karwarensis Subram. & Sekar, Kavaka 18(1-2): 70 (1993) [1990] Fig 15 n
Facesoffungi number: FoF 10160
Saprobic on wood. Sexual morph: Ascomata solitary to gregarious, immersed to erumpent
thought the bark of substrate or superficial on decorticated wood, sitting on thin subiculum
composed of brown, septate hyphae, coriaceous to membranaceous, globose to subglobose, dark
brown to black, the apex collapsing when dry, lacking ostioles, with Quellkörper. Peridium
composed of brown to hyaline cells of textura angularis to textura prismatica, Munk pores present.
Paraphyses absent. Asci polysporous, arising from a hyaline placenta-like tissue, radially extending
into the locule around the Quellkörper from the base, unitunicate, clavate, apex rounded, apical ring
inconspicuous, pedicellate, evanescent. Ascospores overlapping, hyaline, aseptate, boat-shaped to
reniform, smooth-walled, with a filiform, flagella-like appendage at each end, guttulate. Asexual
morph: Undetermined (adapted from Subramanian & Sekar 1990).
Known hosts and distribution – on decayed wood in India (type locality) (Subramanian &
Sekar 1990).
Notes – Janannfeldtia karwarensis is characterized by subglobose ascomata, polysporous asci
arising from a hyaline placenta-like tissue, radially extending into the locule around the
Quellkörper from the base and ascospores with filiform appendage at each end (Subramanian &
Sekar 1990).
Lasiosphaeriopsis D. Hawksw. & Sivan., in Hawksworth, Trans. Br. mycol. Soc. 74(2): 371 (1980)
Saprobic or parasitic on lichen. Sexual morph: Ascomata perithecial, gregarious, superficial,
sitting on a subiculum composed of brown, septate hyphae, carbonaceous, turbinate, obovoid to
subglobose, black, tuberculate, ostiolate, with periphyses. Peridium comprising black carbonaceous
tissues at outer layer, inner layer composed of membranaceous, brown to hyaline cells of textura
angularis to textura prismatica, Munk pores present. Paraphyses absent or cylindrical, septate,
constricted at the septum. Asci 2-, 4- to 8-spored, unitunicate, clavate to cylindrical, apex rounded,
apical ring inconspicuous, with long pedicel, evanescent. Ascospores uni- to bi-seriate, broadly
fusiform, hyaline, aseptate when young, becoming brown to dark brown, 3-septate, rarely multi-
septate, sometimes constricted at the septum, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Hawksworth 1980, Pérez-Ortega & Halici 2008).
Notes – Lasiosphaeriopsis, a lichenicolous genus, typified by L. salisburyi has tuberculate
ascomata, clavate asci and broadly fusiform ascospores (Hawksworth 1980). Lasiosphaeriopsis
species were reported on Amygdalaria, Peltigera, Pilophorus, Porpidia, Stereocaulon in America,
Asia and Europe (Pérez-Ortega & Halici 2008). This genus was initially placed in Coronophorales
(Hawksworth 1980), and subsequently assigned to Nitschkiaceae (Eriksson & Hawksworth 1987,
Hyde et al. 2020e, Index Fungorum 2021).
Lasiosphaeriopsis salisburyi D. Hawksw. & Sivan., in Hawksworth, Trans. Br. mycol. Soc. 74(2):
373 (1980) Fig. 18
Facesoffungi number: FoF 10064
Saprobic or parasitic on lichen. Sexual morph: Ascomata 170–310 × 210–370 µm (x
̄ = 245 ×
310 µm, n = 10), perithecial, gregarious, superficial, sitting on a subiculum composed of brown,
septate hyphae, 3–7 µm wide, carbonaceous, turbinate, obovoid to subglobose, black, tuberculate,
ostiolate, with periphyses. Peridium 35–110 µm wide, comprising three layers, outer layer
composed of black carbonaceous tissues; middle layer composed of membranaceous, dark brown to
pale brown cells of textura angularis, rarely Munk pores present; inner layer composed of
924
membranaceous, hyaline cells of textura prismatica. Paraphyses 4–9 µm wide, cylindrical, septate,
constricted at the septum, evanescent. Asci (90–)100–110(–120) × (14–)16–18(–22) µm (x
̄ = 105 ×
17 µm, n = 20), 2- to 4-spored, unitunicate, clavate to cylindrical, apex rounded, apical ring
inconspicuous, pedicellate, evanescent. Ascospores (22–)25–30(–35) × 6.5–9 µm (x
̄ = 27 × 8 µm, n
= 50), bi-seriate, broadly fusiform, hyaline, aseptate when young, becoming 3-septate and brown to
olivaceous brown at middle cells, constricted at the septum, smooth-walled, guttulate. Asexual
morph: Undetermined (adapted from Hawksworth 1980).
Material examined – UK, England, Lancashire, Ainsdale, Hillside Sandhills, on thallus of
Peltigera rufescens, 23 March 1951, G. Salisbruy (IMI-80468, holotype).
Known hosts and distribution – on thallus of Peltigera rufescens (Peltigeraceae) in the UK
(type locality) (Hawksworth 1980).
Notes – Lasiosphaeriopsis salisburyi is similar to Rhagadostoma lichenicola, but differs from
it in the possessing of 3-septate ascospores, whereas those of the latter are 0–1-septate
(Hawksworth 1980).
Figure 18 – Lasiosphaeriopsis salisburyi: (IMI-80468, holotype). a Material label. b–d Ascomata
on lichen. e Subiculum composed of brown, septate hyphae. f Peridium. g Ostiole. h–i Asci with
paraphyses. j Munk pore. k–o Ascospores. Scale bars: b = 1 mm, c = 500 µm, d = 200 µm, e = 100
µm, f–i = 50 µm, k–o = 20 µm, j = 10 µm.
Loranitschkia Lar.N. Vassiljeva, Mikol. Fitopatol. 24(3): 207 (1990)
Saprobic on wood. Sexual morph: Ascomata scattered to gregarious, superficial, sitting on
thin subiculum, pyriform to subglobose, tuberculate, black, the apex collapsing when dry, lacking
925
ostioles. Peridium composed of brown to hyaline cells of porous texture. Paraphyses absent. Asci
8-spored, unitunicate, clavate, apex rounded, apical ring inconspicuous, pedicellate, evanescent.
Ascospores bi-seriate, hyaline, clavate, with one middle septum, apex rounded, attenuated in a
filiform, flagella-like, sometimes 1-septate appendage at base, smooth-walled, guttulate. Asexual
morph: Undetermined (adapted from Vasilyeva et al. 2010).
Notes – The monotypic Loranitschkia was introduced by Vasilyeva (1990) from Russia, and
it is characterized by tuberculate ascomata, clavate asci and clavate ascospores with a long flagella-
like appendage (Vasilyeva et al. 2010). This genus is placed in Nitschkiaceae based on the
characteristics of ascomata similar to Nitschkia (Vasilyeva et al. 2010).
Loranitschkia viticola Lar.N. Vassiljeva [as ‘Loranitschki’], Mikol. Fitopatol. 24(3): 207 (1990)
Fig. 15i
Facesoffungi number: FoF 10161
Saprobic on wood. Sexual morph: Ascomata scattered to gregarious, superficial, sitting on
thin subiculum, pyriform to subglobose, black, the apex collapsing when dry, lacking ostioles.
Peridium composed of brown to hyaline cells of porous texture. Paraphyses absent. Asci 8-spored,
unitunicate, clavate, apex rounded, apical ring inconspicuous, pedicellate, evanescent. Ascospores
bi-seriate, hyaline, clavate, with one middle septum, apex rounded, attenuated in a filiform,
flagella-like, sometimes 1-septate appendage at base, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Vasilyeva et al. 2010).
Known hosts and distribution – on decayed wood of Vitis in Russia (type locality) (Vasilyeva
et al. 2010).
Notes – The ascomata of Loranitschkia viticola are similar to Nitschkia species, but the wall
in Loranitschkia viticola is thicker than in Nitschkia species, in addition, the former has porous
texture of ascomata walls, different from the rounded to angular wall cells of the latter (Vasilyeva
et al. 2010).
Nitschkia G.H. Otth ex P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 13 (1873)
Saprobic on wood. Sexual morph: Subiculum absent or composed of brown to dark brown,
septate, branched, sinuous hyphae with metallic iridescence. Ascomata perithecial or without an
external opening, gregarious, erumpent or superficial, sitting on or in a subiculum, carbonaceous or
coriaceous, black, turbinate, cupuliform to subglobose, tuberculate, the apex collapsing when dry,
with periphyses inside of the inconspicuous ostioles or lacking ostioles. Peridium composed of
membranaceous, brown to hyaline cells of textura angularis to textura prismatica, sometimes
Munk pores present. Paraphyses absent or unbranched, septate. Asci 4- to 8-spored, unitunicate,
clavate to cylindrical, apex blunt or rounded, apical ring inconspicuous, with long pedicel,
evanescent. Ascospores bi-seriate to overlapping, hyaline to brown, aseptate or septate, allantoid to
subcylindrical, sometimes slightly curved, smooth-walled, mostly with guttules. Asexual morph:
Undetermined (adapted from Fitzpatrick 1923, Subramanian & Sekar 1990).
Notes – Fuckel (1870b) introduced the generic name Nitschkia and described N. fuckelii
based on Sphaeria cupularis Fries. Subsequently, Karsten (1873) stated that S. cupularis Fries is
illegitimate as it has an earlier homonym S. cupularis Pers.; thus, he clarified the genus Nitschkia
and introduced N. cupularis as generic type. This genus is characterized by turbinate ascomata,
clavate asci and allantoid ascospores and currently includes 58 species (Hyde et al. 2020e, Index
Fungorum 2021). Molecular data is available for five species, N. calyculus, N. grevillii, N.
meniscoidea, N. pezizoidea, N. tetraspora (Huhndorf et al. 2004b, Mugambi & Huhndorf 2010). In
this study, N. calyculus, N. grevillii and N. tetraspora clustered as a sister clade of
Acanthonitschkea (73%ML/0.95BY, Fig. 1) and N. meniscoidea nests in Nitschkiaceae
(100%ML/1.00BY, Fig. 1), but N. pezizoidea is basal to Bertiaceae (62%ML, Fig. 1).
Nitschkia cupularis (Pers.) P. Karst., Bidr. Känn. Finl. Nat. Folk 23: 81 (1873) Fig. 19
Basionym – Sphaeria cupularis Pers., Observ. mycol. (Lipsiae) 1: 65 (1796)
926
Facesoffungi number: FoF 01122
Saprobic on wood. Sexual morph: Ascomata 325–550 × 210–450 µm (x
̄ = 500 × 350 µm, n =
10), gregarious, erumpent through the bark of host to superficial, coriaceous to membranaceous,
black, turbinate, cupuliform to subglobose, tuberculate, the apex collapsing when dry, lacking
ostioles, with a small papillate ostiole-like structure without an external opening and periphyses-
like hyphae. Peridium 30–55 µm wide, comprising three layers, outer layer composed of black
tissue; middle layer composed of membranaceous, dark brown to brown cells of textura angularis,
Munk pores present; inner layer composed of membranaceous, hyaline cells of textura prismatica.
Paraphyses 5–12 µm wide, cylindrical, unbranched, septate, constricted at the septum. Asci (45–
)50–60(–70) × (7–)8–10(–12) µm (x
̄ = 58 × 9 µm, n = 30), 8-spored, unitunicate, clavate, apex
rounded, pedicellate, evanescent. Ascospores (7.5–)10–12(–14) × 2–4 µm (x
̄ = 10.5 × 2.5 µm, n =
50), bi-seriate, hyaline, aseptate, allantoid to subcylindrical, slightly curved, smooth-walled, mostly
with guttules. Asexual morph: Undetermined.
Material examined – Austria, Niederösterreich, Wienerwald, Rekawinkel, on decayed wood
of Prunus padus, September 1949, F. Petrak (S-F117662); Germany, on decayed wood of Acer and
Prunus, K.W. Fuckel (BR-5020096960571).
Known hosts and distribution – on decayed wood of Acer, Prunus, Rubus, Sambucus,
Syringa, Tilia and Ulmus in Austria, Germany (type locality), Russia and the USA (Karsten 1873,
Vasilyeva et al. 2010).
Notes – We were unable to obtain the type material. Therefore, we re-examined the authentic
specimens (F117662 and 5020096960571) collected by Petrak and Fuckel, respectively.
Rhagadostoma Körb., Parerga lichenol. (Breslau) 5: 472 (1865)
Saprobic or parasitic on lichen. Sexual morph: Subiculum absent or composed of brown to
dark brown, septate, branched, sinuous hyphae. Ascomata perithecial, gregarious, superficial,
sitting on or in a subiculum, carbonaceous, black, globose to subglobose, tuberculate, the apex
collapsing when dry, with periphyses inside of the inconspicuous ostioles. Peridium composed of
membranaceous, brown to hyaline cells of textura angularis to textura prismatica, Munk pores
present. Paraphyses absent or unbranched or branched, septate. Asci 2- to 8-spored, unitunicate,
clavate, apex rounded and thickening, apical ring inconspicuous, pedicellate, evanescent.
Ascospores bi-seriate to overlapping, hyaline to brown, 1-septate, rarely multi-septate, ellipsoidal,
broadly fusiform to cylindrical, sometimes slightly curved, smooth-walled, mostly with guttules.
Asexual morph: Undetermined (adapted from Navarro-Rosinés & Hladun 1994, Navarro-Rosinés
et al. 1999).
Notes – Körber (1865) established Rhagadostoma based on R. corrugatum. Subsequently,
Saccardo (1882a) stated that R. corrugatum is a synonym of Bertia lichenicola, and Keissler (1930)
introduced R. lichenicola as the current name of B. lichenicola and R. corrugatum. However, the
first identified R. corrugatum is regarded as the name of the type species of Rhagadostoma
(Maharachchikumbura et al. 2016, Hyde et al. 2020e). This lichenicolous genus was initially placed
in Sordariales (Navarro-Rosinés & Hladun 1994, Navarro-Rosinés et al. 1999). Huhndorf et al.
(2004b) proposed that Rhagadostoma is morphologically related to Bertia but its phylogenetic
position remains unknown. However, subsequently, this genus was assigned to Nitschkiaceae
(Lumbsch & Huhndorf 2010, Hyde et al. 2020e, Index Fungorum 2021).
Rhagadostoma corrugatum Körb., Parerga lichenol. (Breslau) 5: 473 (1865) Fig. 20
Facesoffungi number: FoF 10065
Saprobic or parasitic on lichen. Sexual morph: Subiculum absent or composed of brown to
dark brown, septate, branched, sinuous hyphae 60–90 µm wide. Ascomata 300–420 µm (x
̄ = 370
µm, n = 10) diam., perithecial, gregarious, superficial, sitting on or in a subiculum, carbonaceous,
black, globose to subglobose, tuberculate, the apex collapsing when dry, with periphyses inside of
the inconspicuous ostioles. Peridium 60–90 µm wide, comprising three layers, outer layer
composed of black tissue; middle layer composed of membranaceous, brown to olivaceous brown
927
cells of textura angularis, Munk pores present; inner layer composed of membranaceous, hyaline
cells of textura prismatica. Paraphyses absent. Asci 2- to 4-spored, unitunicate, clavate, apex
rounded and thickening, apical ring inconspicuous, pedicellate, evanescent. Ascospores (40–)45–
55(–68) × (6–)8–10(–15) µm (x
̄ = 50 × 9.5 µm, n = 50), bi-seriate to overlapping, hyaline to brown,
1-septate, rarely multi-septate, ellipsoidal, broadly fusiform to cylindrical, sometimes slightly
curved, smooth-walled, mostly with guttules. Asexual morph: Undetermined.
Figure 19 – Nitschkia cupularis: a, c, e, h, m–n, t–u (S-F117662); b, d, f–g, i–l, o–s (BR-
5020096960571). a–b Material. c–d Ascomata erumpent through the bark of host. e, g Ascoma
cross section. f Munk pores (arrow). h Ostiole-like structure without an external opening.
i Peridium. j–n Asci. o Paraphyses. p–u Ascospores. Scale bars: c–d = 1 mm, g = 200 µm, e = 100
µm, h–i = 50 µm, f, j–o = 20 µm, p–u = 5 µm.
Material examined – Norway, Oppland, Sønderland, Nymoen, on Solorina crocea growing
on Calamagrostis, 4 September 1984, O.E. Eriksson and J.Z. Yue (NY-03380689).
Known hosts and distribution – on thallus of Solorina crocea in Austria, Finland, Germany,
Italy, Norway, Spain (type locality) and the UK (Körber 1865, Saccardo 1882a, Olivier 1907,
Hafellner 2015).
928
Notes – Rhagadostoma corrugatum was found on Solorina crocea (Körber 1865). We were
unable to obtain the type material and, therefore, re-examined an authentic sample on S. crocea.
Figure 20 – Rhagadostoma corrugatum: (NY-03380689). a Material. b–d Ascomata on the host.
e Ascoma cross section. f Munk pores (arrow). g Subiculum composed of brown hyphae.
h Peridium. i Ascus. j–q Ascospores. Scale bars: b = 1 mm, c–d = 500 µm, e = 200 µm, h = 100
µm, g, i = 50 µm, f, j–q = 20 µm.
Rhagadostomella Etayo, Biblthca Lichenol. 84: 109 (2002)
Saprobic or parasitic on lichen. Sexual morph: Ascomata perithecial, gregarious, erumpent to
superficial, sitting on a subiculum, carbonaceous, olivaceous brown or dark brown to black,
globose to subglobose, tuberculate, the apex collapsing when dry, ostiolate, periphysate. Peridium
composed of membranaceous, brown cells of textura epidermoidea to textura angularis.
Paraphyses absent. Asci 4- to 8-spored, unitunicate or bitunicate, broadly clavate, apex blunt to
rounded and thickening, apical ring conspicuous after the ascospores released, pedicellate,
evanescent. Ascospores overlapping at the apex, hyaline, aseptate to 1- or 2-septate, fusiform to
929
cylindrical, sometimes slightly curved, smooth-walled, mostly with guttules. Asexual morph:
Undetermined (adapted from Etayo 2002, Flakus et al. 2019).
Notes – Rhagadostomella, a lichenicolous genus, was introduced by Etayo (2002) based on
R. gregaria from Colombia. A second species R. hypolobariella which has bitunicate asci and
growing on Lobariella pallida in Bolivia was introduced (Flakus et al. 2019). This genus is placed
in Nitschkiaceae but no molecular data is available for the Rhagadostomella (Etayo 2002, Flakus et
al. 2019).
Rhagadostomella gregaria Etayo, Biblthca Lichenol. 84: 109 (2002) Fig. 15r–s
Facesoffungi number: FoF 10162
Saprobic or parasitic on lichen. Sexual morph: Ascomata perithecial, gregarious, erumpent to
superficial, sitting on a subiculum, carbonaceous, dark brown to black, globose to subglobose,
tuberculate, the apex collapsing when dry, ostiolate, periphysate. Peridium composed of
membranaceous, brown cells of textura epidermoidea to textura angularis. Paraphyses absent. Asci
8-spored, unitunicate, broadly clavate, apex blunt and thickening, apical ring conspicuous after the
ascospores released, pedicellate, evanescent. Ascospores overlapping at the apex, hyaline, 1-septate,
fusiform to cylindrical, sometimes slightly curved, smooth-walled, mostly with guttules. Asexual
morph: Undetermined (adapted from Etayo 2002).
Known hosts and distribution – on thallus of Sticta weigelii in Colombia (type locality) and
Ecuador (Etayo 2002, 2017).
Notes – Rhagadostomella gregaria was collected on Sticta weigelii in Colombia and Ecuador
and is characterzied by tuberculate ascomata, broadly clavate asci and fusiform to cylindrical
ascospores (Etayo 2002, 2017).
Tortulomyces Lar.N. Vassiljeva, S.L. Stephenson, Chernyshev & K.D. Hyde, Mycoscience 54(1):
111 (2013)
Saprobic on wood. Sexual morph: Subiculum composed of brown, branched, septate, sinuous
hyphae. Ascomata gregarious, superficial, sitting on a subiculum, brown to black, turbinate,
subglobose, tuberculate, the apex collapsing when dry, lacking ostioles. Peridium composed of
brown to hyaline cells of textura angularis to textura prismatica. Paraphyses cylindrical, septate.
Asci 8-spored, unitunicate, clavate to cylindrical, apex rounded, apical ring inconspicuous, with
long pedicel, evanescent. Ascospores bi-seriate to overlapping, pale brown, 1-septate, ellipsoidal,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Vasilyeva et al. 2013).
Notes – The monotypic genus Tortulomyces was introduced by Vasilyeva et al. (2013) for T.
thailandicus growing on dead branches in Thailand. Tortulomyces was placed in Nitschkiaceae, but
no molecular data is available for this genus (Vasilyeva et al. 2013).
Tortulomyces thailandicus Lar.N. Vassiljeva, S.L. Stephenson, Chernyshev & K.D. Hyde,
Mycoscience 54(1): 111 (2013) Fig. 15u
Facesoffungi number: FoF 10163
Saprobic on wood. Sexual morph: Subiculum composed of brown, branched, septate, sinuous
hyphae. Ascomata gregarious, superficial, sitting on a subiculum, brown to black, turbinate,
subglobose, tuberculate, the apex collapsing when dry, lacking ostioles. Peridium composed of
brown to hyaline cells of textura angularis to textura prismatica. Paraphyses cylindrical, septate.
Asci 8-spored, unitunicate, clavate to cylindrical, apex rounded, apical ring inconspicuous, with
long pedicel, evanescent. Ascospores bi-seriate to overlapping, pale brown, 1-septate, ellipsoidal,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Vasilyeva et al. 2013).
Known hosts and distribution – on decayed wood in Thailand (type locality) (Vasilyeva et al.
2013).
Notes – Tortulomyces thailandicus grows on dead branches in Thailand and has subglobose
ascomata without ostioles, clavate asci and ellipsoidal ascospores, but there is no available
molecular data for this species (Vasilyeva et al. 2013).
930
Scortechiniaceae Huhndorf, A.N. Mill. & F.A. Fernández, in Huhndorf, Miller & Fernández,
Mycol. Res. 108(12): 1387 (2004)
Saprobic on wood. Sexual morph: Subiculum absent or composed of numerous or rare, brown
to dark brown, branched, septate hyphae, with dichotomously branched, tapering or rounded ends.
Ascomata scattered or gregarious, semi-immersed or erumpent or superficial, sitting on or in a
subiculum, carbonaceous or coriaceous, brown to black, turbinate, globose, obpyriform to
subglobose, the apex collapsing when dry, tuberculate, glabrous or spiny, lacking ostioles, with a
conical Quellkörper. Peridium usually outer layer composed of carbonaceous, black tissue; inner
layer composed of membranaceous, brown to hyaline cells of textura angularis to textura
prismatica, sometimes Munk pores present. Paraphyses absent or filiform, branched or
unbranched, septate, evanescent. Asci 8- or multi-spored, unitunicate, clavate to cylindrical,
pedicellate or sessile, apex blunt or rounded, without apical ring, evanescent. Ascospores bi-seriate
to overlapping, hyaline to brown or olivaceous brown, allantoid, ellipsoidal, ovoid to oval, straight
to slightly curved, usually aseptate, smooth-walled, sometimes with mucilaginous appendages,
mostly with guttules. Asexual morph: Conidiophores erect, solitary, straight to flexuous.
Conidiogenous cells hyaline, smooth-walled, subcylindrical. Conidia hyaline, aseptate, smooth-
walled, ellipsoidal to ovoid, guttulate (adapted from Huhndorf et al. 2004b, Hyde et al. 2020e).
Type genus – Scortechinia Sacc.
Notes – Scortechiniaceae was established by Huhndorf et al. (2004b) based on the well-
supported clade of Euacanthe, Neofracchiaea and Scortechinia in LSU sequence analyses. The
Quellkörper was accepted as a main character to define this family (Huhndorf et al. 2004b). Later,
Biciliospora, Coronophorella, Cryptosphaerella, Pseudocatenomycopsis, Scortechiniella,
Scortechiniellopsis and Tympanopsis, were accommodated in this family based on phylogenetic
results (Mugambi & Huhndorf 2010, Crous et al. 2013, Hyde et al. 2020e). The members in this
family are characterized by turbinate ascomata with Quellkörper, clavate asci and hyaline
ascospores (Huhndorf et al. 2004b, Mugambi & Huhndorf 2010).
We found that Cryptosphaerella should be redefined. Initially, Cryptosphaerella was
introduced as a genus with similar morphology to Cryptosphaeria (Diatrypaceae) (Saccardo
1882a). Later, it was considered a member of Coronophoraceae (von Höhnel 1907).
Cryptosphaerella is typified by C. nitschkei which has perithecial, immersed ascomata with ostioles
and polysporous asci (Nitschke 1870, Saccardo 1882a). Some species with similar characteristics
(such as C. anonae, C. coffaeiphila and C. cordobensis) were subsequently introduced to this
genus, but no molecular data is available for these species (Spegazzini 1909, 1922, 1926).
Mugambi & Huhndorf (2010) sequenced six ‘Cryptosphaerella’ species lacking ostioles in the
ascomata with Quellkörper. Therefore, Cryptosphaerella was assigned to Scortechiniaceae based
on phylogenetic analyses of these six species. However, the type of Cryptosphaerella does not have
unique Quellkörpers and Munk pores. We therefore transfer Cryptosphaerella to Sordariomycetes
genera incertae sedis and redefine the six species introduced by Mugambi & Huhndorf (2010) as
Pseudocryptosphaerella and Neocryptosphaerella in Scortechiniaceae based on morphology and
phylogenetic results.
Biciliospora Petr., Sydowia 6(5-6): 429 (1952)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, rounded ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, carbonaceous, brown to black, globose to
subglobose, the apex collapsing when dry, tuberculate, glabrous, lacking ostioles, with a central,
conical Quellkörper at apex. Peridium outer layer composed of carbonaceous, black tissue; inner
layer composed of membranaceous, brown to hyaline cells of textura angularis to textura
prismatica, Munk pores present. Paraphyses absent. Asci 8-spored, arising from a hyaline placenta-
like tissue, radially extending into the locule around the Quellkörper from the base, unitunicate,
clavate, pedicellate, apex rounded, without apical ring, evanescent. Ascospores bi-seriate to
overlapping, hyaline, ellipsoidal to fusiform, aseptate, smooth-walled, guttulate, with a filiform
931
appendage at each end and usually recurved at opposite sides. Asexual morph: Undetermined
(adapted from Petrak 1952).
Notes – The monotypic Biciliospora was introduced by Petrak (1952) for B. velutina and
initially placed in Coronophorales. Subsequently, this genus was assigned in Coronophoraceae and
even accepted as a synonym of Nitschkia (Müller & von Arx 1973, Nannfeldt 1975, Subramanian
& Sekar 1990). However, molecular data of Biciliospora revealed that this genus nested in
Scortechiniaceae (Mugambi & Huhndorf 2010). In this study, Biciliospora is sister to
Scortechiniella (85%ML/1.00BY, Fig. 1) in Scortechiniaceae.
Biciliospora velutina Petr., Sydowia 6(5-6): 429 (1952) Fig. 21
Facesoffungi number: FoF 10066
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae 4.5–7.5 µm wide, with dichotomously branched, rounded ends. Ascomata
340–450 µm (x
̄ = 380 µm, n = 10) diam., scattered or gregarious, superficial, sitting in a subiculum,
carbonaceous, brown to black, globose to subglobose, the apex collapsing when dry, tuberculate,
glabrous, lacking ostioles, with a central, conical Quellkörper at apex. Peridium 25–45 µm wide,
comprising three layers, outer layer composed of carbonaceous, black tissue; middle layer
composed of membranaceous, brown to reddish-brown cells of textura angularis, Munk pore
present; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses
absent. Asci (65–)70–85(–90) × (14–)16–20(–22) µm (x
̄ = 75 × 18 µm, n = 20) at the ascospore-
bearing part, 8-spored, arising from a hyaline placenta-like tissue, radially extending into the locule
around the Quellkörper from the base, unitunicate, clavate, pedicellate, apex rounded, without
apical ring, evanescent. Ascospores 25–30(–35) × 5–8 µm (x
̄ = 28 × 6 µm, n = 50), bi-seriate to
overlapping, hyaline, ellipsoidal to fusiform, aseptate, smooth-walled, guttulate, with a filiform
appendage at each end and usually recurved at opposite sides. Asexual morph: Undetermined.
Material examined – Porto Rico, Utaada, on fallen branches in coffee grove, 20 October
1920, F.S. Earle (W-11051, holotype).
Known hosts and distribution – on fallen branches in coffee grove in Porto Rico (type
locality) and Kenya (Petrak 1952, Mugambi & Huhndorf 2010).
Notes – Biciliospora velutina was introduced by Petrak (1952) from Porto Rico and was
sequenced by Mugambi & Huhndorf (2010). In this study, B. velutina is sister to Scortechiniella
similis (85%ML/1.00BY, Fig. 1), differing from the latter in having polysporous asci with oval
ascospores (von Arx & Müller 1954).
Coronophorella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1507 (1909)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, tapering ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, tuberculate, glabrous, lacking ostioles, with a central, conical
Quellkörper touching the bottom of the locule. Peridium composed of membranaceous, brown to
hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses absent.
Asci 8-spored, arising from a hyaline placenta-like tissue, radially extending into the locule around
the Quellkörper from the base, unitunicate, clavate to cylindrical, pedicellate, apex rounded,
without apical ring, evanescent. Ascospores bi-seriate to overlapping, hyaline, reniform to allantoid,
aseptate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Penzig &
Saccardo 1897, Subramanian & Sekar 1990).
Notes – The monotypic Coronophorella was initially placed in Coronophorales, and was
considered to be a synonym of Nitschkia, Scortechinia and Tympanopsis based on morphology
(Penzig & Saccardo 1897, von Arx & Müller 1954, Müller & Arx 1955, Nannfeldt & Santesson
1975). However, this genus was an independent branch of Scortechiniaceae in phylogenetic result
(Mugambi & Huhndorf 2010). In this study, Coronophorella is basal to Biciliospora,
Pseudocryptosphaerella, Scortechiniella and Scortechiniellopsis in Scortechiniaceae (Fig. 1).
932
Figure 21 – Biciliospora velutina: (W-11051, holotype). a Material. b–d Ascomata sitting in a
subiculum on host. e Ascoma cross section (in Congo red reagent). f Peridium. g Munk pores
(arrow). h Septate hyphae. i–j Asci. k–r Ascospores. Scale bars: b = 1 mm, c–d = 500 µm, e = 200
µm, f, h–j = 50 µm, g, k–r = 20 µm.
Coronophorella chaetomioides (Penz. & Sacc.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 1507 (1909) Fig. 24n–o
Basionym – Enchnoa chaetomioides Penz. & Sacc., Malpighia 11(9-10): 390 (1897)
Facesoffungi number: FoF 10164
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, tapering ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, tuberculate, glabrous, lacking ostioles, with a central, conical
Quellkörper almost or touching the bottom of the locule. Peridium composed of membranaceous,
brown to hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses
absent. Asci 8-spored, arising from a hyaline placenta-like tissue, radially extending into the locule
around the Quellkörper from the base, unitunicate, clavate to cylindrical, pedicellate, apex rounded,
without apical ring, evanescent. Ascospores bi-seriate to overlapping, hyaline, reniform to allantoid,
aseptate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Penzig &
Saccardo 1897, Subramanian & Sekar 1990).
933
Known hosts and distribution – on decayed wood in Ghana, India, Indonesia (type locality),
Kenya and Sierra Leone (Penzig & Saccardo 1897, Nannfeldt & Santesson 1975, Subramanian &
Sekar 1990, Mugambi & Huhndorf 2010).
Notes – Coronophorella chaetomioides was originally introduced as Enchnoa chaetomioides
from Indonesia (Penzig & Saccardo 1897). It is characterized by ascomata without ostioles, clavate
asci and reniform to allantoid ascospores (Penzig & Saccardo 1897, Subramanian & Sekar 1990).
Mugambi & Huhndorf (2010) accepted this species in Scortechiniaceae based on molecular data. In
this study, C. chaetomioides nests in Scortechiniaceae (Fig. 1).
Euacanthe Theiss., Annls mycol. 15(3/4): 272 (1917)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, tapering ends. Ascomata densely
gregarious, superficial, sitting on a subiculum, coriaceous, black, turbinate, cupuliform to
subglobose, the apex collapsing when dry, tuberculate, with spiculate, erect setae, lacking ostioles,
with a central, conical Quellkörper. Peridium composed of membranaceous, brown to hyaline cells
of textura angularis to textura prismatica, Munk pores present. Paraphyses absent. Asci 8-spored,
arising from a hyaline placenta-like tissue, radially extending into the locule around the
Quellkörper from the base, unitunicate, clavate, pedicellate, apex rounded, without apical ring,
evanescent. Ascospores bi-seriate to overlapping, hyaline, reniform, aseptate, smooth-walled,
guttulate. Asexual morph: Undetermined (adapted from Engler 1895, Subramanian & Sekar 1990).
Notes – The monotypic Euacanthe was introduced based on Meliolopsis usambarensis
(Theissen 1917). Nannfeldt & Santesson (1975) transferred this genus as Acanthonitschkea
(Nitschkiaceae) and placed E. usambarensis as a synonym of A. foveolata. Subsequently,
Subramanian & Sekar (1990) assigned Euacanthe in Coronophorales and accepted E. foveolata
based on A. foveolata as the only species in this genus. Mugambi & Huhndorf (2010) sequenced E.
foveolata and found it nested in Scortechiniaceae. We use the original name, E. usambarensis, as
the current name of type species to avoid confusion in future work. In this study, Euacanthe is
sister to Pseudocatenomycopsis (0.97BY, Fig. 1) in Scortechiniaceae.
Euacanthe usambarensis (Rehm) Theiss., Annls mycol. 15(3/4): 272 (1917) Fig. 22
Basionym – Meliolopsis usambarensis Rehm, in Hennings, Pflanzenw. Ost-Afrikas
Nachbarg., Teil C: 31 (1895)
Facesoffungi number: FoF 10067
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae 6–14 µm wide, with dichotomously branched, tapering ends. Ascomata
630–1250 × 420–690 µm (x
̄ = 985 × 550 µm, n = 10), densely gregarious, superficial, sitting on a
subiculum, coriaceous, black, turbinate, cupuliform to subglobose, the apex collapsing when dry,
tuberculate, with spiculate, erect setae 10–15 µm wide, lacking ostioles, with a central, conical
Quellkörper. Peridium 60–90 µm wide, membranaceous, comprising two layers, outer layer
composed of dark brown to brown cells of textura angularis, Munk pores present; inner layer
composed of hyaline cells of textura prismatica. Paraphyses absent. Asci (14–)18–21(–23) × 7–9
µm (x
̄ = 20 × 8 µm, n = 20) at ascospore-bearing part, 8-spored, arising from a hyaline placenta-
like tissue, radially extending into the locule around the Quellkörper from the base, unitunicate,
clavate, pedicellate, apex rounded, without apical ring, evanescent. Ascospores (4.5–)5–6(–7) ×
2.5–4 µm (x
̄ = 5.5 × 3 µm, n = 50), bi-seriate to overlapping, hyaline, reniform, aseptate, smooth-
walled, guttulate. Asexual morph: Undetermined (adapted from Engler 1895, Subramanian & Sekar
1990).
Material examined – Tanzania, Usambara, Kombolo-Waldbestände, on decayed wood,
February 1893, C. Holst (S-F12537, holotype); ibid. (S-F12538, isotype).
Known hosts and distribution – on decayed wood in India, Kenya and Tanzania (type
locality) (Engler 1895, Subramanian & Sekar 1990, Mugambi & Huhndorf 2010).
934
Notes – Euacanthe usambarensis was introduced based on Meliolopsis usambarensis
(Theissen 1917). Nannfeldt & Santesson (1975) proposed that Sphaeria foveolata and E.
usambarensis were synonym of Acanthonitschkea foveolata. Later, Subramanian & Sekar (1990)
accepted E. foveolata as type species based on A. foveolata, and Mugambi & Huhndorf (2010)
sequenced E. foveolata (GKM1221). However, we use the original E. usambarensis as its generic
type. This species is sister to the asexual species Pseudocatenomycopsis rothmanniae (0.97BY, Fig.
1) in Scortechiniaceae based on phylogenetic result.
Figure 22 – Euacanthe usambarensis: a–b, f, h–j, m–n, q (S-F12537, holotype); c–e, g, k–l, o–p (S-
F12538, isotype). a–d Ascomata sitting on a subiculum on host. e–f Ascoma cross section.
g, j Conical Quellkörper. h Munk pores (arrow). i Peridium. k–n Asci. o Septate hyphae.
p–q Ascospores. Scale bars: a = 2 mm, b–c = 1 mm, d = 500 µm, e–f = 200 µm, g, i–j, o = 100 µm,
h = 20 µm, k–n, p–q = 10 µm.
935
Neocryptosphaerella S.K. Huang & K.D. Hyde, gen. nov.
Etymology: Name refers to genus being similar to Cryptosphaerella.
Facesoffungi number: FoF 10165; Index Fungorum number: IF558354
Saprobic on wood. Sexual morph: Ascomata solitary or scattered, immersed to erumpent,
black, turbinate, the apex collapsing when dry, tuberculate, sitting in a subiculum, lacking ostioles,
with a central, conical Quellkörper. Peridium composed of brown to hyaline cells of textura
angularis to textura prismatica, Munk pores present. Paraphyses absent. Asci polysporous,
unitunicate, clavate, with long pedicel, apex rounded, without apical ring, evanescent. Ascospores
overlapping, hyaline, allantoid to cylindrical, slightly curved, aseptate, smooth-walled, guttulate.
Asexual morph: Undetermined (adapted from Mugambi & Huhndorf 2010).
Notes – We establish Neocryptosphaerella based on Cryptosphaerella celata (type) and C.
globosa. Mugambi & Huhndorf (2010) introduced six ‘Cryptosphaerella’ species having ascomata
lacking ostioles and a Quellkörper. The characteristics of the type of Cryptosphaerella are ostiolate
ascomata lacking a Quellkörper (Saccardo 1882a) (see notes for Cryptosphaerella). Therefore,
Cryptosphaerella should not belong to Scortechiniaceae and these six species do not belong to
Cryptosphaerella based on morphology. Phylogenetically, C. celata and C. globosa form an
independent clade and the remaining four species form another clade in Scortechiniaceae
(Mugambi & Huhndorf 2010). The phylogenetic position of the type of Cryptosphaerella is
undetermined. We therefore introduce Neocryptosphaerella for C. celata and C. globosa. This
genus is sister to Biciliospora, Coronophorella, Pseudocryptosphaerella, Scortechiniella and
Scortechiniellopsis in Scortechiniaceae, and has ascomata lacking ostioles, Quellkörper and
polysporous asci with allantoid to cylindrical ascospores (Mugambi & Huhndorf 2010, this study,
0.98BY, Fig. 1).
Neocryptosphaerella celata (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb. nov.
Basionym – Cryptosphaerella celata Mugambi & Huhndorf, Mycologia 102(1): 197 (2010)
Facesoffungi number: FoF 10166; Index Fungorum number: IF558348
Saprobic on wood. Sexual morph: Ascomata solitary or scattered, immersed to erumpent,
black, turbinate, the apex collapsing when dry, tuberculate, sitting in a subiculum, lacking ostioles,
with a central, conical Quellkörper. Peridium composed of brown to hyaline cells of textura
angularis to textura prismatica, Munk pores present. Paraphyses absent. Asci polysporous,
unitunicate, clavate, with long pedicel, apex rounded, without apical ring, evanescent. Ascospores
overlapping, hyaline, allantoid to cylindrical, slightly curved, aseptate, smooth-walled, guttulate.
Asexual morph: Undetermined (adapted from Mugambi & Huhndorf 2010).
Known hosts and distribution – on decorticated wood in Kenya (type locality) (Mugambi &
Huhndorf 2010).
Notes – Neocryptosphaerella celata is sister to N. globosa (84%ML/1.00BY, Fig. 1),
differing in the size of ascospores (Mugambi & Huhndorf 2010).
Neocryptosphaerella globosa (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb. nov.
Basionym – Cryptosphaerella globosa Mugambi & Huhndorf, Mycologia 102(1): 201 (2010)
Index Fungorum number: IF558353
Neofracchiaea Teng, Sinensia, Shanghai 9: 255 (1938)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, rounded ends. Ascomata densely
gregarious, superficial, sitting in a subiculum, coriaceous, black, globose to subglobose, the apex
collapsing when dry, tuberculate, lacking ostioles, with a central, conical Quellkörper. Peridium
composed of membranaceous, brown to hyaline cells of textura angularis to textura prismatica,
Munk pores present. Paraphyses absent. Asci polysporous, arising from a hyaline placenta-like
tissue, radially extending into the locule around the Quellkörper from the base, unitunicate, clavate,
pedicellate, apex rounded, without apical ring, evanescent. Ascospores overlapping, hyaline,
936
cylindrical to oblong, slightly curved, aseptate, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Saccardo 1882b).
Notes – The monotypic Neofracchiaea was introduced by Teng (1938) based on Cucurbitaria
callista, which was considered to be a member of Cryptosphaerella, Fracchiaea or Nitschkia based
on similar charcteristics (Saccardo 1882a, Fitzpatrick 1924, Nannfeldt & Santesson 1975).
However, Neofracchiaea nested in Scortechiniaceae as an independent clade based on phylogenetic
results (Huhndorf et al. 2004b, Mugambi & Huhndorf 2010). In this study, Neofracchiaea is sister
to Scortechinia and Tympanopsis (Fig. 1).
Neofracchiaea callista (Berk. & M.A. Curtis) Teng, Sinensia, Shanghai 9: 255 (1938) Fig. 23
Basionym – Cucurbitaria callista Berk. & M.A. Curtis, N. Amer. Fung.: no. 821 (1875)
Facesoffungi number: FoF 10068
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae 3–7 µm wide, with dichotomously branched, rounded ends. Ascomata
200–350 µm (x
̄ = 260 µm, n = 20) diam., densely gregarious, superficial, sitting in a subiculum,
coriaceous, black, globose to subglobose, the apex collapsing when dry, tuberculate, lacking
ostioles, with a central, conical Quellkörper. Peridium 20–55 µm wide, membranaceous,
comprising two layers, outer layer composed of brown to olivaceous brown cells of textura
angularis, Munk pores present; inner layer composed of hyaline cells of textura prismatica.
Paraphyses absent. Asci (30–)40–50(–60) × (9–)12–15(–18) µm (x
̄ = 45 × 14 µm, n = 20) at
ascospore-bearing part, polysporous, arising from a hyaline placenta-like tissue, radially extending
into the locule around the Quellkörper from the base, unitunicate, clavate, pedicellate, apex
rounded, without apical ring, evanescent. Ascospores (5.5–)6.5–8(–10) × 1.5–2.6 µm (x
̄ = 7.5 × 2
µm, n = 50), overlapping, hyaline, cylindrical to oblong, slightly curved, aseptate, smooth-walled,
guttulate. Asexual morph: Undetermined.
Material examined – USA, Maryland, Beltsville, on decayed wood, October 1950, F. Petrak
(PDD-54147); ibid. 16 June 1950 (PDD-60976).
Known hosts and distribution – on decayed wood in China and the USA (type locality)
(Berkeley 1875, Saccardo 1882a, Teng 1938).
Notes – We were unable to obtain type material and therefore, re-examined the authentic
samples collected by Petrak from the USA.
Pseudocatenomycopsis Crous & L.A. Shuttlew., in Crous et al., Persoonia 31: 221 (2013)
Saprobic on wood. Sexual morph: Undetermined. Asexual morph: Mycelium hyaline,
branched, septate. Conidiophores erect, hyaline to brown, straight to flexuous, branched, septate.
Conidiogenous cells hyaline to brown, subcylindrical, with flattened. Ramoconidia hyaline to
brown, 0–1-septate, ellipsoidal to subcylindrical, smooth-walled, forming conidial chains. Conidia
hyaline, ellipsoidal to ovoid, smooth-walled (adapted from Crous et al. 2013).
Notes – The monotypic Pseudocatenomycopsis was introduced for P. rothmanniae (Crous et
al. 2013). This asexual genus is considered to be similar to Catenomycopsis in morphology and in
phylogenetic position is sister to Euacanthe (Crous et al. 2013, this study, Fig. 1).
Pseudocatenomycopsis rothmanniae Crous & L.A. Shuttlew., in Crous et al., Persoonia 31: 221
(2013)
Facesoffungi number: FoF 10167
Saprobic on wood. Sexual morph: Undetermined. Asexual morph: Mycelium hyaline,
branched, septate. Conidiophores erect, hyaline to brown, straight to flexuous, branched, septate.
Conidiogenous cells hyaline to brown, subcylindrical, with flattened. Ramoconidia hyaline to
brown, 0–1-septate, ellipsoidal to subcylindrical, smooth-walled, forming conidial chains. Conidia
hyaline, ellipsoidal to ovoid, smooth-walled (adapted from Crous et al. 2013).
Known hosts and distribution – on stems of Rothmannia engleriana (Rubiaceae) in Zambia
(type locality) (Crous et al. 2013).
937
Notes – Pseudocatenomycopsis rothmanniae is sister to Euacanthe usambarensis (0.97BY,
Fig. 1)
Figure 23 – Neofracchiaea callista: a–c, e, h–j, i, n (PDD-54147); d, f–h, j–m (PDD-60976).
a Material. b–d Ascomata sitting in a subiculum on host. e–f Ascoma cross section. g Peridium.
h Septate hyphae. i Munk pores (arrow). j–m Asci. n Ascospores. Notes: e–f, h, j–m are stained in
Congo red reagent. Scale bars: b = 1 mm, c–d = 200 µm, e–f, j = 100 µm, g–h, k–n = 20 µm, i = 10
µm.
Pseudocryptosphaerella S.K. Huang & K.D. Hyde, gen. nov.
Etymology: Name refers to genus being similar to Cryptosphaerella.
Facesoffungi number: FoF 10168; Index Fungorum number: IF558359
Saprobic on wood. Sexual morph: Ascomata solitary or scattered, immersed to erumpent,
black, turbinate, the apex collapsing when dry, tuberculate, sitting in a subiculum, lacking ostioles,
with a central, conical Quellkörper. Peridium composed of brown to hyaline cells of textura
angularis to textura prismatica, Munk pores present. Paraphyses absent. Asci 8- to multi-spored,
unitunicate, clavate, with long pedicel, apex rounded, without apical ring, evanescent. Ascospores
938
overlapping, hyaline, ellipsoidal cylindrical to broadly fusiform, slightly curved, aseptate, smooth-
walled, guttulate. Asexual morph: Undetermined (adapted from Mugambi & Huhndorf 2010).
Notes – Mugambi & Huhndorf (2010) introduced and sequenced four ‘Cryptosphaerella’
species, C. costaricensis, C. cylindriformis, C. elliptica and C. malindensis. These species clustered
in a clade and are characterized by lacking ostioles in the ascomata and a Quellkörper, polysporous
asci and ellipsoidal to broadly fusiform ascospores (Mugambi & Huhndorf 2010). In this study,
they cluster with 100%ML/1.00BY support (Fig. 1) and are sister to Biciliospora, Scortechiniella
and Scortechiniellopsis in Scortechiniaceae. Therefore, we accept these species as members of
Pseudocryptosphaerella based on phylogenetic result.
Pseudocryptosphaerella elliptica (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb. nov.
Basionym – Cryptosphaerella elliptica Mugambi & Huhndorf, Mycologia 102(1): 201 (2010)
Facesoffungi number: FoF 10169; Index Fungorum number: IF558355
Saprobic on wood. Sexual morph: Ascomata solitary or scattered, immersed to erumpent,
black, turbinate, the apex collapsing when dry, tuberculate, with tooth-like setae, sitting in a
subiculum, lacking ostioles, with a central, conical Quellkörper. Peridium composed of brown to
hyaline cells of textura angularis to textura prismatica, Munk pores present. Paraphyses absent.
Asci polysporous, unitunicate, clavate, with long pedicel, apex rounded, without apical ring,
evanescent. Ascospores overlapping, hyaline, ellipsoidal to broadly fusiform, aseptate, smooth-
walled, guttulate. Asexual morph: Undetermined (adapted from Mugambi & Huhndorf 2010).
Known hosts and distribution – on decayed wood in Ecuador (type locality) (Mugambi &
Huhndorf 2010).
Notes – Pseudocryptosphaerella elliptica is sister to P. cylindriformis (100%ML/1.00BY,
Fig. 1), differing from the latter which has cylindrical to oblong ascospores.
Pseudocryptosphaerella costaricensis (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb.
nov. Basionym – Cryptosphaerella costaricensis Mugambi & Huhndorf, Mycologia 102(1): 199
(2010)
Index Fungorum number: IF558356
Pseudocryptosphaerella cylindriformis (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb.
nov. Basionym – Cryptosphaerella cylindriformis Mugambi & Huhndorf, Mycologia 102(1): 199
(2010)
Index Fungorum number: IF558357
Pseudocryptosphaerella malindensis (Mugambi & Huhndorf) S.K. Huang & K.D. Hyde, comb.
nov. Basionym – Cryptosphaerella malindensis Mugambi & Huhndorf [as 'malindensis'],
Mycologia 102(1): 202 (2010)
Index Fungorum number: IF558358
Scortechinia Sacc., Syll. fung. (Abellini) 9: 604 (1891)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, spinous ends. Ascomata densely
gregarious, superficial, sitting in a subiculum, carbonaceous, black, with metallic iridescence,
turbinate, globose to subglobose, the apex collapsing when dry, glabrous, lacking ostioles, with a
central, conical Quellkörper. Peridium composed of membranaceous, brown to hyaline cells of
textura angularis to textura prismatica, Munk pores present. Paraphyses absent. Asci 8-spored,
arising from a hyaline placenta-like tissue, radially extending into the locule around the
Quellkörper from the base, unitunicate, clavate, pedicellate, apex rounded, without apical ring,
evanescent. Ascospores bi-seriate to overlapping, hyaline to brown, ellipsoidal to oval, straight or
939
curved, aseptate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from
Subramanian & Sekar 1990).
Notes – Scortechinia was introduced by Saccardo & Berlese (1885) for S. culcitella. This
genus subsequently was considered as a synonym of Nitschkia based on similar morphology
(Nannfeldt & Santesson 1975). However, S. acanthostroma, S. conferta and S. diminuspora were
sequenced by Huhndorf et al. (2004b) and Mugambi & Huhndorf (2010). These strains are sister to
Tympanopsis in Scortechiniaceae (Fig. 1). Subramanian & Sekar (1990) combined Tympanopsis
confertula as Scortechinia conferta based on their similar characteristics, which is also confirmed
by molecular analysis (Fig. 1).
Scortechinia culcitella (Berk. & Ravenel) Speg., Anal. Soc. cient. argent. 26(1): 29 (1888)
Fig. 24a–m
Basionym – Sphaeria culcitella Berk. & Ravenel, in Berkeley & Curtis, Proc. Amer. Acad.
Arts & Sci. 4: 128 (1860)
Facesoffungi number: FoF 10170
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae 6.5–14 µm wide, with dichotomously branched, spinous ends. Ascomata
220–400 × 140–260 µm (x
̄ = 300 × 220 µm, n = 10), densely gregarious, superficial, sitting in a
subiculum, carbonaceous, black, with metallic iridescence, turbinate, globose to subglobose, the
apex collapsing when dry, glabrous, lacking ostioles, with a central, conical Quellkörper. Peridium
30–75 µm wide, composed of membranaceous, brown to hyaline cells of textura angularis to
textura prismatica, Munk pores present. Paraphyses absent. Asci 8-spored, arising from a hyaline
placenta-like tissue, radially extending into the locule around the Quellkörper from the base,
unitunicate, clavate, pedicellate, apex rounded, without apical ring, evanescent. Ascospores 9–13×
4–5 µm (x
̄ = 11 × 4.5 µm, n = 20), bi-seriate to overlapping, hyaline to brown, ellipsoidal to oval,
straight or curved, aseptate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from
Spegazzini 1888).
Material examined – USA, Carolina, on decayed trunk of Ostrya or Acer (PC-MNHN-PC-
PC0121050).
Known hosts and distribution – on decayed wood of Betula, Cerasus and Prunus in India,
Paraguay (type locality) and the USA (Spegazzini 1888, Subramanian & Sekar 1990).
Notes – Scortechinia culcitella was introduced based on Sphaeria culcitella (Spegazzini
1888). Nannfeldt & Santesson (1975) subsequently transferred it as Nitschkia acanthostroma (= S.
acanthostroma) based on morphology. Scortechinia acanthostroma was sequenced by Mugambi &
Huhndorf (2010), but there is no molecular data available for the type S. culcitella.
Scortechiniella Arx & E. Müll., Beitr. Kryptfl. Schweiz 11(no. 1): 382 (1954)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, rounded ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, glabrous, lacking ostioles, with a central, conical Quellkörper at
apex. Peridium composed of membranaceous, brown to hyaline cells of textura angularis to textura
prismatica. Paraphyses absent. Asci polysporous, arising from a hyaline placenta-like tissue,
radially extending into the locule around the Quellkörper from the base, unitunicate, clavate,
pedicellate, apex rounded, without apical ring, evanescent. Ascospores overlapping, hyaline,
ellipsoidal to oval, aseptate, smooth-walled, guttulate, with a filiform appendage at each end.
Asexual morph: Undetermined (adapted from von Arx & Müller 1954).
Notes – The monotypic Scortechiniella was introduced for S. similis which has similar
morphology to Scortechinia, differing in multi-spored asci and ascospores with appendages (von
Arx & Müller 1954). Mugambi & Huhndorf (2010) accepted it as a member of Scortechiniaceae
based on its molecular data. In this study, Scortechiniella is sister to Biciliospora (85%ML/1.00BY,
Fig. 1). However, the latter has fusiform ascospores.
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Figure 24 – Scortechinia culcitella: a–h, j–m (PC-MNHN-PC-PC0121050); i (redrawn from
Subramanian & Sekar 1990). a Material. b–d Gregarious ascomata sitting in a subiculum.
e Sectional view of Ascomata. f–h Hyphae of subiculum. i Asci. j–m Ascospores; Coronophorella
chaetomioides: n–o (redrawn from Subramanian & Sekar 1990). n Sectional view of Ascoma.
o Ascus. Scale bars: b–e = 200 µm, f, n = 100 µm, g–h = 50 µm, o = 20 µm, i = 10 µm, j–m = 5
µm.
Scortechiniella similis (Bres.) Arx & E. Müll., Beitr. Kryptfl. Schweiz 11(no. 1): 383 (1954)
Fig. 25l–n
Basionym – Trichosphaerella similis Bres., in Wildeman, Bull. Jard. bot. État Brux. 4(1): 7
(1914)
Facesoffungi number: FoF 10171
941
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, rounded ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, glabrous, lacking ostioles, with a central, conical Quellkörper at
apex. Peridium composed of membranaceous, brown to hyaline cells of textura angularis to textura
prismatica. Paraphyses absent. Asci polysporous, arising from a hyaline placenta-like tissue,
radially extending into the locule around the Quellkörper from the base, unitunicate, clavate,
pedicellate, apex rounded, without apical ring, evanescent. Ascospores overlapping, hyaline,
ellipsoidal to oval, aseptate, smooth-walled, guttulate, with a filiform appendage at each end.
Asexual morph: Undetermined (adapted from von Arx & Müller 1954).
Known hosts and distribution – on decayed wood in Puerto Rico and Zaire (type locality)
(von Arx & Müller 1954, Mugambi & Huhndorf 2010).
Notes – Scortechiniella similis is similar to Biciliospora velutina in having ascomata without
ostioles, polysporous asci and ascospores with filiform appendages, but the latter has fusiform
ascospores. Phylogenetically, S. similis is sister to B. velutina based on multi-gene analysis
(85%ML/1.00BY, Fig. 1).
Scortechiniellopsis Sivan., Trans. Br. mycol. Soc. 62(1): 36 (1974)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, tapering ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, glabrous, lacking ostioles, with a central, conical Quellkörper at
apex, almost or touching the bottom of the locule. Peridium composed of membranaceous, brown
to hyaline cells of textura angularis to textura prismatica. Paraphyses filiform. Asci polysporous,
arising from a hyaline placenta-like tissue, radially extending into the locule around the
Quellkörper from the base, unitunicate, clavate to cylindrical, pedicellate, apex rounded, without
apical ring, evanescent. Ascospores overlapping, hyaline, reniform, allantoid to ovoid, aseptate,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Sivanesan 1974).
Notes – The monotypic Scortechiniellopsis was initially placed in Coronophorales (Sivanesan
1974). However, Mugambi & Huhndorf (2010) accepted it in Scortechiniaceae based on
phylogenetic result. In this study, Scortechiniellopsis is sister to Biciliospora and Scortechiniella
(83%ML/1.00BY, Fig. 1), differing from it in the possessing of ascospores without appendage.
Scortechiniellopsis leonensis Sivan., Trans. Br. mycol. Soc. 62(1): 37 (1974) Fig. 25a–f
Facesoffungi number: FoF 10172
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae, with dichotomously branched, tapering ends. Ascomata scattered or
gregarious, superficial, sitting in a subiculum, coriaceous, brown to black, globose to subglobose,
the apex collapsing when dry, glabrous, lacking ostioles, with a central, conical Quellkörper at
apex, almost or touching the bottom of the locule. Peridium composed of membranaceous, brown
to hyaline cells of textura angularis to textura prismatica. Paraphyses filiform. Asci polysporous,
arising from a hyaline placenta-like tissue, radially extending into the locule around the
Quellkörper from the base, unitunicate, clavate to cylindrical, pedicellate, apex rounded, without
apical ring, evanescent. Ascospores overlapping, hyaline, reniform, allantoid to ovoid, aseptate,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Sivanesan 1974).
Material examined – Sierra Leone, on decayed branch of Cynometra leonensis, 27 September
1954, F.C. Deighton (IMI-58117, holotype).
Known hosts and distribution – on decayed wood of Cynometra leonensis (Fabaceae) in
Sierra Leone (type locality) and Kenya (Mugambi & Huhndorf 2010).
Notes – We re-examined the holotype specimen. Unfortunately, we did not find any
structures, except for some dark brown and septate hyphae.
942
Figure 25 – Scortechiniellopsis leonensis: a–b (IMI-58117, holotype); c–f (redrawn from
Sivanesan 1974). a Material. b Hyphae on host. c Sectional view of Ascoma. d Hyphae of
subiculum. e Ascus. f Ascospores; Cryptosphaerella nitschkei: g (IMI-K-M251354); h–i (NY-
02934200); j–k (redrawn from Saccardo 1882b). g–h Immersed ascomata. i Ascoma cross section.
943
j Ascus. k Ascospores; Scortechiniella similis: l–n (redrawn from von Arx & Müller 1954).
l Sectional view of ascoma. m Ascus. n Ascospores. Scale bars: g = 1 mm, h = 500 µm, b–c, i, l =
200 µm, d, j, m = 50 µm, e, k, n = 20 µm, f = 10 µm.
Tympanopsis Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 19(no. 2): 24 (1894)
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae. Ascomata scattered to gregarious, superficial, sitting in a subiculum,
carbonaceous to coriaceous, black, turbinate, globose to subglobose, the apex collapsing when dry,
tuberculate, glabrous, lacking ostioles, with a central, conical Quellkörper. Peridium usually outer
layer composed of carbonaceous, black tissue; inner layer composed of membranaceous, brown to
hyaline cells of textura angularis to textura prismatica, sometimes Munk pores present.
Paraphyses absent. Asci 8-spored, arising from a hyaline placenta-like tissue, radially extending
into the locule around the Quellkörper from the base, unitunicate, clavate, pedicellate, apex
rounded, without apical ring, evanescent. Ascospores bi-seriate to overlapping, hyaline to brown or
olivaceous brown, ellipsoidal to oval, straight or curved, aseptate, smooth-walled, guttulate.
Asexual morph: Undetermined (adapted from Starbäck 1894).
Notes – Tympanopsis was introduced by Starbäck (1894) for T. euomphala. Nannfeldt &
Santesson (1975) subsequently accepted it as Nitschkia. However, the phylogenetic position of T.
uniseriate is an independent clade in Scortechiniaceae (Mugambi & Huhndorf 2010). In this study,
Tympanopsis is sister to Scortechinia (Fig. 1).
Tympanopsis euomphala (Berk. & M.A. Curtis) Starbäck, Bih. K. svenska VetenskAkad. Handl.,
Afd. 3 19(no. 2): 24 (1894) Fig. 26
Basionym – Sphaeria euomphala Berk. & M.A. Curtis, Grevillea 4(no. 32): 141 (1876)
Facesoffungi number: FoF 10069
Saprobic on wood. Sexual morph: Subiculum composed of numerous, brown to dark brown,
branched, septate hyphae 4–8 µm wide, with rounded ends. Ascomata 390–700 × 400–600 µm (x
̄ =
580 × 480 µm, n = 10), scattered to gregarious, superficial, sitting in a subiculum, carbonaceous to
coriaceous, black, turbinate, globose to subglobose, the apex collapsing when dry, tuberculate,
glabrous, lacking ostioles, with a central, conical Quellkörper. Peridium 40–130 µm wide,
comprising three layers, outer layer composed of carbonaceous, black tissue; middle layer
composed of membranaceous, brown to reddish-brown cells of textura angularis, Munk pore
present; inner layer membranaceous, composed of hyaline cells of textura prismatica. Paraphyses
absent. Asci 20–25 × 8–12 µm (x
̄ = 23 × 10 µm, n = 10) at ascospore-bearing part, 8-spored, arising
from a hyaline placenta-like tissue, radially extending into the locule around the Quellkörper from
the base, unitunicate, clavate, pedicellate, apex rounded, without apical ring, evanescent.
Ascospores 6–9 × 2.5–3.5 µm (x
̄ = 7 × 3 µm, n = 50), bi-seriate to overlapping, hyaline to
olivaceous brown, ellipsoidal, straight or slightly curved, aseptate, smooth-walled, guttulate.
Asexual morph: Undetermined.
Material examined – UK, on decayed wood, associated with Hyporxylon rubiginosum, 26
September 1935, E.W. Mason (IMI-13384).
Known hosts and distribution – on decayed wood of Fraxinus in the UK and USA (type
locality) (Berkeley 1876).
Notes – Tympanopsis euomphala was transferred as Nitschkia confertula (Nannfeldt &
Santesson 1975), but no molecular data is available. We were unable to obtain type material.
Therefore, we re-examined an authentic material determined by Nannfeldt (mentioned in the label
of material).
Hypocreales Lindau, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1): 343 (1897)
Notes – Hypocreales accommodates fourteen families, Bionectriaceae, Calcarisporiaceae,
Clavicipitaceae, Cocoonihabitaceae, Cordycipitaceae, Flammocladiellaceae, Hypocreaceae,
Myrotheciomycetaceae, Nectriaceae, Niessliaceae, Ophiocordycipitaceae, Sarocladiaceae,
944
Stachybotryaceae, and Tilachlidiacea (Hyde et al. 2020e, Wijayawardene et al. 2020). They are
widely distributed in tropical and subtropical regions as saprophytic, symbiotic or parasitic
organisms (Maharachchikumbura et al. 2016, Hyde et al. 2020e). However, the multi-gene analysis
proved that these families belong in Hypocreales, and they usually have membranaceous ascomata
(Hyde et al. 2020e).
Figure 26 – Tympanopsis euomphala: a–l, n–p (IMI-13384); m (redrawn from Starbäck 1894).
a Material. b–c Ascomata. d–e Ascoma cross section. f Munk pores (arrow). g Hyphae of
subiculum. h–m Asci. n–p Ascospores. Scale bars: b = 1 mm, c = 500 µm, d = 200 µm, e, g = 100
µm, m = 20 µm, f, h–l = 10 µm, n–p = 5 µm.
Niessliaceae Kirschst., Annls mycol. 37(1/2): 89 (1939)
Saprobic or parasitic on wood, leaves or isolated from soil, sometimes fungicolous and
lichenicolous. Sexual morph: Ascomata perithecial or cleistothecial, solitary or gregarious, semi-
immersed to superficial, sometimes sitting on a subiculum, cupuliform, globose to subglobose,
yellow, orange, brown or black, membranaceous, tuberculate or smooth, glabrous or with setae or
irregular appendages, collapsing or collapsing laterally or not collapsing, with or lacking ostioles.
Peridium composed of membranaceous, yellow or brown or olivaceous brown or reddish brown to
945
hyaline cells of textura angularis to textura epidermoidea to textura prismatica. Paraphyses absent
or filiform or cylindrical, septate, branched or unbranched. Asci 8-spored, unitunicate, cylindrical to
clavate, pedicellate, apex rounded or blunt, apical ring indistinct or absent, evanescent. Ascospores
uni- or bi-seriate or overlapping, hyaline to brown, ellipsoidal to fusiform, straight or curved, 0–1-
septate, verrucose or smooth-walled, sometimes faintly striate, mostly with guttules. Asexual
morph: Hyphomycetous. Mycelium hyaline to light brown, superficial, effused. Conidiophores
mononematous, macronematous, hyaline, unbranched, aseptate. Conidia oval to ellipsoidal, usually
aseptate, hyaline, smooth-walled, guttulate. Coelomycetous. Conidiomata globose to subglobose,
dark brown to black. Peridium composed of brown cells of textura angularis, becoming hyaline
towards the inner conidiogenous region. Conidiophores hyaline, cylindrical to subcylindrical,
septate. Conidia oval to ellipsoidal, 0–2-septate, hyaline, smooth-walled, guttulate (adapted from
Samuels & Barr 1997, Hyde et al. 2020e).
Type genus – Niesslia Auersw.
Notes – The family Niessliaceae was established by Kirschstein (1939) to currently
accommodate 21 genera (Hyde et al. 2020e, Wijayawardene et al. 2020). They usually have
perithecial ascomata, unitunicate asci with inamyloid apical ring and hyaline, ellipsoidal to
fusiform ascospores (Samuels & Barr 1997, Hyde et al. 2020e). Eight genera have molecular data,
Eucasphaeria, Myrtacremonium, Neoeucasphaeria, Niesslia, Pseudohyaloseta, Rosasphaeria,
Trichosphaerella, Valetoniellopsis, and they are aggregated in Niessliaceae (Hyde et al. 2020e, this
study, 83%ML/1.00BY, Fig. 1).
Atronectria Etayo, in Etayo & Rosato, Biblthca Lichenol. 98: 52 (2008)
Lichenicolous. Sexual morph: Ascomata perithecial, solitary or scattered, immersed to
erumpent, pyriform to subglobose, black, membranaceous, tuberculate, glabrous, collapsing or
collapsing laterally when dry, ostiolate, periphysate. Peridium composed of membranaceous, brown
to hyaline cells of textura angularis to textura epidermoidea to textura prismatica. Paraphyses
cylindrical, branched, septate, constricted at the septum. Asci 8-spored, unitunicate, cylindrical,
pedicellate, apex blunt, apical ring indistinct, evanescent. Ascospores bi-seriate, hyaline to brown,
oblong to ellipsoidal, straight or slightly curved, 1-septate, smooth-walled, guttulate. Asexual
morph: Undetermined (adapted from Etayo & Sancho 2008, Flakus et al. 2019).
Notes – The lichenicolous Atronectria was introduced by Etayo & Sancho (2008) for A.
magellanica. The second species, A. lobariellae, was collected on Lobariella crenulata from
Bolivia (Flakus et al. 2019). These two species have perithecial ascomata, cylindrical asci and
hyaline ascospores (Etayo & Sancho 2008, Flakus et al. 2019), however no molecular data is
available for the Atronectria.
Atronectria magellanica Etayo, in Etayo & Sancho, Biblthca Lichenol. 98: 52 (2008) Fig. 27a–g
Facesoffungi number: FoF 10173
Lichenicolous. Sexual morph: Ascomata perithecial, solitary or scattered, immersed to
erumpent through the bark of host, pyriform to subglobose, black, membranaceous, tuberculate,
glabrous, collapsing or collapsing laterally when dry, ostiolate, periphysate. Peridium composed of
membranaceous, brown to hyaline cells of textura epidermoidea to textura prismatica. Paraphyses
cylindrical, branched, septate, constricted at the septum. Asci 8-spored, unitunicate, cylindrical,
pedicellate, apex blunt, apical ring indistinct, evanescent. Ascospores bi-seriate, hyaline to brown,
ellipsoidal to fusiform, straight or slightly curved, 1-septate, constricted at the septum, smooth-
walled, guttulate. Asexual morph: Undetermined (adapted from Etayo & Sancho 2008).
Material examined – Chile, Navarino, Lago Róbalo, S54º58’40.0”, W67º 41’24.7”, c. 420m,
on thallus of Pseudocyphellaria coriifolia on Nothofagus pumilio; 19 January 2005, J. Etayo
(MAF-Etayo 22938, holotype).
Known hosts and distribution – on Pseudocyphellaria coriifolia that grow on Nothofagus in
Chile (type locality) (Etayo & Sancho 2008).
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Notes – We re-examined the holotype of Atronectria magellanica, but apart from the fragile
ascomata, we did not observe other structures.
Circinoniesslia Samuels & M.E. Barr, Can. J. Bot. 75(12): 2166 (1998) [1997]
Fungicolous, on ascomata of Bionectria or Nectria species. Sexual morph: Ascomata
perithecial, solitary or scattered, superficial, globose, yellow, membranaceous, surrounded by
white, circinately coiled appendages, collapsing when dry, ostiolate with an unconspicuous papilla,
periphysate. Peridium composed of membranaceous, yellow to hyaline cells of textura globulosa to
textura angularis to textura prismatica. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical,
pedicellate, apex rounded, evanescent. Ascospores bi-seriate, hyaline, ellipsoidal, straight or
slightly curved, 1-septate, striate, guttulate. Asexual morph: Undetermined (adapted from Samuels
& Barr 1997).
Notes – The monotypic Circinoniesslia was found on ascomata of Bionectria or Nectria and
has a special crown-shaped appendage on ascomata with striate ascospores (Samuels & Barr 1997).
Samuels & Barr (1997) considered that the superficial and setose ascomata is the main feature of
Niessliaceae, and Circinoniesslia meets this condition.
Circinoniesslia nectriae Samuels & M.E. Barr, Can. J. Bot. 75(12): 2166 (1998) [1997] Fig. 28
Facesoffungi number: FoF 10070
Fungicolous, on ascomata of Bionectria or Nectria species. Sexual morph: Ascomata 260–
360 µm (x
̄ = 300 µm, n = 10) diam., perithecial, solitary or scattered, superficial, globose, yellow,
membranaceous, surrounded by white, circinately coiled appendages, collapsing when dry, ostiolate
with an unconspicuous papilla, periphysate. Peridium 25–65 µm wide, membranaceous, comprising
three layers, outer layer composed of hyaline cells of textura globulosa; middle layer composed of
yellow cells of textura angularis; inner layer composed of hyaline cells of textura prismatica.
Paraphyses absent. Asci (45–)50–60(–65) × 7–11 µm (x
̄ = 55 × 9 µm, n = 20), 8-spored,
unitunicate, cylindrical, pedicellate, apex rounded, evanescent. Ascospores (10–)12–13(–15) ×
(4.5–)5–6(–7) µm (x
̄ = 12.5 × 5.5 µm, n = 50), bi-seriate, hyaline, ellipsoidal to boat-shaped,
straight or slightly curved, 1-septate, striate, with an apical germ pore, guttulate. Asexual morph:
Undetermined.
Material examined – Puerto Rico, Caribbean National Forest, Luquillo Mts., El Verde
Research Area, alt. 350 m, on Bionectria sp. on decayed wood, 19 February 1996, G.J. Samuels,
H.-J. Schroers (BPI-748359, holotype).
Known hosts and distribution – on ascomata of Bionectria or Nectria species on decayed
wood in Guyana and Puerto Rico (type locality) (Samuels & Barr 1997).
Notes – We re-examined the holotype of Circinoniesslia nectriae and found that ascomata of
C. nectriae grow on the dried ascomata of Bionectria. Samuels & Barr (1997) mentioned ‘the
setose ascomata give the impression of small animals, the mouths of which are fringedd with
graceful whiskers’. We observed cross-sections of the appendages of ascomata and found that it is
composed of loose and globose cells that swell in water.
Cryptoniesslia Scheuer, Mycol. Res. 97(5): 543 (1993)
Saprobic on leaves. Sexual morph: Ascomata perithecial, solitary, immersed to erumpent,
pyriform to subglobose, brown or black, membranaceous, with short setae around the ostioles,
periphysate, collapsing when dry. Setae cylindrical, expanded near the apex, aseptate, branched or
unbranched, dark brown. Peridium composed of membranaceous, brown to hyaline cells of textura
epidermoidea to textura angularis to textura prismatica. Paraphyses cylindrical, unbranched,
septate, constricted at the septum. Asci 8-spored, unitunicate, clavate, pedicellate, apex blunt, apical
ring indistinct, evanescent. Ascospores bi-seriate or overlapping, hyaline, fusiform, with rounded
ends, straight or slightly curved, 1-septate, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Scheuer 1993).
Notes – The monotypic Cryptoniesslia was established by Scheuer (1993) for C. seutlosa,
which has similarly immersed ascomata with special setae to Niesslia and Trichosphaerella.
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Figure 27 – Atronectria magellanica: a–d (MAF-Etayo 22938, holotype); e–g (redrawn from Etayo
& Sancho 2008). a Material. b–d Ascomata on the host. e Sectional view of ascoma. f Ascus and
paraphyses. g Ascospores; Myrmaeciella endoleuca: h (HBG-2347); i–j (redrawn from Jaklitsch &
Voglmayr 2011). h Ascomata on the host. i Ascus. j Ascospores; Eucasphaeria capensis: k–o
(redrawn from Crous et al. 2007). k Ascus. l Ascospores. m Conidiophores. n–o Conidia. Scale
bars: b = 2 mm, h = 1 mm, c = 500 µm, d = 200 µm, e = 50 µm, f, i, k, m = 20 µm, j, l, n–o = 10
µm, g = 5 µm.
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Figure 28 – Circinoniesslia nectriae: (BPI-748359, holotype). a Material. b–d Ascomata on the
host. e Ascoma cross section. f Ostiole with periphyses. g Crown appendage cross section.
h Peridium. i–k Asci. l–o Ascospores. Notes: j–k stained in Melzer’s reagent. Scale bars: b = 1 mm,
c–d = 200 µm, e = 100 µm, h = 50 µm, f–g, i–k = 20 µm, l–o = 5 µm.
Cryptoniesslia setulosa Scheuer, Mycol. Res. 97(5): 543 (1993) Fig. 29
Facesoffungi number: FoF 10071
Saprobic on leaves. Sexual morph: Ascomata 100–200 µm (x
̄ = 155 µm, n = 10) diam.,
perithecial, solitary, immersed to erumpent, pyriform to subglobose, brown or black,
membranaceous, with short setae around the ostioles, periphysate, collapsing when dry. Setae
cylindrical, expanded near the apex, aseptate, branched or unbranched, dark brown. Peridium 15–
40 µm wide, membranaceous, comprising three layers, brown cells of textura epidermoidea on
surface view; middle layer composed of brown to olivaceous brown cells of textura angularis;
949
inner layer composed of hyaline cells of textura prismatica. Paraphyses cylindrical, unbranched,
septate, constricted at the septum. Asci (25–)30–35(–40) × (8–)10–13(–15) µm (x
̄ = 32 × 12 µm, n
= 20) at ascospore-bearing part, 8-spored, unitunicate, clavate, pedicellate, apex blunt, apical ring
indistinct, evanescent. Ascospores (15–)20–24 × 2.5–3(–4) µm (x
̄ = 23 × 2.8 µm, n = 30), bi-seriate
or overlapping, hyaline, fusiform, with rounded ends, straight or slightly curved, aseptate when
young, becoming 1-septate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted
from Scheuer 1993).
Material examined – UK, North Devon, Barnstaple, sand dune system of Braunton Burrows,
on decayed leaves of Carex arenariae, 5 August 1988, C. Scheuer (BPI-802277, isotype).
Known hosts and distribution – on decayed leaves of Carex arenariae (Cyperaceae) in the
UK (type locality) (Scheuer 1993).
Notes – Cryptoniesslia setulosa was introduced by Scheuer (1993) in having immersed
ascomata with special setae surrounded by ostioles, but sequence data are not available.
Eucasphaeria Crous, in Crous, Mohammed, Glen, Verkley & Groenewald, Fungal Diversity 25: 21
(2007)
Parasitic on leaves and leaf litters. Sexual morph: Ascomata perithecial, solitary or scattered,
semi-immersed to superficial, globose to subglobose, black, membranaceous, surrounded by
hyaline hyphae, ostiolate, periphysate. Peridium composed of membranaceous, brown cells of
textura angularis. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded or blunt, apical ring distinct. Ascospores bi-seriate, hyaline, ellipsoidal to fusiform, straight
or curved, 1-septate, verrucose or smooth-walled, sometimes faintly striate, mostly with guttules.
Asexual morph: Coelomycetous. Conidiomata semi-immersed, globose to subglobose, acervuloid,
dark brown to black. Peridium composed of brown cells of textura angularis, becoming hyaline
towards inner conidiogenous region. Conidiophores hyaline, cylindrical to subcylindrical, septate,
branched, giving rise to 1–2 conidiogenous cells. Conidiogenous cells fusiform to ellipsoidal,
hyaline, phialidic, straight to slightly curved. Conidia in natural environment, cylindrical to oblong,
slightly curved, 0–1-septate, hyaline, smooth-walled, guttulate; in culture sporulating, falcate, 0–2-
septate, hyaline, smooth-walled, guttulate (adapted from Crous et al. 2007, 2016).
Notes – Eucasphaeria was established by Crous et al. (2007), and the type E. capensis was
sequenced by Vu et al. (2019). Subsequently, Eucasphaeria rustici and E. proteae were introduced
based on morphology and phylogenetic results (Crous et al. 2016, 2020a). In this study,
Eucasphaeria is sister to Neocucasphaeria and Rosasphaeria (99%ML/1.00BY, Fig. 1).
Eucasphaeria capensis Crous, in Crous, Mohammed, Glen, Verkley & Groenewald, Fungal
Diversity 25: 21 (2007) Fig. 27k–o
Facesoffungi number: FoF 10174
Parasitic on leaves and leaf litters. Sexual morph: Ascomata perithecial, solitary or scattered,
semi-immersed to superficial, globose to subglobose, black, membranaceous, surrounded by
hyaline hyphae, ostiolate, periphysate. Peridium composed of membranaceous, brown cells of
textura angularis. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded or blunt, apical ring distinct. Ascospores bi-seriate, hyaline, ellipsoidal to fusiform, straight
or curved, 1-septate, verrucose or smooth-walled, sometimes faintly striate, mostly with guttules.
Asexual morph: Coelomycetous. Conidiomata semi-immersed, globose to subglobose, acervuloid,
dark brown to black. Peridium composed of brown cells of textura angularis, becoming hyaline
towards the inner conidiogenous region. Conidiophores hyaline, cylindrical to subcylindrical,
septate, branched, giving rise to 1–2 conidiogenous cells. Conidiogenous cells fusiform to
ellipsoidal, hyaline, phialidic, straight to slightly curved. Conidia in natural environment,
cylindrical to oblong, slightly curved, aseptate, hyaline, smooth-walled, guttulate; in culture
sporulating, falcate, 0–2-septate, hyaline, smooth-walled, guttulate (adapted from Crous et al.
2007).
950
Known hosts and distribution – on leaves and leaf litters of Eucalyptus in South Africa (type
locality) (Crous et al. 2007).
Notes – Eucasphaeria capensis was collected from South Africa and has black ascomata,
fuisform ascospores and coelomycetous asexual morph (Crous et al. 2007). Its type strain, E.
capensis (CBS 120028) was sequenced by Vu et al. (2019), and it is sister to E. rustici
(98%ML/0.94BY, Fig. 1).
Figure 29 – Cryptoniesslia setulosa: a–f, h–k, m–o (BPI-802277, isotype); g, l (redrawn from
Scheuer 1993). a Material. b Immersed ascomata. c Ascoma cross section. d, f Ostiole with
periphyses (arrow points to setae). e Peridium. g Setae. h Ascus with blunt and thick apex. i–k Asci.
l Paraphysis. m–o Ascospores. Notes: h stained in Melzer’s reagent. Scale bars: b = 200 µm, c =
100 µm, d = 50 µm, e–f, i–l = 20 µm, g–h, m–o = 10 µm.
951
Malmeomyces Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 32 (1899)
Saprobic on bamboo. Sexual morph: Ascomata perithecial, solitary to scattered, superficial,
subglobose, usually collapsing as cupuliform, yellow, membranaceous to coriaceous, surrounded by
erect, septate, unbranched, tapering setae with rounded end, ostioles inconspicuous, periphysate.
Peridium composed of membranaceous, yellow to hyaline cells of textura angularis to textura
prismatica. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex
blunt, apical ring distinct, evanescent. Ascospores bi-seriate or overlapping, hyaline, fusiform,
straight or slightly curved, 1–3-septate, smooth-walled, guttulate. Asexual morph: Undetermined
(adapted from Starbäck 1899).
Notes – The monotypic Malmeomyces was initially placed in Hypocreaceae (Starbäck 1899),
and subsequently accepted at Niessliaceae based on ascomata with erect setae (Lumbsch &
Huhndorf 2010, Maharachchikumbura et al. 2016, Hyde et al. 2020e, Wijayawardene et al. 2020).
Malmeomyces pulchellus Starbäck [as 'pulchella'], Bih. K. svenska VetenskAkad. Handl., Afd. 3
25(no. 1): 32 (1899) Fig. 30
Facesoffungi number: FoF 10072
Saprobic on bamboo. Sexual morph: Ascomata 130–250 µm (x
̄ = 180 µm, n = 10) diam.,
perithecial, solitary to scattered, superficial, subglobose, usually collapsing as cupuliform, yellow,
membranaceous to coriaceous, surrounded by erect, septate, unbranched, tapering setae, 6–10 µm
wide at base, with rounded end, ostioles inconspicuous, periphysate. Peridium 25–60 µm wide,
composed of membranaceous, yellow to hyaline cells of textura angularis to textura prismatica.
Paraphyses absent. Asci (30–)37–40(–45) × (8–)10–15 µm (x
̄ = 39 × 12 µm, n = 20) at ascospore-
bearing part, 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex blunt, apical ring
distinct, evanescent. Ascospores (10–)12–15(–18) × 2.5–4 µm (x
̄ = 14 × 3 µm, n = 50), bi-seriate or
overlapping, hyaline, fusiform, straight or slightly curved, aseptate when young, becoming 1-
septate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Starbäck 1899).
Material examined – Brazil, Rio Grande do Sul, col. Ijuhy, on decayed of Bambusa
eujusdam, 1 April 1893, A. Malme (S-F46453, holotype).
Known hosts and distribution – on decayed leaves of Bambusa eujusdam (Bambuseae) in
Brazil (type locality) (Starbäck 1899).
Notes – Malmeomyces pulchellus was collected on decayed leaves of bamboo in Brazil and
has setose ascomata, clavate asci and 1–3-septate ascospores (Starbäck 1899). In this study, we re-
examined the holotype and found that it has 0–1-septate ascospores, but no 2–3-septate ascospores.
Melchioria Penz. & Sacc., Malpighia 11(9-10): 399 (1897)
Saprobic or parasitic on leaves or stems. Sexual morph: Ascomata perithecial, densely
gregarious, superficial, globose to subglobose, black, carbonaceous to coriaceous, glabrous, not
collapsing, ostiolate, periphysate. Peridium composed of dark brown to hyaline cells of textura
angularis to textura prismatica. Paraphyses absent or filiform. Asci 8-spored, unitunicate, clavate,
pedicellate, apical ring indistinct, evanescent. Ascospores bi-seriate or overlapping, hyaline,
fusiform, straight or slightly curved, 1-septate, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Spegazzini 1884, Penzig & Saccardo 1897).
Notes – Melchioria was initially considered similar to Sphaeria with black ascomata and
hyaline, fusiform ascospores (Penzig & Saccardo 1897). This genus was subsequently listed in
Niessliaceae (Lumbsch & Huhndorf 2007). However, Melchioria does not have setose or hairy
ascomata similar to other Niessliaceae members, and there is no molecular data available for this
genus. We maintain Melchioria in Niessliaceae pending future work.
Melchioria leucomelaena Penz. & Sacc., Malpighia 11(9-10): 399 (1897) Fig. 31
Facesoffungi number: FoF 10073
Saprobic on stems. Sexual morph: Ascomata 290–450 µm (x
̄ = 380 n = 10) diam., perithecial,
densely gregarious, superficial, globose to subglobose, tuberculate, black, carbonaceous to
952
coriaceous, glabrous, not collapsing, ostiolate, periphysate. Peridium 45–70 µm wide, comprising
three layers, outer layer composed of carbonaceous, black tissue; middle layer composed of
membranaceous, dark brown cells of textura angularis; inner layer membranaceous, composed of
hyaline cells of textura prismatica. Paraphyses absent. Asci 100–120(–140) × (20–)24–26(–30) µm
(x
̄ = 115 × 25 µm, n = 20) at ascospore-bearing part, 8-spored, unitunicate, clavate, pedicellate,
apical ring indistinct, evanescent. Ascospores (45–)52–60(–70) × (6–)8–10(–14) µm (x
̄ = 57 × 9.5
µm, n = 50), bi-seriate or overlapping, hyaline, fusiform, straight or slightly curved, 1-septate,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Penzig & Saccardo 1897).
Material examined – Indonesia, Jawa, Jawa Barat (West), Tjibodas, on decayed stems of
Elettaria, N. Nyman (S-F74326); ibid. on stems of Amomum, 1907, v. Höhnel (S-F82127).
Known hosts and distribution – on decayed stems of Elettaria in Indonesia (type locality)
(Penzig & Saccardo 1897).
Figure 30 – Malmeomyces pulchellus: (S-F46453, holotype). a Material. b–c Ascomata on the host.
d Ascoma cross section. e Squashed ostiole with periphyses. f Septate setae. g Peridium. h–k Asci.
l–m Ascospores. Notes: m stained in cotton blue reagent. Scale bars: b–c = 500 µm, d = 100 µm, f–
g = 50 µm, e, h–k = 20 µm, l–m = 10 µm.
Notes – Melchioria leucomelaena was found on the rotten stems of Elettaria in Indonesia and
is characterized by black ascomata with fusiform ascospores (Penzig & Saccardo 1897) but no
953
molecular data is available for this species. In this study, the type material is unable to obtain.
Therefore, we re-examined two authentic materials collected from Indonesia by Nyman and Höhnel
respectively.
Figure 31 – Melchioria leucomelaena: a–b, d, f–n (S-F74326); c, e (S-F82127). a Material.
b–c Ascomata on the host. d Ascomata. e–f Peridium. g–h Evanescent asci. i–n Ascospores. Scale
bars: b–c = 1 mm, d = 500 µm, e–h = 50 µm, i–n = 20 µm.
Miyakeomyces Hara, Bot. Mag., Tokyo 27(no. 317): (248) (1913)
Parasitic on leaves. Sexual morph: Ascomata perithecial, densely gregarious, penetrated,
globose to subglobose, black, coriaceous to membranaceous, glabrous, collapsing when dry,
ostiolate, periphysate. Peridium composed of membranaceous, olivaceous brown to hyaline cells of
textura intricata to textura angularis to textura prismatica. Paraphyses absent. Asci 8-spored,
unitunicate, cylindrical to clavate, pedicellate, apex rounded, apical ring indistinct, evanescent.
Ascospores bi-seriate, hyaline, ellipsoidal to fusiform, straight or slightly curved, 2–4-septate,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from von Höhnel 1918).
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Notes – The monotypic Miyakeomyces was found to be parasitic on leaves of Phyllostachys
(Hara 1913). von Höhnel (1918) observed that M. bambusae makes the leaf yellow and thick, and
its fruiting bodies appear on both sides of the leaf.
Miyakeomyces bambusae Hara [as ‘Miyakeamyces’], Bot. Mag., Tokyo 27(no. 317): (248) (1913)
Fig. 32
Facesoffungi number: FoF 10074
Parasitic on leaves. Sexual morph: Ascomata densely aggregate into an ellipsoid, gradually
thinning towards the edge, 450–520 × 500–650 µm (x
̄ = 480 × 570 µm, n = 5) diam., perithecial,
penetrated, globose to subglobose, 100–150 µm (x
̄ = 120 µm, n = 10) diam., black, coriaceous to
membranaceous, glabrous, collapsing when dry, ostiolate, periphysate. Peridium 16–40 µm wide,
membranaceous, comprising three layers, outer layer composed of olivaceous brown cells of
textura intricata; middle layer composed of olivaceous brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Paraphyses absent. Asci (50–)60–70 × 11–13 µm
(x
̄ = 65 × 12 µm, n = 10), 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded,
apical ring indistinct, evanescent. Ascospores (20–)23–26(–30) × (4–)5–6(–7) µm (x
̄ = 25 × 5 µm, n
= 20), bi-seriate, hyaline, ellipsoidal to fusiform, straight or slightly curved, aseptate, smooth-
walled, guttulate. Asexual morph: Undetermined.
Material examined – Japan, Gifu, prov. Mino, Kawauye-mura, on leaves of Phyllostachys
bambusoides, 11 November 1911, K. Hara (S-F10914, Isolectotype).
Known hosts and distribution – on leaves of Phyllostachys bambusoides (Poaceae) in Japan
(type locality) (Hara 1913).
Notes – Miyakeomyces bambusae was introduced as a parasitic species on bamboo leaves and
in having 2–4-septate, fusiform ascospores (von Höhnel 1918). In this study, we only found
aseptate ascospores in the isolectotype material.
Myrmaeciella Lindau, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1(1): 478 (1897)
Saprobic on wood. Sexual morph: Ascomata perithecial, densely gregarious, superficial,
globose to subglobose, black, membranaceous, glabrous, ostioles inconspicuous. Peridium
composed of dark brown cells of textura angularis. Paraphyses absent. Asci 8-spored, unitunicate,
clavate, pedicellate or sessile, evanescent. Ascospores uni- to bi-seriate, hyaline, ellipsoidal to
oblong, straight or slightly curved, 1-septate, guttulate. Asexual morph: Undetermined (adapted
from Saccardo 1880, Theissen 1910, Jaklitsch & Voglmayr 2011).
Notes – Myrmaeciella was treated as a synonym of Hypocreopsis (Müller & von Arx 1962,
Rogerson 1970). Rossman et al. (1999) subsequently transferred Myrmaeciella to Niessliaceae
based on their similar membranaceous ascomata, inamyloid asci and ellipsoidal ascospores.
Myrmaeciella endoleuca (Sacc.) Lindau, in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig)
1(1): 478 (1897) Fig. 27h–j
Basionym – Myrmaecium endoleucum Sacc., Michelia 2(no. 6): 138 (1880)
Facesoffungi number: FoF 10175
Saprobic on wood. Sexual morph: Ascomata perithecial, densely gregarious, superficial,
globose to subglobose, black, membranaceous, glabrous, ostioles inconspicuous. Peridium
composed of dark brown cells of textura angularis. Paraphyses absent. Asci 8-spored, unitunicate,
clavate, pedicellate, evanescent. Ascospores uni- to bi-seriate, hyaline, ellipsoidal, 1-septate,
guttulate. Asexual morph: Undetermined (adapted from Saccardo 1880, Jaklitsch & Voglmayr
2011).
Material examined – Switzerland, on decayed wood, 11 February 1962, E. Müller and J.A.
von Arx (HBG-2347).
Known hosts and distribution – on decayed wood in Switzerland and the USA (type locality)
(Saccardo 1880).
955
Notes – The type material of Myrmaeciella endoleuca could not be obtained. Therefore, we
re-examined the authentic materials collected by Müller and Arx. A kind of hyphomycetes with
globose conidia contaminated this specimen, and except for the shriveled ascomata, no other
features were observed in this study.
Figure 32 – Miyakeomyces bambusae: (S-F10914, Isolectotype). a Material. b–c Ascomata on the
host (b on front of the leaf, c on back of the leaf). d–e Ascomata cross section. f Ascoma.
g Peridium. h–i Olivaceous brown cells of textura intricata. j Squashed ostiole. k–p Asci. Scale
bars: b–c = 500 µm, d = 200 µm, e = 100 µm, f, h–i = 50 µm, g, j–p = 20 µm.
Myrtacremonium Crous, in Crous et al., Persoonia 38: 259 (2017)
Parasitic on leaves. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium hyaline, septate, branched hyphae. Conidiophores mononematous, macronematous,
hyaline, straight to flexuous. Conidiogenous cells terminal, hyaline, subcylindrical, with minute
flared collarette at apex. Conidia solitary, oblong, aseptate, hyaline, smooth-walled, guttulate
(adapted from Crous et al. 2017).
956
Notes – The hyphomycetous genus Myrtacremonium was accepted in Niessliaceae based on
phylogenetic result (Crous et al. 2017). In this study, Myrtacremonium nests in Niessliaceae
(70%ML/0.99BY, Fig. 1).
Myrtacremonium eucalypti Crous, in Crous et al., Persoonia 38: 259 (2017) Fig. 33m
Facesoffungi number: FoF 10176
Parasitic on leaves. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium hyaline, septate, branched hyphae. Conidiophores mononematous, macronematous,
hyaline, straight to flexuous. Conidiogenous cells terminal, hyaline, subcylindrical, with minute
flared collarette at apex. Conidia solitary, oblong, aseptate, hyaline, smooth-walled, guttulate
(adapted from Crous et al. 2017).
Known hosts and distribution – on leaves of Eucalyptus globulus (Myrtaceae) in Australia
(type locality) (Crous et al. 2017).
Notes – The monotypic Myrtacremonium was introduced by Crous et al. (2017) for M.
eucalypti.
Neoeucasphaeria Crous, in Crous et al., Persoonia 41: 293 (2018)
Parasitic on Eucalyptus. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium hyaline, septate, branched hyphae. Conidiophores mononematous, macronematous,
densely gregarious, hyaline, straight to flexuous, branched. Conidiogenous cells terminal, hyaline,
subcylindrical, phialidic, flexuous. Conidia solitary, ellipsoidal, aseptate, hyaline, smooth-walled,
guttulate (adapted from Crous et al. 2018).
Notes – The monotypic Neoeucasphaeria was collected from Eucalyptus in Australia (Crous
et al. 2018). In this study, Neoeucasphaeria is sister to Rosasphaeria in Niessliaceae (Fig. 1).
Rosasphaeria has cylindrical, curved, 1-septate conidia different from Neoeucasphaeria (Jaklitsch
& Voglmayr 2012).
Neoeucasphaeria eucalypti Crous, in Crous et al., Persoonia 41: 293 (2018)
Facesoffungi number: FoF 10177
Parasitic on Eucalyptus. Sexual morph: Undetermined. Asexual morph: Hyphomycetous.
Mycelium hyaline, septate, branched hyphae. Conidiophores mononematous, macronematous,
densely gregarious, hyaline, straight to flexuous, branched. Conidiogenous cells terminal, hyaline,
subcylindrical, phialidic, flexuous. Conidia solitary, ellipsoidal, aseptate, hyaline, smooth-walled,
guttulate (adapted from Crous et al. 2018).
Known hosts and distribution – on Eucalyptus in Australia (type locality) (Crous et al. 2018).
Notes – The hyphomycetous Neoeucasphaeria eucalypti was introduced by Crous et al.
(2018) from Australia. It is sister to Rosasphaeria moravica (Fig. 1), differing from it in the
possessing of ellipsoidal conidia (Jaklitsch & Voglmayr 2012).
Niesslia Auersw., in Gonnermann & Rabenhorst, Myc. Europ. Pyren. 5-6: 30 (1869)
Saprobic on wood, leaves or stems. Sexual morph: Ascomata perithecial, solitary or
gregarious, superficial, sometimes sitting on a subiculum, globose to subglobose, or collapsing as
cupuliform, brown or black, membranaceous, surrounded by brown to black setae, ostioles
inconspicuous, periphysate. Peridium composed of membranaceous, yellow or brown to hyaline
cells of textura angularis to textura prismatica. Paraphyses absent. Asci 8-spored, unitunicate,
cylindrical to clavate, pedicellate, apex rounded or blunt, apical ring indistinct or absent,
evanescent. Ascospores uni- or bi-seriate or overlapping, hyaline, ellipsoidal, cylindrical to
fusiform, straight or curved, usually 1-septate, smooth-walled, guttulate. Asexual morph:
Hyphomycetous. Conidiophores mononematous, macronematous, hyaline, unbranched. Conidia
formed in chains or oval to ellipsoidal, 0–1-septate, hyaline, smooth-walled, guttulate (adapted
from Gams et al. 2019).
957
Notes – Niesslia was introduced by Auerswald (1869) for N. chaetomium and it is
characterized by spiny ascomata, ellipsoidal ascospores and acremonium-like asexual morph
(Gams et al. 2019). Gams et al. (2019) clarified the affinities of Niesslia species by reviewing the
authentic specimens. Several Niesslia species were sequenced (Crous et al. 2019a, Gams et al.
2019, Vu et al. 2019), but there is no molecular data available about the type N. chaetomium. In this
study, species of Niesslia are scattered in Niessliaceae (Fig. 1).
Niesslia chaetomium (Ces. & De Not.) Auersw., in Gonnermann & Rabenhorst, Myc. Europ.
Pyren. 5-6: 30 (1869) Fig. 33j–l
Basionym – Venturia chaetomium Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc. 4): 225
(1863)
Facesoffungi number: FoF 01128
Saprobic on coniferous substrates. Sexual morph: Ascomata perithecial, solitary or scattered,
superficial, globose to subglobose, or collapsing as cupuliform, black, membranaceous, ostiolate,
surrounded by long, brown to black, conical, aseptate, tapering setae. Peridium composed of
membranaceous, brown cells of textura angularis. Paraphyses absent. Asci 8-spored, unitunicate,
cylindrical, pedicellate, apex blunt, evanescent. Ascospores uni- or bi-seriate, hyaline, ellipsoidal to
fusiform, straight or slightly curved, usually 1-septate, smooth-walled, guttulate. Asexual morph:
Undetermined (adapted from Gams et al. 2019).
Known hosts and distribution – on decayed wood of Pinus and Juniperus in Czech Republic
(type locality), France, Germany, the UK and Venezuela (Corda 1838, Gams et al. 2019).
Notes – Niesslia was established based on N. chaetomium (Auerswald 1869), and afterwards,
this species was treated as a synonym of N. exosporioides (Index Fungorum 2021). Gams et al.
(2019) examined specimens related to Niesslia, and proposed that the substrates of N. chaetomium
and N. exosporioides are different. Niesslia chaetomium was considered a synonym of N. pusilla
(Gams et al. 2019), but no molecular data available for these two species.
Niesslia exosporioides (Desm.) G. Winter, Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 197 (1885)
Fig. 33a–i
Basionym – Sphaeria exosporioides Desm., Nouv. Not.: 39 (1843)
Facesoffungi number: FoF 10178
Saprobic on grasses and similar substrates. Sexual morph: Ascomata 120–210 µm (x
̄ = 170
µm, n = 5), perithecial, solitary or scattered, superficial, globose to subglobose, or collapsing as
cupuliform, black, membranaceous, surrounded by long, brown to black, filiform, septate, tapering
setae, 4–9 µm wide at the base, ostioles inconspicuous, periphysate. Peridium 15–30 µm wide,
composed of membranaceous, brown cells of textura angularis. Paraphyses absent. Asci 8-spored,
unitunicate, cylindrical, pedicellate, apex blunt, evanescent. Ascospores 9–12 × 1–1.6 µm (x
̄ = 10 ×
1.2 µm, n = 20), uni- or bi-seriate, hyaline, ellipsoidal to fusiform, straight or slightly curved,
usually 1-septate, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Conidiophores
mononematous, macronematous, hyaline, unbranched. Conidia cylindrical, hyaline, smooth-walled,
guttulate (adapted from Gams et al. 2019).
Material examined – Austria, Niederdonau, Hainburg, Hundsheimer Kogel, on dead stems of
Carex pilosa, May 1940, F. Petrak (PRM-717659).
Known hosts and distribution – on decayed leaves or stems of Carex, Luzula and Festuca in
Argentina, Austria, Belgium and France (type locality) (Gams et al. 2019).
Notes – Niesslia exosporioides was introduced by Winter (1885) based on Sphaeria
exosporioides. It was considered as a synonym of N. chaetomium (Index Fungorum 2021).
However, Gams et al. (2019) disagreed with this statement. In this study, we re-examined authentic
material of N. exosporioides collected by Petrak and found that it has ascomata with filiform,
septate setae, differing from N. chaetomium that has ascomata with erect, spinous, aseptate setae
(Corda 1838, Gams et al. 2019).
958
Figure 33 – Niesslia exosporioides: a–i (PRM-717659). a Material. b–c Ascomata on the host.
d Peridium. e Septate seta. f–i Ascospores; Niesslia chaetomium: j–l (redrawn from Corda 1838).
j Ascoma. k ascus. l Ascospores; Myrtacremonium eucalypti: m (redrawn from Crous et al. 2017).
m Conidiophores and conidia; Paraniesslia tuberculata: n–o (redrawn from Tsui et al. 2001).
n Ascus. o Ascospores. Scale bars: b–c = 500 µm, n = 50 µm, d, m = 20 µm, e, o = 10 µm, f–i = 5
µm.
Niesslia nolinae (A.W. Ramaley) W. Gams, in Gams, Stielow, Gräfenhan & Schroers, Mycol.
Progr. 18(1-2): 34 (2019) Fig. 34
Basionym – Hyaloseta nolinae A.W. Ramaley, Mycotaxon 79: 269 (2001)
Facesoffungi number: FoF 10179
Saprobic on leaves or leaf litter. Sexual morph: Ascomata perithecial, solitary or gregarious,
superficial, sitting on a subiculum with hyaline hyphae, globose to subglobose, collapsing when
dry, brown, membranaceous, surrounded by setae, ostioles inconspicuous, periphysate. Setae
cylindrical, aseptate, hyaline, tapering, with swollen apex. Peridium composed of membranaceous,
brown cells of textura epidermoidea. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical,
959
pedicellate, apex rounded, evanescent. Ascospores bi-seriate, hyaline, ellipsoidal to oblong, 1-
septate, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Mycelium white to orange,
superficial, effuse. Conidiophores mononematous, macronematous, hyaline, unbranched, aseptate.
Conidia (2–)3.5–4(–4.5) × 1–2 µm (x
̄ = 3.8 × 1.5 µm, n = 50), terminal, oval to ellipsoidal,
aseptate, hyaline, smooth-walled, guttulate (adapted from Ramaley 2001).
Material examined – USA, New Mexico, Lincoln Co., Valley of Fires, on leaf litter of Nolina
micrantha, 22 October 1997, A.W. Ramaley (BPI-871322, holotype).
Known hosts and distribution – on leaves of Nolina micrantha (Asparagaceae) in the USA
(type locality) (Ramaley 2001).
Notes – Niesslia nolinae initially introduced as Hyaloseta nolinae from the USA (Ramaley
2001). Gams et al. (2019) subsequently accepted this species in Niesslia based on its ascomata with
hyaline setae similar to N. pandani. Niesslia nolinae was sequenced by Lombard et al. (2015) and
Vu et al. (2019). In this study, N. nolinae is sister to N. gamsii (97%ML/1.00BY, Fig. 1), differing
from the latter that has oblong, clavate or ampulliform conidia (Ashrafi et al. 2017).
Nitschkiopsis Nannf. & R. Sant., in Nannfeldt, Svensk bot. Tidskr. 69: 322 (1975)
Lichenicolous. Sexual morph: Ascomata perithecial, solitary or scattered, semi-immersed to
superficial, globose to subglobose, or collapsing as cupuliform, brown to black, with metallic
iridescence, membranaceous, surrounded by two types of setae, ostiolate, periphysate. Setae oblong
to cylindrical, hyaline, with swollen apex; conical, spinous, brown to dark brown. Peridium
composed of membranaceous, pale brown to reddish brown to hyaline cells of textura angularis to
textura prismatica. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex
blunt, apical ring indistinct, evanescent. Ascospores uni- or bi-seriate, hyaline, ellipsoidal to oval,
0–1-septate, smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Nannfeldt &
Santesson 1975).
Notes – The monotypic Nitschkiopsis was introduced by Nannfeldt & Santesson (1975) for N.
stictarum. This lichenicolous genus was initially related to Lasiosphaeriaceae and Nitschkiaceae
(Nannfeldt & Santesson 1975). However, Tretiach (2002) transferred it as Niesslia stictarum and
introduced another similar lichenicolous species, Nie. robusta. In this study, we found that
Nitschkiopsis has perithecial ascomata with hyaline setae similar to Nie. nolinae; and spinous setae
similar to Nie. chaetomium, differing from it in the possessing of ellipsoidal ascospores. Since
Niesslia is polyphyletic (Fig. 1), we accept the Nitschkiopsis with two kinds of setae as an
independent genus in Niessliaceae pending molecular data.
Nitschkiopsis stictarum Nannf. & R. Sant., Svensk bot. Tidskr. 69(3): 322 (1975) Fig. 35
Facesoffungi number: FoF 10075
Lichenicolous. Sexual morph: Ascomata 70–130 µm (x
̄ = 105 µm, n = 10) diam., perithecial,
solitary or scattered, semi-immersed to superficial, globose to subglobose, or collapsing as
cupuliform, brown to black, with metallic iridescence, membranaceous, surrounded by two types of
setae, ostiolate, periphysate. Setae oblong to cylindrical, hyaline, with swollen apex, 45–85 × 3.5–5
µm; conical, spinous, brown to dark brown, 9–25 × 3–12 µm. Peridium 10–20 µm wide,
membranaceous, comprising three layers, outer layer composed of pale brown cells of textura
angularis; middle layer composed of reddish brown to brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Paraphyses absent. Asci (20–)25–30(–40) × (4–
)5–6.5 µm (x
̄ = 28 × 5.4 µm, n = 20) at ascospore-bearing part, 8-spored, unitunicate, cylindrical,
pedicellate, apex blunt, apical ring indistinct, evanescent. Ascospores (5–)6–8(–8.5) × (1.5–)2–
2.5(–3) µm (x
̄ = 7.2 × 2.3 µm, n = 50), uni- or bi-seriate, hyaline, ellipsoidal to oval, 0–1-septate,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Nannfeldt & Santesson
1975).
Material examined – Kenya, Central, Nanyku. Mt. Kenys, W slope, National Park Road
(Naro Moru Track), in the bamboo zone, Alt. 2700–3100 m, on Sticta ambavillaria on bamboo
(Arundinaria alpina), 23 January 1970, R. Santesson (UPS-UPS:BOT:L-112114, holotype); ibid.
960
(S-F10356, Isotype); ibid. (UPS-UPS:BOT:F-521855); Tanzania, Arusha, Mt. Meru, E slope, c. 2
km N of Kitoto Camp, on Hypericum lanceolatum in a high and relatively dense ericaceous
bushland (in the montane forest belt), on Sticta ambavillaria, 7 January 1971, R. Santesson (UPS-
UPS:BOT:F-521848).
Figure 34 – Niesslia nolinae: a, f–k (BPI-871322, holotype); b–e, l (redrawn from Ramaley 2001).
a Material. b Ascoma. c Seta with swollen apex. d Ascus. e Ascospores. f Dry culture. g Mycelium.
h, k–l Conidiophores and conidia. i Conidia. j Sporodochium. Scale bars: g = 500 µm, j = 200 µm,
b = 100 µm, c–e, h–i, k–l = 10 µm.
961
Figure 35 – Nitschkiopsis stictarum: a, c, e, g–i, k–m, o–p, u (UPS-UPS:BOT:L-112114,
holotype); b, d, f, j, n, r–t, w–z (S-F10356, Isotype); v (UPS-UPS:BOT:F-521855); q (UPS-
UPS:BOT:F-521848). a–b Material. c–d Ascomata on Sticta ambavillaria. e–f Ascomata.
g–h Ascoma cross section. i Ostiole with periphyses. j–k Peridium. l Conical seta. m–n Cylindrical
setae. o–t Asci. u–z Ascospores. Notes: n, w–z stained in Congo red reagent. Scale bars: c–d = 100
µm, g–h = 50 µm, i, m–n = 20 µm, j–l, o–t = 10 µm, u–z = 5 µm.
Known hosts and distribution – on Sticta ambavillaria in Kenya (type locality), Tanzania,
Peru, New Guinea (Nannfeldt & Santesson 1975, Gams et al. 2019).
Notes – Nitschkiopsis stictarum recorded so far found on Sticta ambavillaria (Nannfeldt &
Santesson 1975, Gams et al. 2019). It was transferred as Niesslia stictarum (Tretiach 2002),
however no molecular data is available for the N. stictarum.
962
Paraniesslia C.K.M. Tsui, K.D. Hyde & Hodgkiss, Mycologia 93(5): 1002 (2001)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary or scattered, semi-immersed
to superficial, pyriform, globose to subglobose, pale brown to brown, membranaceous, ostioles
surrounded by cylindrical, brown to black, septate setae, periphysate. Peridium membranaceous,
brown cells of textura epidermoidea on surface view; inner layer composed of brown to hyaline
cells of textura angularis to textura prismatica. Paraphyses filiform, septate, evanescent. Asci 8-
spored, unitunicate, clavate, pedicellate, apex blunt, evanescent. Ascospores uni- or bi-seriate or
overlapping, hyaline when young, becoming olivaceous brown, ellipsoidal to fusiform, 1-septate,
slightly constricted at the septum, tuberculate, guttulate. Asexual morph: Undetermined (adapted
from Tsui et al. 2001).
Notes – The monotypic Paraniesslia was introduced by Tsui et al. (2001) for P. tuberculata.
It is similar to Niesslia with setose ascomata and clavate asci, differing from it in the possessing of
tuberculate, ellipsoidal ascospores (Tsui et al. 2001). There is no molecular data available for
Paraniesslia.
Paraniesslia tuberculata C.K.M. Tsui, K.D. Hyde & Hodgkiss, Mycologia 93(5): 1002 (2001)
Fig. 33n–o
Facesoffungi number: FoF 10180
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary or scattered, semi-immersed
to superficial, pyriform, globose to subglobose, pale brown to brown, membranaceous, ostioles
surrounded by cylindrical, brown to black, septate setae, periphysate. Peridium membranaceous,
brown cells of textura epidermoidea on surface view; inner layer composed of brown to hyaline
cells of textura angularis to textura prismatica. Paraphyses filiform, septate, evanescent. Asci 8-
spored, unitunicate, clavate, pedicellate, apex blunt, evanescent. Ascospores uni- or bi-seriate or
overlapping, hyaline when young, becoming olivaceous brown, ellipsoidal to fusiform, 1-septate,
slightly constricted at the septum, tuberculate, guttulate. Asexual morph: Undetermined (adapted
from Tsui et al. 2001).
Known hosts and distribution – on submerged wood in China (type locality) (Tsui et al.
2001).
Notes – Paraniesslia tuberculate was collected on submerged wood in China (Tsui et al.
2001). It is similar to Niesslia, but can be differentiated from its olivaceous brown, tuberculate and
ellipsoidal ascospores (Tsui et al. 2001).
Pseudohyaloseta Tibpromma & K.D. Hyde, in Tibpromma et al., Fungal Diversity:
10.1007/s13225-018-0408-6, [113] (2018)
Saprobic on wood or leaves. Sexual morph: Ascomata perithecial, scattered or gregarious,
semi-immersed to superficial, globose to subglobose, black, membranaceous, surrounded by
cylindrical, hyaline setae with swollen apex, ostiolate, periphysate. Peridium membranaceous,
composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses filiform,
septate, unbranched. Asci 6–8-spored, unitunicate, clavate, pedicellate, apical ring distinct.
Ascospores uni- or bi-seriate, hyaline to subhyaline, cylindrical to fusiform, slightly curved, 1-
septate, slightly constricted at the septum, tuberculate, guttulate, with a mucilaginous sheath.
Asexual morph: Undetermined (adapted from Tibpromma et al. 2018).
Notes – The monotypic Pseudohyaloseta was established by Tibpromma et al. (2018) for P.
pandanicola. This genus is closely related to Niesslia nolinae based on phylogenetic result.
However, the latter has different ellipsoidal ascospores without a sheath (Tibpromma et al. 2018).
In this study, Pseudohyaloseta is sister to the clade of N. antarctica, N. gamsii (asexual morph) and
N. nolinae (100%ML/1.00BY, Fig. 1), which have ellipsoidal ascospores without a sheath.
Pseudohyaloseta pandanicola Tibpromma & K.D. Hyde, in Tibpromma et al., Fungal Diversity:
10.1007/s13225-018-0408-6, [113] (2018)
Facesoffungi number: FoF 10181
963
Saprobic on wood or leaves. Sexual morph: Ascomata perithecial, scattered or gregarious,
semi-immersed to superficial, globose to subglobose, black, membranaceous, surrounded by
cylindrical, hyaline setae with swollen apex, ostiolate, periphysate. Peridium membranaceous,
composed of brown to hyaline cells of textura angularis to textura prismatica. Paraphyses filiform,
septate, unbranched. Asci 6–8-spored, unitunicate, clavate, pedicellate, apical ring distinct.
Ascospores uni- or bi-seriate, hyaline to subhyaline, cylindrical to fusiform, slightly curved, 1-
septate, slightly constricted at the septum, tuberculate, guttulate, with a mucilaginous sheath.
Asexual morph: Undetermined (adapted from Tibpromma et al. 2018).
Known hosts and distribution – on dead roots, stems or leaves of Freycinetia and Pandanus
in Thailand (type locality) (Tibpromma et al. 2018).
Notes – Pseudohyaloseta pandanicola was first collected on dead roots of Freycinetia
javanica, and Pandanus sp. in Thailand (Tibpromma et al. 2018).
Pseudonectriella Petr., Sydowia 13(1-6): 127 (1959)
Saprobic or parasitic on wood. Sexual morph: Ascomata perithecial, solitary, superficial,
turbinate, obpyriform to subglobose, or deep collapsing as cupuliform, 3–8-loculate, dark brown to
black, membranaceous, tuberculate, glabrous, ostioles inconspicuous, periphysate. Peridium
composed of membranaceous, brown to pale brown to hyaline cells of textura globulosa to textura
angularis to textura prismatica. Paraphyses rare, filiform, septate. Asci 8-spored, unitunicate,
cylindrical, pedicellate, apex rounded, apical ring absent, evanescent. Ascospores uni- or bi-seriate,
hyaline, ellipsoidal to oval, 0–1-septate, smooth-walled, guttulate. Asexual morph: Coelomycetous.
Conidiomata superficial, turbinate, obpyriform to subglobose, or deep collapsing as cupuliform, 3–
8-loculate, dark brown to black, membranaceous, tuberculate. Peridium composed of
membranaceous, brown to pale brown cells of textura globulosa to textura angularis to textura
prismatica, becoming hyaline towards inner conidiogenous region. Conidiophores erect, hyaline,
cylindrical to subcylindrical, unbranched. Conidia terminal, oval to ellipsoidal, aseptate, hyaline,
smooth-walled, guttulate (adapted from Petrak 1959).
Notes – The monotypic Pseudonectriella is typified by P. ahmadii which has black ascomata,
cylindrical asci and ellipsoidal ascospores (Petrak 1959). Lumbsch & Huhndorf (2007) placed this
genus in Niessliaceae. However, no sequence data is available for the Pseudonectriella.
Pseudonectriella ahmadii Petr., Sydowia 13(1-6): 128 (1959) Fig. 36
Facesoffungi number: FoF 10076
Saprobic or parasitic on wood. Sexual morph: Ascomata 430–600 µm (x
̄ = 500 µm, n = 10)
diam., perithecial, solitary, superficial, turbinate, obpyriform to subglobose, or deep collapsing as
cupuliform, 3–8-loculate, 130–200 µm (x
̄ = 160 µm, n = 10) diam., dark brown to black,
membranaceous, tuberculate, glabrous, ostioles inconspicuous, periphysate. Peridium 20–45 µm
wide, membranaceous, comprising three layers, outer layer composed of brown to pale brown cells
of textura globulosa; middle layer composed of pale brown to hyaline cells of textura angularis;
inner layer composed of hyaline cells of textura prismatica. Paraphyses absent. Asci (20–)25–30(–
35) × (2.5–)3–5(–7) µm (x
̄ = 28 × 4 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate,
apex rounded, apical ring absent, evanescent. Ascospores (2.5–)3–3.5(–4.5) × (1–)1.5–2.5 µm (x
̄ =
3.4 × 1.8 µm, n = 50), uni- or bi-seriate, hyaline, ellipsoidal to oval, 0–1-septate, smooth-walled,
guttulate. Asexual morph: appeared in the same locule with asci or in conidiomata. Conidiomata
130–200 µm (x
̄ = 160 µm, n = 10) diam., superficial, turbinate, obpyriform to subglobose, or deep
collapsing as cupuliform, 3–8-loculate, dark brown to black, membranaceous, tuberculate.
Peridium 28–45 µm wide, membranaceous, comprising three layers, outer layer composed of
brown to pale brown cells of textura globulosa; middle layer composed of pale brown to hyaline
cells of textura angularis; inner layer composed of hyaline cells of textura prismatica, towards
inner conidiogenous region. Conidiophores 5–8(–12) × 0.5–1.5 µm (x
̄ = 7.5 × 1 µm, n = 20), erect,
hyaline, cylindrical to subcylindrical, unbranched. Conidia 1–2.5 × 0.7–1.5 µm (x
̄ = 2 × 1 µm, n =
50), terminal, oval to globose, aseptate, hyaline, smooth-walled, guttulate.
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Material examined – Pakistan, Lahore, Changa Manga, on branches of Cadaba farinosa, 24
March 1954, S. Ahmad (BPI-1110420, holotype); ibid. (S-F8085, Isotype).
Known hosts and distribution – on wood of Cadaba farinosa (Capparaceae) in Pakistan (type
locality) (Petrak 1959).
Notes – Petrak (1959) collected Pseudonectriella ahmadii on the wood of Cadaba in Pakistan
and found that its sexual morph is similar to Nectria cinnabarina. In this study, we re-examined the
type material and found both mature asci and conidiophores producing conidia in the same locule
(Fig. 36 u).
Pseudorhynchia Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1206 (1909)
Saprobic leaves or stems. Sexual morph: Ascomata perithecial, scattered or gregarious, semi-
immersed to superficial, globose to subglobose, brown or black, membranaceous, tuberculate,
inconspicuous ostioles surrounded by long setae. Setae composed of filiform, brown, septate,
tapering hyphae, with metallic iridescence. Peridium composed of membranaceous, brown to
hyaline cells of textura epidermoidea to textura angularis to textura prismatica. Paraphyses absent.
Asci 8-spored, unitunicate, clavate, pedicellate, apex rounded, apical ring indistinct, evanescent.
Ascospores bi-seriate, hyaline, fusiform, straight or slightly curved, 0–1-septate, faintly striate,
guttulate. Asexual morph: Undetermined (adapted from Penzig & Saccardo 1897, Dulymamode et
al. 2001).
Notes – Pseudorhynchia was established by von Höhnel (1909a) based on Ceratostomella
polyrrhyncha. The second species, P. mauritiana, was collected from leaves of Cordemoya
integrifolia in Mauritius. They are characterized by ascomata with ostioles surrounded by long
setae and fusiform ascospores (Penzig & Saccardo 1897, Dulymamode et al. 2001). Lumbsch &
Huhndorf (2007) placed this genus in Niessliaceae, but no sequence data is available.
Pseudorhynchia polyrrhyncha (Penz. & Sacc.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl.,
Abt. 1 118: 1206 (1909) Fig. 37
Basionym – Ceratostomella polyrrhyncha Penz. & Sacc., Malpighia 11(9-10): 408 (1897)
Facesoffungi number: FoF 10077
Saprobic leaves or stems. Sexual morph: Ascomata 110–170 µm (x
̄ = 140 µm, n = 10),
perithecial, scattered or gregarious, semi-immersed to superficial, globose to subglobose, brown or
black, membranaceous, tuberculate, inconspicuous ostioles surrounded by long setae. Setae 420–
500 × 25–55 µm, composed of filiform, brown, septate, tapering hyphae, with metallic iridescence.
Peridium 15–45 µm wide, membranaceous, comprising three layers, brown cells of textura
epidermoidea on surface view; middle layer composed of brown to olivaceous brown cells of
textura angularis; inner layer composed of hyaline cells of textura prismatica. Paraphyses absent.
Asci (50–)65–80(–90) × (7–)9–12(–15) µm (x
̄ = 70 × 10 µm, n = 20), 8-spored, unitunicate, clavate,
pedicellate, apex rounded, apical ring indistinct, evanescent. Ascospores (15–)17–20(–25) × (2.5–
)3–5(–5.5) µm (x
̄ = 18 × 4 µm, n = 50), bi-seriate, hyaline, fusiform, straight or slightly curved, 0–
1-septate, faintly striate, guttulate. Asexual morph: Undetermined.
Material examined – Venezuela, Edo. Miranda, Guatopo National Park, alt. 500–600 m,
10º03’N, 66º26’W, Trail between Agua Blanca and La Cruceta, on dead leaves of Heliconia sp.,
27–30 November 1990, G.J. Samuels, B. Hein and S.M. Huhndorf (BPI-802960).
Known hosts and distribution – on decayed leaves or stems of Elettaria in Indonesia (type
locality) and Venezuela (Penzig & Saccardo 1897).
Notes – The type material is unable to obtain. Therefore, we re-examined the authentic
material collected by Samuels, Hein and Huhndorf in Venezuela.
Rosasphaeria Jaklitsch & Voglmayr, Fungal Diversity 52(1): 93 (2012)
Saprobic on twigs of Rosa. Sexual morph: Ascomata perithecial, solitary or scattered,
immersed, pyriform to subglobose, black, membranaceous, glabrous, ostiolate, periphysate.
Peridium composed of membranaceous, brown to hyaline cells of textura angularis to textura
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prismatica. Paraphyses absent. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate or
sessile, apex rounded, apical ring indistinct, evanescent. Ascospores bi-seriate, hyaline, cylindrical
to fusiform, straight or slightly curved, 1-septate, smooth-walled, guttulate. Asexual morph:
Hyphomycetous. Mycelium white to orange, superficial, effused. Conidiophores mononematous,
macronematous, hyaline, unbranched, cylindrical to oblong, aseptate. Conidia terminal, cylindrical,
usually curved, 1-septate, hyaline, smooth-walled, guttulate (adapted from Jaklitsch & Voglmayr
2012).
Figure 36 – Pseudonectriella ahmadii: a–c, e, q–r, s, w (BPI-1110420, holotype); d, f–p, t–v (S-
F8085, Isotype). a Material. b–d Ascomata on the host. e–f Ascomata cross section. g Peridium of
966
ascomata. h–n Asci. o–r Ascospores. s Peridium of conidiomata. t Conidiophores and conidia.
u Mature asci and conidiophores in the same locule. v Wall cells of ascomata towards inner
conidiogenous region (enlarged view of the conidiophores part of u). w Conidiophores and conidia
in conidiomata. Notes: u–w stained in cotton blue reagent. Scale bars: b–d = 200 µm, e–f = 100
µm, g, u = 50 µm, h–k, s = 20 µm, l–n, t, v–w = 10 µm, o–r = 2 µm.
Figure 37 – Pseudorhynchia polyrrhyncha: (BPI-802960). a Material. b–d Ascomata on the host.
e Squashed ascoma. f Seta. g–h Ostiole. i Ascomata cross section. j Peridium. k–l, p Asci.
m–n Ascospores. o Apical ascus. Scale bars: b, e = 500 µm, c–d, f, i = 200 µm, g–h, j, p = 50 µm,
k–l = 20 µm, m–o = 10 µm.
967
Notes – The monotypic Rosasphaeria was established by Jaklitsch & Voglmayr (2012) based
on Stigmatea moravica. This genus was placed in Niessliaceae based on phylogenetic result
(Jaklitsch & Voglmayr 2012). In this study, Rosasphaeria is sister to Neoeucasphaeria (Fig. 1),
differing from the latter that has ellipsoidal, aseptate and hyaline conidia (Crous et al. 2018).
Rosasphaeria moravica (Petr.) Jaklitsch & Voglmayr, Fungal Diversity 52(1): 93 (2012) Fig. 38
Basionym – Stigmatea moravica Petr., Annls mycol. 12(5): 473 (1914)
Facesoffungi number: FoF 10078
Saprobic on twigs of Rosa. Sexual morph: Ascomata 110–220 µm (x
̄ = 160 µm, n = 10)
diam., perithecial, solitary or scattered, immersed, pyriform to subglobose, black, membranaceous,
glabrous, ostiolate, periphysate. Peridium 10–30 µm wide, composed of membranaceous, brown to
hyaline cells of textura angularis to textura prismatica. Paraphyses absent. Asci (50–)67–82(–90) ×
(5–)7–8(–12) µm (x
̄ = 75 × 7.5 µm, n = 20), 8-spored, unitunicate, cylindrical to clavate, pedicellate
or sessile, apex rounded, apical ring indistinct, evanescent. Ascospores (15–)20–25(–30) × (2–)2.5–
4.5(–5) µm (x
̄ = 22 × 3.5 µm, n = 50), bi-seriate, hyaline, cylindrical to fusiform, straight or slightly
curved, 1-septate, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Mycelium white to
orange, superficial, effused. Conidiophores mononematous, macronematous, hyaline, unbranched,
cylindrical to oblong, aseptate. Conidia terminal, cylindrical, usually curved, 1-septate, hyaline,
smooth-walled, guttulate (adapted from Jaklitsch & Voglmayr 2012).
Material examined – Czech Republic, Olomoucky kraj, M. Weisskirchen, Hrabuvka, on dead
twigs of Rosa canina, 18 April 1912, F. Petrak (S-F86647, holotype).
Known hosts and distribution – on decayed twigs of Rosa canina (Rosaceae) in Austria,
Czech Republic (type locality) and Sweden (Jaklitsch & Voglmayr 2012).
Notes – Jaklitsch & Voglmayr (2012) introduced Rosasphaeria moravica in Niessliaceae.
This species is sister to Neoeucasphaeria eucalypti and nests in Niessliaceae (Jaklitsch &
Voglmayr 2012, this study, Fig. 1).
Taiwanascus Sivan. & H.S. Chang, Mycol. Res. 101(2): 176 (1997)
Saprobic on wood. Sexual morph: Ascomata cleistothecial, solitary, superficial, globose,
brown, membranaceous, tuberculate, with setae. Setae cylindrical, aseptate, brown, straight,
smooth-walled, with branched ends. Peridium composed of membranaceous, olivaceous brown
cells of textura angularis. Paraphyses absent. Asci 4-spored, unitunicate, cylindrical, pedicellate,
apex rounded, apical ring absent, evanescent. Ascospores bi-seriate to overlapping, hyaline,
fusiform or ellipsoidal, straight or slightly curved, aseptate or 1-septate, smooth-walled, guttulate.
Asexual morph: Undetermined (adapted from Sivanesan & Chang 1997, Rajeshkumar & Rossman
2013).
Notes – Taiwanascus was introduced by Sivanesan & Chang (1997) for T. tetrasporus.
Rajeshkumar & Rossman (2013) subsequently introduced second species, T. samuelsii, from India.
These species have cleistothecial ascomata with special branched setae, but the former has fusiform
ascospores and the latter has ellipsoidal (Sivanesan & Chang 1997, Rajeshkumar & Rossman
2013). There is no sequence data available for this genus.
Taiwanascus tetrasporus Sivan. & H.S. Chang, Mycol. Res. 101(2): 176 (1997) Fig. 39
Facesoffungi number: FoF 10079
Saprobic on wood. Sexual morph: Ascomata 94–250 µm (x
̄ = 120 µm, n = 10) diam.,
cleistothecial, solitary, superficial, globose, brown, membranaceous, tuberculate, with setae. Setae
16–32 × 3–7 µm (x
̄ = 25 × 5 µm, n = 20), cylindrical, aseptate, brown, straight, smooth-walled,
with 2–8-branched ends, each vertex has 6–8 tiny denticles. Peridium 5–10 µm wide, composed of
membranaceous, olivaceous brown cells of textura angularis. Paraphyses absent. Asci (15–)20–
25(–28) × 4–6 µm (x
̄ = 23 × 5 µm, n = 20) at ascospore-bearing part, 4-spored, unitunicate,
cylindrical, pedicellate, apex rounded, apical ring absent, evanescent. Ascospores (14–)18–22 ×
(0.6–)1–1.6 µm (x
̄ = 20 × 1.2 µm, n = 50), overlapping, hyaline, fusiform, straight or slightly
968
curved, aseptate, rarely 1-septate, smooth-walled, guttulate. Asexual morph: Undetermined
(adapted from Sivanesan & Chang 1997).
Material examined – China, Taiwan, Taipei, Wulai, on dead wood, 18 January 1994, H.S.
Chang (IMI-364835, holotype).
Figure 38 – Rosasphaeria moravica: (S-F86647, holotype). a Material. b Ascomata on the host.
c Immersed ascomata. d–e Ascomata cross section. f Peridium. g–h Asci (h enlarged apical ring).
i–n Ascospores. Scale bars: d–e = 100 µm, f–n = 20 µm.
969
Figure 39 – Taiwanascus tetrasporus: (IMI-364835, holotype). a Material. b–c Ascomata on the
host. d Squashed ascoma. e Seta. f Surface view of ascoma. g–h Asci. i Ascospores. Scale bars: b =
1 mm, c = 500 µm, d = 50 µm, f–h = 20 µm, e. i = 10 µm.
Known hosts and distribution – on decayed wood in China (type locality) (Sivanesan &
Chang 1997).
Notes – Sivanesan & Chang (1997) introduced Taiwanascaceae based on Taiwanascus
tetrasporus and Samuels & Barr (1997) treated this family as a synonym of Niessliaceae.
Therefore, T. tetrasporus was placed in Niessliaceae, but sequence data is not available.
Trichosphaerella E. Bommer, M. Rousseau & Sacc., in Bommer & Rousseau, Bull. Soc. R. Bot.
Belg. 29(no. 1): 261 (1890)
Saprobic or parasitic on wood or stems, sometimes fungicolous. Sexual morph: Ascomata
perithecial, scattered or gregarious, superficial, sometimes sitting on a subiculum, globose to
subglobose, brown, membranaceous, glabrous or with brown setae, ostiolate, periphysate. Peridium
composed of membranaceous, brown to hyaline cells of textura angularis to textura prismatica.
Paraphyses absent. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate or sessile, apex
rounded, apical ring indistinct or absent, evanescent. Ascospores uni- or bi-seriate or overlapping,
hyaline, ellipsoidal to fusiform, straight or slightly curved, usually 1-septate, constricted at the
septum to form ascoconidia, guttulate. Ascoconidia globose to subglobose, hyaline, smooth-walled
970
or spinulose or verrucose, sometimes becoming red in Melzer’s reagent. Asexual morph:
Hyphomycetous. Conidiophores mononematous, macronematous, hyaline, branched, straight or
curved. Conidia globose to ellipsoidal, usually aseptate, hyaline, smooth-walled or verrucose
(adapted from Müller & Samuels 1982, Samuels 1983, Samuels & Barr 1997).
Notes – Trichosphaerella was introduced by Bommer & Rousseau (1890) for T. decipiens.
von Höhnel (1903) subsequently introduced Bresadolella with B. aurea. However, Samuels & Barr
(1997) treated B. aurea as a synonym of T. decipiens based on similar morphology and associated
with acremonium-like fungi. Lombard et al. (2015) sequenced the T. ceratophora which is sister to
Hyaloseta nnolinae in Niessliaceae. In this study, T. ceratophora is sister to Valetoniellopsis,
differing from the latter that has glabrous ascomata with ellipsoidal ascospores. However, there is
no molecular data for the type T. decipiens.
Trichosphaerella decipiens Sacc., E. Bommer & M. Rousseau, in Bommer & Rousseau, Bull. Soc.
R. Bot. Belg. 29(no. 1): 261 (1890) Fig. 40
Facesoffungi number: FoF 10080
Saprobic or parasitic on wood or stems, sometimes fungicolous. Sexual morph: Ascomata
perithecial, scattered or gregarious, superficial, sitting on a subiculum with hyaline, coiled,
verrucose, septate hyphae, globose to subglobose, brown, membranaceous, with brown, cylindrical,
tapering setae, 74–120 × 9–16 µm, ostiolate, periphysate. Peridium composed of membranaceous,
brown to hyaline cells of textura angularis to textura prismatica. Paraphyses absent. Asci 24–28(–
32) × 3–5 µm (x
̄ = 27 × 4.5 µm, n = 20), 8-spored, unitunicate, cylindrical to clavate, pedicellate or
sessile, apex rounded, apical ring indistinct or absent, evanescent. Ascospores bi-seriate, hyaline,
ellipsoidal to fusiform, straight or slightly curved, 1-septate, constricted at the septum to form
ascoconidia, guttulate. Ascoconidia globose to subglobose, hyaline, verrucose. Asexual morph:
Hyphomycetous. Conidiophores mononematous, macronematous, hyaline, branched, straight.
Conidia terminal, oval to ellipsoidal, aseptate, hyaline, smooth-walled (adapted from Bommer &
Rousseau 1890, Höhnel 1903, Samuels & Barr 1997).
Material examined – Austria, Niederösterreich, Pressbaum, on decayed wood of Fagus,
September 1903, v. Höhnel (S-F10868; S-F10869, slide of isotype of Bresadolella aurea).
Known hosts and distribution – on wood or basidiomycete in Austria, Belgium (type
locality), Canada, Estonia, Guyana and Luxembourg (von Höhnel 1903, Samuels & Barr 1997).
Notes – Trichosphaerella decipiens was collected on decorticated brahces of Fagus sylvatica
in Belgium (Bommer & Rousseau 1890). Afterwards, von Höhnel (1903) introduced Bresadolella
for B. aurea collected on decayed wood of Fagus in Austria. Samuels & Barr (1997) accepted B.
aurea as a synonym of T. decipiens based on similar characteristics and obtained its culture from a
material made in Guiana. However, there is no sequence data available for T. decipiens.
Valetoniella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1499 (1909)
Saprobic on wood. Sexual morph: Ascomata perithecial, solitary or scattered, superficial on
decorticated wood, sometimes sitting on a subiculum, globose to subglobose, translucent and pale
brown when fresh, brown or black when dry, membranaceous, tuberculate, ostiolate, periphysate,
collapsing when dry. Setae cylindrical, brown, with several conical, brown to pale brown
ornamentals at apex. Peridium composed of membranaceous, yellow or pale brown to hyaline cells
of textura angularis to textura prismatica. Paraphyses absent. Asci 8-spored, unitunicate, clavate,
pedicellate, apex rounded, apical ring absent, evanescent. Ascospores bi-seriate or overlapping,
hyaline, usually becoming red in Melzer’s reagent, ellipsoidal or fusiform, 1-septate, smooth-
walled, faintly striate, guttulate. Asexual morph: Undetermined (adapted from von Höhnel 1909b,
Samuels & Barr 1997).
Notes – Valetoniella was established by von Höhnel (1909b) for V. crucipila, which was found on
decorticated wood and accompanied Nectria species. Afterwards, Samuels & Barr (1997)
introduced V. claviornata and V. pauciornata based on their ascomata that have obvious setae with
several conical ornamentals at apex, but no molecular data is available for the Valetoniella.
971
Figure 40 – Trichosphaerella decipiens (= Bresadolella aurea): b–c, e–f (S-F10868, isotype);
a, d (S-F10869, isotype); g–i (redrawn from Samuels & Barr 1997). a–b Squashed ascoma. c Seta.
d–g Asci (g ascoconidia in asci). h Hyphae of subiculum. i Conidiophores and conidia. – Scale
bars: a = 200 µm, b = 100 µm, c = 50 µm, d–e = 20 µm, f–i = 10 µm.
Valetoniella crucipila Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1500 (1909)
Fig. 41
Facesoffungi number: FoF 10081
Saprobic on wood. Sexual morph: Ascomata 160–290 µm (x
̄ = 240 µm, n = 10), perithecial,
solitary or scattered, superficial on decorticated wood, sometimes sitting on a subiculum, globose to
subglobose, translucent and pale brown when fresh, brown or black when dry, membranaceous,
tuberculate, ostiolate, periphysate, collapsing when dry. Setae 35–45 × 5–8 µm (x
̄ = 40 × 6 µm, n =
20), cylindrical, brown, with 2–6 conical, brown to pale brown ornamentals at apex. Peridium 25–
35 µm wide, membranaceous, comprising two layers, outer layer composed of yellow or pale
brown cells of textura angularis; inner layer composed of hyaline cells of textura prismatica.
Paraphyses absent. Asci (35–)42–46(–60) × (7–)8–10 µm (x
̄ = 45 × 9 µm, n = 20), 8-spored,
unitunicate, clavate, pedicellate, apex rounded, apical ring absent, evanescent. Ascospores (7.5–)8–
10(–12) × (3–)3.5–4.5(–5.2) µm (x
̄ = 9 × 4 µm, n = 50), bi-seriate or overlapping, hyaline, usually
972
becoming red in Melzer’s reagent, ellipsoidal or fusiform, 1-septate, smooth-walled, faintly striate,
guttulate. Asexual morph: Undetermined (adapted from von Höhnel 1909b, Samuels & Barr 1997).
Figure 41 – Valetoniella crucipila: a–b, d, fj (PDD-41743); c, e, k (PDD-50069). a Material.
b–c Ascomata on the host. d Ascoma cross section. e Squashed ascoma. f Peridium. g Setae.
h–i Asci. j–k Ascospores. Notes: d. f. j from Permanent slide; h–i, k stained in Melzer’s reagent.
Scale bars: b = 500 µm, c = 200 µm, d = 100 µm, e, k = 50 µm, f–i = 20 µm, j = 10 µm.
Material examined – New Zealand, Auckland, Titirangi beach reserve, on ascomata of
Nectria haematococca and N. mammoidea, 18 September 1980, G.J. Samuels and P.R. Johnston
(PDD-41743); New Zealand, Southland, Catlins state Forest Park, lake Wilkie, on ascomata of
Nectria westlandica, 18 April 1985, G.J. Samuels, P.K. Buchanan and L.M. Kohn (PDD-50069).
Known hosts and distribution – on decayed wood and usually accompanied Nectria species in
Guyana, Indonesia (type locality), New Zealand, Puerto Rico and Venezuela (von Höhnel 1909b,
Samuels & Barr 1997).
Notes – Valetoniella crucipila was first collected on decorticated wood of Albizia moluccana
and accompanied Nectria species (von Höhnel 1909b). Subsequently, Samuels (1983) redescribed
this species from New Zealand. Then, Samuels & Barr (1997) noted that it has been recollected in
Guyana, Puerto Rico and Venezuela. This species is characterized by ascomata with special setae,
clavate asci and ellipsoidal ascospores that turn red in Melzer’s reagent (von Höhnel 1909b,
973
Samuels 1983, Samuels & Barr 1997). In this study, the type material is unable to obtain, thus, we
re-examined the authentic materials collected from New Zealand.
Valetoniellopsis Samuels & M.E. Barr, Can. J. Bot. 75(12): 2175 (1998) [1997]
Saprobic on leaves. Sexual morph: Ascomata perithecial, solitary or scattered, superficial,
globose to subglobose, or collapsing as cupuliform, reddish brown or brown, multiloculated,
membranaceous, tuberculate, glabrous or with brown, septate, lax setae at apex, ostiolate,
periphysate. Peridium composed of membranaceous, brown to hyaline cells of textura angularis to
textura prismatica. Paraphyses absent. Asci 8-spored, unitunicate, clavate, pedicellate or sessile,
apex rounded or blunt, apical ring absent, evanescent. Ascospores uni- or bi-seriate, hyaline,
cylindrical to oblong, straight or slightly curved, 1-septate, smooth-walled, guttulate. Asexual
morph: Hyphomycetous. Conidiophores mononematous, macronematous, hyaline, unbranched,
straight. Conidia terminal, ellipsoidal, aseptate, hyaline, smooth-walled, guttulate (adapted from
Samuels & Barr 1997).
Notes – The monotypic Valetoniellopsis is typified by V. laxa and placed in Niessliaceae
(Samuels & Barr 1997). It is similar to Circinoniesslia with setose ascomata and acremonium-like
asexual morph (Samuels & Barr 1997). Subsequently, the type V. laxa (CBS:191.97) was
sequenced and nested in Niessliaceae (Tang et al. 2007, Vu et al. 2019). In this study,
Valetoniellopsis is sister to Trichosphaerella in Niessliaceae (Fig. 1).
Valetoniellopsis laxa Samuels & M.E. Barr, Can. J. Bot. 75(12): 2175 (1998) [1997] Fig. 42
Facesoffungi number: FoF 10082
Saprobic on leaves. Sexual morph: Ascomata (50–)100–500(–700) µm (x
̄ = 370 µm, n = 10),
perithecial, solitary or scattered, superficial, globose to subglobose, or collapsing as cupuliform,
reddish brown or brown to black, multiloculated, membranaceous, tuberculate, glabrous or with
brown, septate, lax setae at apex, ostiolate, periphysate. Peridium 18–85 µm wide, membranaceous,
comprised of two layers, outer layer composed of brown cells of textura angularis; inner layer
composed of hyaline cells of textura prismatica. Paraphyses absent. Asci (18–)24–28(–32) × (5–
)6–7.5(–8.5) µm (x
̄ = 25 × 6.5 µm, n = 20), 8-spored, unitunicate, clavate, pedicellate or sessile,
apex rounded or blunt, apical ring absent, evanescent. Ascospores (5–)5.5–6.5(–8) × (1.5–)2–2.5
µm (x
̄ = 6 × 2.2 µm, n = 50), uni- or bi-seriate, hyaline, cylindrical to oblong, straight or slightly
curved, 0–1-septate, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Conidiophores
mononematous, macronematous, hyaline, unbranched, straight. Conidia terminal, ellipsoidal,
aseptate, hyaline, smooth-walled, guttulate (adapted from Samuels & Barr 1997).
Material examined – USA, Louisiana, Iberia Parish, Avery Island, Jungle Garden, on dead
leaves of palm, 16 August 1996, G.J. Samuels, M. Blackwell, M. Camara (BPI-748357, holotype).
Known hosts and distribution – on dead rachis of palm in the USA (type locality) (Samuels &
Barr 1997).
Notes – Valetoniellopsis laxa was introduced by Samuels & Barr (1997) from the USA. In
this study, this species is sister to Trichosphaerella ceratophora (Fig. 1), but can be differentiated
from V. laxa that has cylindrical to oblong ascospores without ascoconidia (Samuels & Barr 1997).
Other genera in Hypocreomycetidae
Glomerellales genera incertae sedis
Ascocodinaea Samuels, Cand. & Magni, Mycologia 89(1): 156 (1997)
Fungicolous. Sexual morph: Ascomata perithecial, solitary or scattered, superficial, globose
to subglobose, black, membranaceous, tuberculate, with cylindrical, unbranched, dark brown,
tapering setae, collapsing when dry, ostiolate, periphysate. Peridium composed of brown to hyaline
cells of textura epidermoidea to textura prismatica. Paraphyses filiform, septate, branched. Asci 8-
spored, unitunicate, cylindrical, pedicellate, apex blunt, apical ring indistinct, evanescent.
Ascospores uni-seriate, ellipsoidal to fusiform, straight or slightly curved, hyaline, aseptate,
becoming brown median cells and hyaline end cells, usually 3-septate, constricted at the middle
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septum, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Conidiophores
mononematous, macronematous, hyaline, unbranched, brown to dark brown, swollen at base.
Conidiogenous cells monophialidic, enteroblastic, thicken. Conidia terminal, oval to ellipsoidal,
straight or slightly curved, 0–1-septate, hyaline, smooth-walled, guttulate (adapted from Samuels et
al. 1997).
Notes – Two Ascocodinaea species, A. polyporicola and A. stereicola (type), were collected
from basidiomata of Polyporus sp., Stereum sp. and Trichaptum sp. in the USA (Samuels et al.
1997). This genus was initially placed in Hypocreomycetidae (Lumbsch & Huhndorf 2007).
However, Huhndorf et al. (2004a) analyzed LSU sequence data of A. stereicola and found that this
species is sister to Colletotrichum gloeosporioides (= Glomerella cingulata). Subsequently,
Ascocodinaea was transferred to Glomerellales genera incertae sedis (Hyde et al. 2020e,
Wijayawardene et al. 2020). In this study, Ascocodinaea is basal to Plectosphaerellaceae in
Glomerellales as an independent clade (Fig. 1).
Figure 42 – Valetoniellopsis laxa: a–j (BPI-748357, holotype); k (redrawn from Samuels & Barr
1997). a Material. b Ascomata on the host. c Ascoma. d Ascomata cross section. e A locule.
f Peridium. g–i Asci. j Ascospores. k Conidiophores and conidia. Scale bars: c–d = 100 µm, e = 50
µm, f, k = 20 µm, g–i = 10 µm, j = 5 µm.
Ascocodinaea stereicola Samuels, Cand. & Magni, Mycologia 89(1): 156 (1997) Fig. 43
Facesoffungi number: FoF 10083
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Fungicolous. Sexual morph: Ascomata 150–180 × 120–180 µm (x
̄ = 160 × 155 µm, n = 10),
perithecial, solitary or scattered, superficial, globose to subglobose, black, membranaceous,
tuberculate, with cylindrical, unbranched, dark brown, tapering setae, 50–90 × 3–8 µm, collapsing
when dry, ostiolate, periphysate. Peridium 15–30 µm wide, comprising two layers, outer layer
composed of brown cells of textura epidermoidea; inner layer composed of hyaline cells of textura
prismatica. Paraphyses 1.5–3 µm wide, filiform, septate, branched. Asci (90–)95–110(–120) × (5–
)6–7(–9) µm (x
̄ = 100 × 6.5 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex blunt,
apical ring indistinct, evanescent. Ascospores (9–)10–13(–15) × (4–)5–7(–8) µm (x
̄ = 11 × 5.5 µm,
n = 50), uni-seriate, ellipsoidal to broadly fusiform, straight or slightly curved, hyaline, aseptate,
becoming brown median cells and hyaline end cells, usually 3-septate, constricted at the middle
septum, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Conidiophores 2–8 µm wide,
mononematous, macronematous, hyaline, unbranched, brown to dark brown, swollen at base.
Conidiogenous cells monophialidic, enteroblastic, thicken. Conidia (6–)7.5–9(–12) × (2–)3–4(–6)
µm (x
̄ = 8.5 × 3.5 µm, n = 50), terminal and lateral, oval to ellipsoidal, straight or slightly curved,
0–1-septate, hyaline, smooth-walled, guttulate (adapted from Samuels et al. 1997).
Material examined – USA, Kentucky. Laurel Co., Daniel Boone National Forest, Laurel
River Lake Recreation Area, Cane Creek Wildlife Refuge, alt. 350 m, on Stereum sp., 27
September 1995, G.J. Samuels (BPI-749458, holotype); ibid. (BPI-749459, isotype).
Known hosts and distribution – on Stereum in the USA (type locality) (Samuels et al. 1997).
Notes – The fungicolous Ascocodinaea stereicola usually found on Stereum (Samuels et al.
1997). It is closely related to members of Glomerellales based on LSU sequence data (Huhndorf et
al. 2004b, Hyde et al. 2020e, this study, Fig. 1).
Torpedosporales E.B.G. Jones, Abdel-Wahab & K.L. Pang
Etheirophoraceae Rungjind., Somrith. & Suetrong
Etheirophora Kohlm. & Volkm.-Kohlm., Mycol. Res. 92(4): 414 (1989)
Saprobic on wood in marine habitats. Sexual morph: Ascomata perithecial, solitary,
immersed, pyriform, pale brown, coriaceous, papillate. Papilla conical, brown, ostiolate,
periphysate, sometimes erumpent through the wood surface and surrounded by irregular, brown
clypeus. Peridium composed of pale brown cells of textura angularis. Paraphyses filiform, septate,
branched. Asci 8-spored, unitunicate, cylindrical to clavate, pedicellate, apex rounded or blunt,
apical ring absent, evanescent. Ascospores bi-seriate, hyaline, ellipsoidal, straight or slightly
curved, 1-septate, smooth-walled, with 5–8 filiform appendages at one or each end, guttulate.
Asexual morph: Undetermined (adapted from Kohlmeyer & Volkmann-Kohlmeyer 1989).
Notes – Kohlmeyer & Volkmann-Kohlmeyer (1989) found Etheirophora bijubata (type), E.
blepharospora and E. unijubata on rotten wood collected in Hawaii. They have ellipsoidal
ascospores with several filiform terminal appendages at one or each end (Kohlmeyer & Volkmann-
Kohlmeyer 1989). However, Schoch et al. (2007) sequenced E. blepharospora and E. unijubata.
Subsequently, Jones et al. (2014) established Etheirophoraceae, which accommodated
Etheirophora and Swampomyces. Etheirophora and Swampomyces that form a clade as
Etheirophoraceae (Jones et al. 2014, this study, 100%ML/1.00BY, Fig. 1). Swampomyces is
different from Etheirophora in that its ellipsoidal ascospores lacks appendages (Jones et al. 2014).
Etheirophora bijubata Kohlm. & Volkm.-Kohlm., Mycol. Res. 92(4): 414 (1989) Fig. 44a–n
Facesoffungi number: FoF 10084
Saprobic on wood in marine habitats. Sexual morph: Ascomata 140–260 × 270–475 µm (x
̄ =
200 × 340 µm, n = 10), perithecial, solitary, immersed, pyriform, pale brown, coriaceous, papillate.
Papilla conical, brown, ostiolate, periphysate, sometimes erumpent through the wood surface and
surrounded by irregular, brown clypeus. Peridium 5–18 µm wide, comprising two layers, outer
layer composed of brown to pale brown cells of textura angularis; inner layer composed of hyaline
cells of textura prismatica. Paraphyses 0.6–1.6 µm wide, filiform, septate, branched. Asci (60–)65–
75(–90) × (9–)10–12(–15) µm (x
̄ = 74 × 11.5 µm, n = 20), 8-spored, unitunicate, cylindrical to
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clavate, pedicellate, apex rounded or blunt, apical ring absent, evanescent. Ascospores (14–)16–
18(–21) × (5–)5.5–7(–8) µm (x
̄ = 17 × 6 µm, n = 50), bi-seriate, hyaline, ellipsoidal, straight or
slightly curved, 1-septate, smooth-walled, with 5–8 filiform, appendages at each end, guttulate.
Asexual morph: Undetermined (adapted from Kohlmeyer & Volkmann-Kohlmeyer 1989).
Figure 43 – Ascocodinaea stereicola: b–n (BPI-749458, holotype); a, o–x (BPI-749459, isotype).
a Material (Stereum sp.). b Ascoma on host. c Ascoma cross section. d Peridium. e Ostiole
surrounded by setae. f Paraphyses. g–k Asci. l–n Ascospores. o–q Colonies on dry culture on
surface view. r–x Conidiophores and conidia. Scale bars: p = 2 mm, q = 500 µm, b = 200 µm, c =
100 µm, e–k = 50 µm, d, r–u = 20 µm, v–w = 10 µm, l–n, x = 5 µm.
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Figure 44 – Etheirophora bijubata: a–n (NY-1315470, microslide of holotype). a Material.
b Immersed ascomata. c–e Ascomata cross section. f Ostiole with periphyses. g–h Asci.
i Paraphyses. j–n Ascospores; Juncigena adarca: o–x (NY-1276214, holotype). o Material.
p Immersed ascoma on surface view. q–s Ascomata cross section. t Evanescent ascus.
u–x Ascospores. Notes: t, v–x stained in Melzer’s reagent. Scale bars: b = 500 µm, c–e, p–s = 100
µm, f–g, i, t = 50 µm, h, j–n, u–x = 20 µm.
Material examined – USA, Hawaii. Haena Beach, Hanalei, Kauai (Pacific Ocean),
22°13'18"N, 159°34'16"W (22.222, -159.571), on intertidal wood which embedded in concrete
978
(incubated in a moist chamber for 9 months), 8 March 1987, J.J. Kohlmeyer, B. Volkmann-
Kohlmeyer (NY-1315470, microslide of holotype).
Known hosts and distribution – on intertidal wood in the USA (type locality) (Kohlmeyer &
Volkmann-Kohlmeyer 1989).
Notes – The marine fungus Etheirophora bijubata was introduced as type (Kohlmeyer &
Volkmann-Kohlmeyer 1989), but no sequence data is available.
Swampomyces Kohlm. & Volkm.-Kohlm., Bot. Mar. 30(3): 198 (1987)
Saprobic on wood in marine habitats. Sexual morph: Ascomata perithecial, solitary,
immersed, pyriform, pale brown, with black papilla on surface view, coriaceous, ostiolate,
periphysate. Peridium composed of pale brown to hyaline cells of textura epidermoidea to textura
angularis to textura prismatica. Pseudoparaphyses filiform, septate, branched, attached at the top
and bottom. Asci 8-spored, unitunicate, cylindrical, pedicellate, apex rounded or blunt, apical ring
absent, evanescent. Ascospores uni-seriate, hyaline, ellipsoidal, hyaline to yellowish, septate,
verrucose, guttulate. Asexual morph: Undetermined (adapted from Kohlmeyer & Volkmann-
Kohlmeyer 1987).
Notes – Swamponyces was established for S. armeniacus which was collected on mangrove
wood in Belize (Kohlmeyer & Volkmann-Kohlmeyer 1987). Subsequently, S. aegyptiacus, S.
clavatisporus and S. triseptatus were introduced from mangrove substrates with immersed
ascomata and ellipsoidal, septate ascospores (Hyde & Nakagiri 1992, Abdel-Wahab et al. 2001).
These species were sequenced by Sakayaroj et al. (2005) and Schoch et al. (2007). Thus,
phylogenetically, Swampomyces aegyptiacus and S. clavatisporus were clarified as Fulvocentrum
species in Juncigenaceae; while, Etheirophora and Swampomyces clustered in Etheirophoraceae
(Jones et al. 2014, this study, Fig. 1).
Swampomyces armeniacus Kohlm. & Volkm.-Kohlm., Bot. Mar. 30(3): 200 (1987) Fig. 45
Facesoffungi number: FoF 05390
Saprobic on wood in marine habitats. Sexual morph: Ascomata 300–400 × 350–600 µm (x
̄ =
345 × 480 µm, n = 10), perithecial, solitary, immersed, pyriform, pale brown, with black papilla on
surface view, coriaceous, ostiolate, periphysate. Peridium 24–35 µm wide, carbonaceous, black
tissues at apex, fused with cells of substrate, venter wall comprising two layers, membranaceous,
outer layer composed of pale brown to hyaline cells of textura epidermoidea to textura angularis;
inner layer composed of hyaline cells of textura prismatica. Pseudoparaphyses 1.2–2.5 µm wide,
filiform, septate, branched, attached at the top and bottom. Asci (100–)125–140(–160) × (15–)17–
18(–20) µm (x
̄ = 135 × 17.5 µm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex
rounded or blunt, apical ring absent, evanescent. Ascospores (11–)14–18(–22) × (6.5–)7.5–9(–10)
µm (x
̄ = 16.5 × 8 µm, n = 50), uni-seriate, hyaline, ellipsoidal, hyaline to yellowish, 1-septate,
verrucose, guttulate. Asexual morph: Undetermined (adapted from Kohlmeyer & Volkmann-
Kohlmeyer 1987).
Material examined – Belize, Man-Of-War Cay (Atlantic Ocean). 16°53'00"N, 88°06'15"W
(16.883, -88.104), on driftwood, 23 November 1984, J.J. Kohlmeyer (NY-1271822, paratype).
Known hosts and distribution – on decayed wood of Rhizophora in Australia and Belize (type
locality) (Kohlmeyer & Volkmann-Kohlmeyer 1987, Read et al. 1995).
Notes – Swampomyces armeniacus was introduced by Kohlmeyer & Volkmann-Kohlmeyer
(1987) from Belize. It is sister to S. triseptatus in Etheirophoraceae (Sakayaroj et al. 2005, Schoch
et al. 2007, Jones et al. 2014, this study, 96%ML/1.00BY, Fig. 1), differing from it in the
possessing of 1-septate ascospores, whereas the latter that has 3-septate ascospores.
Juncigenaceae E.B.G. Jones, Abdel-Wahab & K.L. Pang
Juncigena Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. Mar. 40(4): 291 (1997)
Saprobic on leaves or culms in marine habitats. Sexual morph: Ascomata perithecial, solitary,
immersed, pyriform to subglobose, brown, coriaceous, ostiolate, periphysate. Peridium composed
979
of membranaceous, pale brown to hyaline cells of textura angularis to textura prismatica.
Pseudoparaphyses filiform, septate, unbranched, attached at the top and bottom. Asci 8-spored,
unitunicate, cylindrical, pedicellate, apex blunt, apical ring, blue in methylene blue, evanescent.
Ascospores uni- to bi-seriate, ellipsoidal to fusiform, straight or slightly curved, hyaline, 3-septate,
constricted at the septa, smooth-walled, guttulate. Asexual morph: Hyphomycetous. Hyphae
hyaline to pale brown, septate, branched. Conidia developing directly on hyphae or on non-
specialized short lateral conidiogenous cells, solitary, irregularly helicoid, pale brown to dark
brown, 4–9-septate, smooth-walled, guttulate (adapted from Kohlmeyer et al. 1997).
Figure 45 – Swampomyces armeniacus: (NY-1271822, paratype). a Material. b Ascomata on the
host. c–d Ascomata cross section. e Peridium. f Pseudoparaphyses. g Ostiole with periphyses.
h–i Asci. j–q Ascospores. Scale bars: b = 1 mm, c–d = 200 µm, e = 100 µm, f–i = 50 µm, j–q = 10
µm.
Notes – Juncigena was introduced as a monotypic genus for J. adarca in Magnaporthaceae
(Kohlmeyer et al. 1997). Schoch et al. (2007) sequenced the type and placed this genus in
980
Juncigenaceae (Jones et al. 2014). Later, the second species J. fruticosae was introduced from the
hyphomycetous Moheitospora fruticosae based on sequence data (Abdel-Wahab et al. 2010). In
this study, Juncigena is sister to Khaleijomyces and Marinokkulati (75%ML/0.91BY, Fig. 1).
Juncigena adarca Kohlm., Volkm.-Kohlm. & O.E. Erikss., Bot. Mar. 40(4): 291 (1997)
Fig. 44o–x
Facesoffungi number: FoF 10085
Saprobic on leaves or culms in marine habitats. Sexual morph: Ascomata 220–300 × 150–180
µm (x
̄ = 270 × 160 µm, n = 5), perithecial, solitary, immersed, pyriform to subglobose, brown,
coriaceous, ostiolate, periphysate. Peridium 7–35 µm wide, comprising two layers, outer layer
composed of pale brown cells of textura angularis; inner layer composed of hyaline cells of textura
prismatica. Pseudoparaphyses filiform, septate, unbranched, attached at the top and bottom. Asci 8-
spored, unitunicate, cylindrical, pedicellate, apex blunt, apical ring, blue in methylene blue,
evanescent. Ascospores (23–)26–30(–32) × (5–)6–7 µm (x
̄ = 27 × 56 µm, n = 10), uni- to bi-seriate,
ellipsoidal to fusiform, straight or slightly curved, hyaline, 3-septate, constricted at the septa,
smooth-walled, guttulate. Asexual morph: Hyphomycetous. Hyphae hyaline to pale brown, septate,
branched. Conidia developing directly on hyphae or on non-specialized short lateral conidiogenous
cells, solitary, irregularly helicoid, pale brown to dark brown, 4–9-septate, smooth-walled, guttulate
(adapted from Kohlmeyer et al. 1997).
Material examined – USA, North Carolina, Broad Creek (Atlantic Ocean), 34°43'00"N,
76°55'07"W (34.717, -76.919), on dead culms of Juncus roemerianus, 9 October 1994, J.J.
Kohlmeyer (NY-1276214, holotype).
Known hosts and distribution – on dead leaves and culms of Juncus roemerianus (Juncaceae)
in the USA (type locality) (Kohlmeyer et al. 1997).
Notes – Juncigena adarca was collected on leaves of Juncus at atlantic coast in the USA and
its asexual morph produced on seawater agar medium (Kohlmeyer et al. 1997). Later, it was placed
in Juncigenaceae based on phylogenetic results (Schoch et al. 2007, Jones et al. 2014). In this
study, J. adarca is sister to hyphomycetous J. fruticosae (Abdel-Wahab et al. 2010, this study,
93%ML/0.98BY, Fig. 1).
Sordariomycetes genera incertae sedis
Cryptosphaerella Sacc., Syll. fung. (Abellini) 1: 186 (1882)
Saprobic on wood. Sexual morph: Ascomata perithecial, scattered to gregarious, immersed to
erumpent, coriaceous to membranaceous, brown to black, globose to subglobose, ostiolate,
periphysate. Peridium composed of membranaceous, brown to hyaline cells of textura angularis to
textura prismatica. Paraphyses absent. Asci polysporous, unitunicate, cylindrical, pedicellate, apex
rounded, without apical ring, evanescent. Ascospores overlapping, hyaline, allantoid to cylindrical,
aseptate, usually slightly curved, smooth-walled, guttulate. Asexual morph: Undetermined (adapted
from Saccardo 1882b, Spegazzini 1909, 1922, 1926).
Notes – Cryptosphaerella was introduced similar to Cryptosphaeria (Diatrypaceae) in having
ostiolate ascomata and allantoid, aseptate ascospores (Saccardo 1882a). Later, because of the
polysporous asci, it was considered similar to Coronophora (von Höhnel 1907), but sequence data
is not available for the type C. nitschkei. Mugambi & Huhndorf (2010) sequenced six
‘Cryptosphaerella’ species, C. celata, C. costaricensis, C. cylindriformis, C. elliptica, C. globosa
and C. malindensis, which have ascomata lacking ostioles and with Quellkörper. Thus,
Cryptosphaerella was transferred to Scortechiniaceae based on the phylogenetic analysis of these
six species. Mugambi & Huhndorf (2010) believed that Quellkörper is a key feature of
Scortechiniaceae, while the type of Cryptosphaerella does not have this character (Fig. 25i).
Therefore, these six species are related to Scortechiniaceae, but do not belong to Cryptosphaerella.
We have reassigned them based on morphology and phylogenetic result (see notes for
Neocryptosphaerella and Pseudocryptosphaerella). Here, we transfer Cryptosphaerella to
Sordariomycetes genera incertae sedis pending future work.
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Cryptosphaerella nitschkei (Auersw.) Sacc., Syll. fung. (Abellini) 1: 186 (1882) Fig. 25g–k
Basionym – Valsa nitschkei Auersw., in Nitschke, Pyrenomyc. Germ. 2: 163 (1870)
Facesoffungi number: FoF 10182
Saprobic on wood. Sexual morph: Ascomata perithecial, densely gregarious, immersed to
erumpent, coriaceous to membranaceous, brown to black, globose to subglobose, with periphyses
inside of the ostioles. Peridium composed of membranaceous, brown cells of textura angularis.
Paraphyses absent. Asci polysporous, unitunicate, cylindrical, pedicellate, apex rounded, without
apical ring, evanescent. Ascospores overlapping, hyaline, allantoid, aseptate, slightly curved,
smooth-walled, guttulate. Asexual morph: Undetermined (adapted from Saccardo 1882b).
Material examined – GERMANY, Oederan, on rotten wood, September 1871, H.G. Winter
(IMI-K-M251354); ibid. (NY-02934200).
Known hosts and distribution – on decayed wood of Salix in Germany (type locality)
(Saccardo 1882a).
Notes – The type material is unable to obtain. We re-examined two authentic specimens
collected by Winter from Germany, but fruiting bodies are empty.
Discussion
Taxonomy is the science of classification, including naming, describing, classifying plants,
animals and microorganisms (https://www.cbd.int/gti/taxonomy.shtml, Kirk et al. 2008, Enghoff
2009). Initially, fungi were described and classified based on shared morphological characteristics
(Nannfeldt 1932, Lundqvist 1972). Many species collected and examined earlier have been
described only based on morphological data, and some have drawings. This information is not
enough to allow these species to be easily identified in the current research arena. For example,
Cryptosphaerella, was believe to possess Quellkörper (Mugambi & Huhndorf 2010) but in this
study, we observed that the type species of Cryptosphaerella (C. nitschkei) has ostiolate ascomata
instead of Quellkörper. Mugambi & Huhndorf (2010) redefined the families in Coronophorales
based on morphology and LSU, TEF and RPB2 genes analyses. They proposed that usually,
Bertiaceae has robust ascomata with warted or tuberculate walls; Scortechiniaceae has quellkörper
structure in ascomata (Mugambi & Huhndorf 2010), which is similar to what we have observed in
this study. However, based on the review of the type species, we found that it is difficult to
distinguish other Coronophorales families by the number of ascospores per ascus, ascospore
appendages or ascomatal vestiture. We recommend determining the taxonomic placement of each
family by multi-gene analyses. Podospora anserina has been used as a model filamentous organism
for more than 100 years and it is widely used in various research in biological science
(Scheckhuber & Osiewacz 2008, Ament-Velásquez et al. 2020), but its taxonomy has been
controversial. Wang et al. (2019a) combined P. anserina under Triangularia anserina based on its
type strain, CBS 433.50, which was phylogenetically related to Triangularia. Ament-Velásquez et
al. (2020) believes that the widely used name P. anserina should be retained to avoid confusion in
research. In the past two decades, many novel taxa in Ascomycota have been discovered based on
morphological examination coupled with sequence data (Cai et al. 2005, Wang et al. 2019a, Marin-
Felix et al. 2020, Hyde et al. 2020e). Therefore, it is necessary to supplement the descriptions,
photoplates and molecular information of the species that have been discovered. This will improve
the classification system of fungi and allow us to find some overlooked species or characteristics,
like in Synaptospora. Huhndorf et al. (1999) and Réblová (2002) noted that the ascospores of
Synaptospora fuse into several groups in the ascus when mature; however, Huang et al. (2021) re-
examined the type S. petrakii and found that the ascospores began to produce septa when immature
and split into individual spore cells when mature.
We provide descriptions, photoplates or drawings by reviewing type/authentic material
together with the literature in Coronophorales and Niessliaceae. In this study, several type materials
are not reviewed. This is because 1) the sample information is missing, including the herbarium
details; 2) the samples were lost, destroyed or contaminated; 3) the samples are fragile and/or
cannot be transported and 4) the species has been published recently, with detailed photoplates and
982
descriptions. We also found that some morphological characters of some species were hard to or
could not be observed when re-examined, especially for older type material. For example,
Patouillard (1922) noted that Spinulosphaeria thaxteri has hyaline and evanescent paraphyses, but
we did not observe paraphyses in the holotype material of S. thaxteri collected in 1898. Engler
(1895) recorded that Euacanthe usambarensis has elongated fusiform asci with a long pedicel
about 21 µm. We reviewed its holotype material from 1893 and found that most of the pedicels of
the asci could not be observed; thus, we could not provide a measurement for the pedicel.
Some studies have published DNA based information on known species, e.g Cai et al. (2006)
and Vu et al. (2019). These data provided better insights into the taxonomic placement of some
controversial taxa. For example, the polyphyletic Lasiosphaeriaceae with cylindrical to pod-like
ascospores is divided into six families based on phylogenetic results (Vu et al. 2019, Wang et al.
2019a, Marin-Felix et al. 2020, Huang et al. 2021). However, we have not been able to sequence
the reviewed materials in this study. We expect that the sequence data of all species can eventually
be published. This allows us to determine the classification of taxa more accurately, and makes the
taxa easier to be identified and used in other researches. Likewise, it is necessary to recollect and
identify species based on type material collection information, morphology and molecular data.
These will be meaningful for various biological research, such as evolution, ecology and chemistry.
Acknowledgements
This work was jointly supported by the National Natural Science Foundation of China (No.
31760014) and the Science and Technology Foundation of Guizhou Province (No. [2019]2451-3).
Shi-Ke Huang thanks to Ming Zeng, Drs. S. Boonmee, P. Chomnunti, A.H. Ekanayaka, R. Jeewon,
Ji-Chuan Kang, Xiang-Yu Zeng and Qi Zhao for their invaluable suggestions and help in this study.
Kevin D. Hyde acknowledges the future of specialist fungi in a changing climate: baseline data for
generalist and specialist fungi associated with ants, Rhododendron species and Dracaena species
(grant no: DBG6080013) and Impact of climate change on fungal diversity and biogeography in the
Greater Mekong Subregion (grant no: RDG6130001). Shaun Pennycook is thanked for checking
and correcting the Latin names.
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